Coelurosauria Huene, 1914
Definition- (Ornithomimus velox <- Allosaurus fragilis)
(suggested)
Other definitions- (Passer domesticus <- Allosaurus fragilis)
(Holtz, Molnar and Currie, 2004; modified from Holtz, 1996; modified from Gauthier, 1986)
(Passer domesticus <- Allosaurus fragilis, Sinraptor dongi, Carcharodontosaurus
saharicus) (Xu, You, Du and Han, 2011)
= Coelurosauridae Williston, 1925?
= Coelurosauroidea Williston, 1925? sensu Nopsca, 1928
= Coelurosauria sensu Gauthier, 1986
Definition- (Passer domesticus <- Allosaurus fragilis) (modified)
= Metatheropoda Ji and Ji, 2001
= Coelurosauria sensu
Xu, You, Du and Han, 2011
Definition- (Passer domesticus <- Allosaurus fragilis, Sinraptor
dongi, Carcharodontosaurus saharicus)
Comments- Originally conceived by Huene as a clade including what are now coelophysoids, basal coelurosaurs (e.g. Coelurus, Compsognathus) and Ornithomimus, as opposed to Ceratosaurus,
megalosaurs and carnosaurs which were grouped with sauropodomorphs as
Pachypodosauria. This view evolved so that in e.g. 1923, Huene
now included Ceratosaurus, Elaphrosaurus and tyrannosaurids in the clade, then considered sister to his Carnosauria in Theropoda. By 1932, oviraptorosaurs, Saurornithoides and dromaeosaurids were included by Huene. Romer's (1956) textbook set the consensus that tyrannosaurids and probably Ceratosaurus
were carnosaurs instead, making a division between small coelurosaurs
and large carnosaurs implicit. Ostrom's (1969) recognition that Deinonychus
exhibited a mix of Romer's coelurosaur (20 of 35) and carnosaur (13 of
36) characters began the breakdown of this classic division, stating "I
presently have strong reservations about the validity of these two
categories." This collapse is exemplified by Barsbold's 1977
classification restricting Coelurosauria to coelophysoids and coelurids
sensu lato, raising Ornithomimosauria, Oviraptorosauria and
Deinonychosauria to equivalent rank, and Welles' (1984) review of
Triassic and Jurassic theropods that concluded they "can be grouped
into families ... but that the relationships of the families are not
clear." Thulborn (1984) also rejected the dichotomy, placing Archaeopteryx, tyrannosaurids, troodontids, ornithomimids and Avimimus
closer to birds than allosaurids, but has compsognathids and
dromaeosaurids outside what today would be considered Avetheropoda.
Paul's (1988) influential phylogeny abandoned the term
coelurosaur altogether, placing coelophysoids basal to averostrans as
in the current consensus, Compsognathus and Coelurus just outside Avetheropoda, Ornitholestes and Proceratosaurus
as basal carnosaurs, and suggesting a Protoavia equivalent in content
to modern Maniraptoriformes. Around this time, Gauthier's
unpublished thesis proposed our current usage of Coelurosauria, for
"birds, and all theropods that are closer to birds than to
Carnosauria." This excluded coelophysoids, Ceratosaurus and tyrannosaurids, but did include Coelurus, Ornitholestes, Compsognathus,
ornithomimosaurs, oviraptorosaurs, deinonychosaurs and birds.
Gauthier and Padian (1985) published this concept, but no phylogenetic
definition was provided until Gauthier (1986). Coelurosauria's
content has remained largely stable since, with the addition of
tyrannosaurs and therizinosaurs the consensus as of 1994 and of
alvarezsauroids, first recognized as a clade in that same year.
The inclusion of megaraptorans is still debated, with some authors
placing them in Carnosauria.
Coelurosauridae- Ever since the
erection of Coelurosauria, authors have sometimes listed a family
Coelurosauridae, generally as an alternative for Coelurosauria or
Coeluridae. However, a genus Coelurosaurus has never been validly
named, with published listings being misspellings of Coelurus, "Coelosaurus" or Coelurosauravus.
Thus a Coelurosauridae or Coelurosauroidea could not exist under the
ICZN. The earliest authorship for the family name Coelurosauridae
was listed as "Cope, 1882" by Steel (1970) as a junior synonym of
Coeluridae. However, none of Cope's publications for that year
use the term, the closest being his 'Marsh on the classification of the
Dinosauria' which uses Coeluria. Similarly, Hay (1930:184) lists
Coelurosauridae as being used by Williston (1925), but this cannot be
confirmed without a copy of that textbook. Nopcsa (1928:183)
definitely uses Coelurosauroidea as a suborder equivalent to
Coelurosauria (using Megalosauroidea instead of Carnosauria as well),
which given ICZN rules would also establish other family level varients
for that year if Williston's was unreliable.
Metatheropoda- Metatheropoda was named in a cladogram by Ji and Ji (2001)
as a clade within Coelurosauria containing Compsognathus, Sinosauropteryx
and Maniraptoriformes. Which coelurosaurs were excluded is not specified (though
tyrannosauroids are a likely candidate) and the clade was not defined. The caption
merely listed "down-like protofeathers" as a diagnosis, which suggests
it was named to encompass feathered coelurosaurs. This idea is complicated by
their inclusion of Compsognathus, which they place in a new subclade
Aptilonia. Though Aptilonia is not defined either, the etymology and pairing
with Sinosauropteryx's Eoptilonia suggests it implies a lack of feathers
in Compsognathus, though this is in all probability preservational. As
feathers are probably symplesiomorphic for dinosaurs (Tianyulong, Kulindadromeus), Metatheropoda seems
like an unnecessary clade.
Coelurosauria defined- Xu et al.'s (2011) definition differs from the
standard one by including Sinraptor and Carcharodontosaurus as
external specifiers. Sinraptor's inclusion is superfluous, as an (Allosaurus,
Carcharodontosaurus (Sinraptor, Passer)) topology has never been
advocated. If Carcharodontosaurus is a tyrannosauroid (Paul, 1988; Kurzanov,
1989; Molnar et al., 1990), this redefinition would exclude tyrannosauroids
from Coelurosauria. One thing I
object to is the use of Passer as an internal specifier for Coelurosauria,
as birds were nor originally classified as coelurosaurs in Huene, 1914 or by
anyone until the 1970's at least. Huene included what would today be called
coelophysids, coelurids, compsognathids, Ornitholestes and ornithomimids.
The best internal specifier for Coelurosauria in my opinion is Ornithomimus.
It's always been a coelurosaur, and has always been placed closer to birds than
Allosaurus (unlike Compsognathus, Coelurus or Ornitholestes-
Paul, 1988; Novas, 1992). Thus I would suggest (Ornithomimus velox <-
Allosaurus fragilis, Carcharodontosaurus saharicus) as a definition for
Coelurosauria.
References- Cope, 1882. Marsh on the classification of the Dinosauria. American Naturalist. 16, 253-255.
Huene, 1914. Beitr�ge zur geschichte der Archosaurier. Geologie und Pal�ontologie Abhandlungen. 13(7), 1-56.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of
the Geological Society of America. 34, 449-458.
Williston, 1925. The Osteology of the Reptiles. Harvard University Press. 300 pp.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Hay, 1930. Second bibliography and catalogue of the fossil Vertebrata of North
America. Carnegie Institution of Washington Publication. 390(2), 1074 pp.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
viii + 361 pp.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod
from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural
History. 30, 165 pp.
Steel, 1970. Part 14. Saurischia. Handbuch der Pal�oherpetologie/Encyclopedia
of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp.
Barsbold, 1977. O evolutsiy chishcheich dinosavrov. Trudy - Sovmestnaya
Sovetsko-Mongol'skaya Paleontologicheskaya Ekspeditsiya. 4, 48-56.
Gauthier, 1984. A cladistic analysis of the higher systematic
categories of the Diapsida. PhD thesis. University of California,
Berkeley. 564 pp.
Thulborn, 1984. The avian relationships of Archaeopteryx, and the origin of birds. Zoological Journal of the Linnean Society. 82(1-2), 119-158.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology
and comparisons. Palaeontographica, Abteilung A. 185, 85-180.
Gauthier and Padian, 1985. Phylogenetic, functional, and aerodynamic
analyses of the origin of birds and their flight. In Hecht, Ostrom,
Viohl and Wellnhofer (eds.). The Beginnings of Birds: Proceedings of
the International Archaeopteryx Conference, Eichst�tt 1984. Freunde des Jura-Museums Eichst�tt, Eichst�tt. 185-197.
Gauthier, 1986. Saurischian monophyly and the origin of birds. Memoirs of the Californian Academy of Sciences 8, 1-55.
Holtz, 1996. Phylogenetic taxonomy of the Coelurosauria (Dinosauria:
Theropoda). Journal of Paleontology. 70, 536-538. DOI:
10.1017/S0022336000038506
Ji and Ji, 2001. How can we define a feathered dinosaur as a bird? In
Gauthier and Gall (eds.). New Perspectives on the Origin and Early
Evolution of Birds: Proceedings of the International Symposium in Honor
of John H. Ostrom. 43-46.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Samman, 2007. Assessing craniocervical functional morphology
in coelurosaurian theropods. Journal of Vertebrate Paleontology. 27(3), 139A.
Xu and Zhao, 2007. Coelurosaurian phylogeny revisited: Recovering phylogenetic
signals from subtle morphological variations. Journal of Vertebrate Paleontology.
27(3), 169A.
Turner, 2008. Phylogenetic history and body size evolution in coelurosaur theropods.
Journal of Vertebrate Paleontology. 28(3), 154A.
Zhang, Kearns, Benton and Zhou, 2009. The ultrastructure of skin and feathers
of Cretaceous birds and dinosaurs. Journal of Vertebrate Paleontology. 29(3),
206A.
Zanno and Makovicky, 2010. Quantitative analysis of herbivorous ecomorphology
in theropod dinosaurs: Patterns of character correlation and progression. Journal
of Vertebrate Paleontology. Program and Abstracts 2010, 192A.
Loewen, Zanno, Irmis, Sertich and Sampson, 2011. Campanian theropod evolution
and intracontinental endemism on Laramidia. Journal of Vertebrate Paleontology.
Program and Abstracts 2011, 146.
Xu, You, Du and Han, 2011. An Archaeopteryx-like theropod from China
and the origin of Avialae. Nature. 475, 465-470.
Balanoff, Bever, Rowe and Norell, 2012. The origin of the avain brain based
on a volumetric analysis of endocranial evolution within Coelurosauria. Journal
of Vertebrate Paleontology. Program and Abstracts 2012, 59.
Brusatte, 2013. The phylogeny of coelurosaurian theropods (Archosauria: Dinosauria)
and patterns of morphological evolution during the dinosaur-bird transition.
Journal of Vertebrate Paleontology. Program and Abstracts 2013, 96.
Larson, Brown and Evans, 2013. Disparity dynamics of small theropod (Coelurosauria:
Dinosauria) tooth assemblages from the Late Cretaceous of North America. Journal
of Vertebrate Paleontology. Program and Abstracts 2013, 159.
Torices, Bradley and Currie, 2013. Ontogenetic variability in Upper Cretaceous
theropod teeth. Journal of Vertebrate Paleontology. Program and Abstracts 2013,
226.
Bicentenaria Novas, Ezcurra,
Agnolin, Pol and Ortiz, 2012
B. argentina Novas, Ezcurra, Agnolin, Pol and Ortiz, 2012
Early Cenomanian, Late Cretaceous
Candelaros Formation of the Rio Limay Subgroup of the Neuquen Group, Rio Negro,
Argentina
Holotype- (MPCA 865) (adult) incomplete jugal, quadratojugal, incomplete
quadrate, partial ectopterygoid, basisphenoid fragment, surangulars (one incomplete),
angular, incomplete prearticulars, articular
Paratype- (MPCA 866) (at least three adults; ~3 m; ~40 kg) ~130 elements
including two partial premaxillae, seventeen fragmentary dorsal vertebrae including
third and fourth dorsal centra, several dorsal rib fragments including proximal
anterior dorsal rib, six partial sacra, two sacral centra, twenty caudal vertebrae,
two fragmentary scapulae, coracoid fragment, proximal humerus, two distal humeri,
distal radius, three proximal ulnae, eight manual unguals, partial ilium, five
proximal pubes, four incomplete femora (~310 mm), two incomplete tibiae, distal
tibia, partial astragalus, metatarsal I, four fragmentary metatarsals including
distal metatarsals III and IV, fifteen pedal phalanges, eight pedal unguals
(juvenile) maxillary fragment, distal femur
Diagnosis- (modified after Novas et al., 2012) second premaxillary tooth
with mesial serrations limited to base of crown; anterior quadratojugal process
twice as long as dorsal process; lateral quadrate condyle much larger than medial
condyle; surangular with raised trapezoidal dorsal margin in lateral view; retroarticular
process dorsoventrally depressed, transversely wide and spoon-shaped; proximal
humerus anteroposteriorly compressed; deep fossa proximal to ectocondyle on
distal humerus; manual ungual III with proximodorsal lip.
Comments- The material was discovered in 1998. Novas et al. (2012) included
it in a version of the TWG analysis that recovered Bicentenaria as a non-tyrannoraptoran coelurosaur. Brusatte et al. (2014) and Cau (2018) recovered the same placement.
References- Novas, Ezcurra, Agnolin, Pol and Ortiz, 2012. New Patagonian
Cretaceous theropod sheds light about the early radiation of Coelurosauria.
Revista del Museo Argentino de Ciencias Naturales. 14(1), 57-81.
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.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25.
Siamotyrannus Buffetaut,
Suteethorn and Tong, 1996
S. isanensis Buffetaut, Suteethorn and Tong, 1996
Late Barremian, Early Cretaceous
Phu Wiang 9, Sao Khua Formation, Thailand
Holotype- (SM-PW9-1) (~6.5-7 m, adult) incomplete fourth
dorsal vertebra (87 mm), incomplete fifth dorsal vertebra (80 mm),
incomplete sixth dorsal vertebra (80 mm), seventh dorsal centrum (96
mm), anterior eighth dorsal centrum, dorsal rib fragment, incomplete
synsacrum (120, 117, 91, 89, 115, 115 mm), partial first caudal
vertebra (110 mm), partial second caudal vertebra (115 mm), partial
third caudal vertebra (120 mm), fourth caudal vertebral fragment (112
mm), partial fifth caudal vertebra (130 mm), incomplete sixth caudal
vertebra (122 mm), incomplete seventh caudal vertebra (130 mm),
incomplete eighth caudal vertebra (123 mm), incomplete ninth caudal
vertebra (133 mm), incomplete tenth caudal vertebra (126 mm),
incomplete eleventh caudal vertebra (134 mm), incomplete ?twelfth
caudal vertebra (122 mm), incomplete ?thirteenth caudal vertebra (114
mm), nine chevrons (c5 275 mm), ilium (800 mm), pubis (860 mm),
incomplete ischium, ischial fragment
Diagnosis- (after Buffetaut et al., 1996)
partially closed obturator notch in pubis; pubic boot more developed anteriorly than posteriorly.
(after Rauhut, 2003) two parallel vertical ridges on ilial blade above and in front of acetabulum.
(after Samathi et al., 2015) anterior to mid dorsal parapophyses with
long and stalk-like pedicels; notch on dorso-posterior part of
postacetabular process (not shown in the original description where the
pelvic photo was roughly cut out from its background; but see Carrano et
al., 2012: Fig. 6).
Other diagnoses- Buffetaut et al. (1996) originally diagnosed Siamotyrannus
with a long list of characters, many primitive for avetheropods-
proximal caudal vertebrae with tall, slender neural spines; mid caudal
vertebrae with accessory neural spines; proximal chevrons long,
straight and slender; long and relatively low ilium; pubis with long,
straight shaft terminating in massive distal boot; no ambiens crest on
pubis; or coelurosaurs- preacetabular process with incipient cuppedicus
shelf; well defined proximolateral ischial scar; ischium slender. A
final character, ischium curved (ventrally), is common in carnosaurs
and basal coelurosaurs.
Comments-
This was discovered in 1993 and first mentioned by "an incomplete
skeleton of a large theropod" ... "which includes a well preserved
pubis with a a large distal "boot" and by Suteethorn et a. (1995) as "a
partly articulated skeleton of a large theropod, including the dorsal
and caudal vertebrae, the sacrum and a large part of the pelvis."
It was briefly described by Buffetaut et al.
(1996) as a new taxon of tyrannosaurid. Samathi has made a
project of redescribing the specimen in detail, in both his Masters
(2013) and PhD (2019) theses. Meeting abstracts about the dorsal
vertebrae (Samathi et al., 2008; Samathi, 2015) and the project in
general (Samathi et al., 2013, 2015) have been published, but not the
osteology itself.
Buffetaut and Suteethorn (1998) say "a recently discovered theropod tibia from Phu Wiang, in all likelihood referable to Siamotyrannus, is not unlike that of the primitive tyrannosaurid Alectrosaurus",
while Buffetaut and Suteethorn (1999) say "a well-preserved tibia found
at Phu Wiang, together with a partial sacrum resembling the type of Siamotyrannus isanensis,
exhibits tyrannosaurid features and very probably belongs to this
taxon." These were partially addressed by Samathi et al. (2019a)
who stated "the tibia with partial sacrum provisionally assigned to Siamotyrannus
by Buffetaut and Suteethorn (1999) more probably belongs to another
taxon of theropod found in the Phu Wiang locality (see below)", and
while they never reference this again, Phuwiangvenator
is the only taxon that preserves these elements (and has parts
discovered since 1993). Buffetaut and Suteethorn (1999) stated "a
maxilla fragment containing
compressed teeth, also from Phu Wiang, [which] may also belong to Siamotyrannus,
but it is very incomplete", which has since been described as
carcharodontosaurid by Buffetaut and Suteethorn (2012).
Similarly, while Buffetaut and Suteethorn (1998) say "abundant
blade-like, serrated theropod teeth from the Sao Khua Formation
probably belong to Siamotyrannus", Phuwiangvenator and Vayuraptor
have since been described from the formation and likely had similar
teeth. These generic theropod teeth are listed here under Averostra.
Buffetaut et al.'s (1996) original assignment to Tyrannosauridae would
today be recognized as a placement in basal Tyrannosauroidea, as
albertosaurines and tyrannosaurines were listed as more derived.
Pharris (DML, 1997) made persuasive arguments for the assignment of Siamotyrannus
to Sinraptoridae (= Metriacanthosauridae), but neither of these early
studies were based on quantitative analyses. The first
quantitavive analyses to include Siamotyrannus
in a theropod sample including more than tyrannosauroids recovered it
as a carnosaur/allosauroid (Rauhut, 2000 and 2003; Holtz et al., 2004),
although the latter had it sister to Fukuiraptor which is now often recognized as a coelurosaur. More recently, Carrano et al. (2012) found Siamotyrannus to be a derived metriacanthosaurine, echoing Pharris' idea. Samathi (2015) updated Siamotyrannus
in that matrix and found it to have "a 'basal' coelurosaur position",
but as Samathi et al. (2015) noted "the alternative possibility that it
might belong to basal Allosauria or Metriacanthosauridae cannot be
rejected" at only three more steps. Samathi and Chanthasit (2017) reported that in Novas et al.'s tetanurine matrix, Siamotyrannus
falls out as the sister taxon of Tyrannoraptora, with megaraptorans
more stemward. A variant of this is shown in the analysis of
Samanthi et al. (2019b) where it falls out sister to Megaraptora plus
Tyrannoraptora along with Chilantaisaurus and Gualicho.
References- Buffetaut, Suteethorn, Martin, Tong, Chaimanee and Triamwichanon,
1995. New dinosaur discoveries in Thailand. In Wannakao, Srisuk, Youngme and
Lertsirivorakul (eds.). International Conference on Geology,
Geotechnology and Mineral Resources of Indochina (Geo-Indo 1995). 157-161.
Suteethorn, Chaimanee, Triamwichanon, Suksawat, Kamsupha, Kumchoo, Buffetaut,
Martin and Tong, 1995. Thai dinosaurs; An updated review. [Academic Conference, Department of Geology]. 129-133.
Buffetaut, Suteethorn and Tong, 1996. The earliest known
tyrannosaur from the Lower Cretaceous of Thailand. Nature. 381(6584), 689-691.
Pharris, DML 1997. https://web.archive.org/web/20201115172819/http://dml.cmnh.org/1997Jun/msg00271.html
Buffetaut and Suteethorn, 1998. Early Cretaceous dinosaurs from Thailand and
their bearing on the early evolution and biogeographical history of some groups
of Cretaceous dinosaurs. In Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History Bulletin.
14, 205-210.
Buffetaut and Suteethorn, 1999. The dinosaur fauna of the Sao Khua
Formation of Thailand and the beginning of the Cretaceous radiation of
dinosaurs in Asia. Palaeogeography, Palaeoclimatology, Palaeoecology.
150, 13-23.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis. University of Bristol. 440 pp.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 213 pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Samathi, Srikosamatra and Suteethorn, 2008. Additional preserved
vertebrae of Siamotyrannus isanensis (Dinosauria; Theropoda) from the
type locality. 9th Science Exhibition, Faculty of Science, Mahidol
University. 75-76.
Buffetaut and Suteethorn, 2012. A carcharodontosaurid theropod
(Dinosauria, Saurischia) in the Sao Khua Formation (Early Cretaceous, Barremian)
of Thailand. 10th Annual Meeting of the European Association of Vertebrate Palaeontologists. Fundamental. 20, 27-30.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Samathi, 2013. Osteology
and phylogenetic position of Siamotyrannus isanensis
(Dinosauria; Theropoda) from the Lower Cretaceous of Thailand.
Masters Thesis, Ludwig-Maximi�lians-Universit�t
M�nchen. 69 pp.
Samathi, Butler and Chanthasit, 2013. Osteology and phylogenetic position of Siamotyrannus isanensis (Dinosauria; Theropoda) from the Lower Cretaceous of Thailand. Evolution, Ecology and Systematics Conference. [pp]
Samathi, 2015. New information on dorsal vertebrae of Siamotyrannus isanensis
(Dinosauria, Theropoda) from the Lower Cretaceous of Thailand. 13th
Annual Meeting of the European Association of Vertebrate
Palaeontologists. Abstract Volume, 132.
Samathi, Butler and Chanthasit, 2015. A revision of Siamotyrannus isanensis (Dinosauria:Theropoda) from the Early Cretaceous of Thailand. The 2nd International Symposium on Asian Dinosaurs. 30-31.
Samathi and Chanthasit, 2017. Two new basal Megaraptora
(Dinosauria: Theropoda) from the Early Cretaceous of Thailand with
comments on the phylogenetic position of Siamotyrannus and Datanglong. Journal of Vertebrate Paleontology. Program and Abstracts, 188.
Samathi, 2019. Theropod dinosaurs from Thailand and southeast Asia
Phylogeny, evolution, and paleobiogeography. PhD thesis, Rheinischen
Friedrich-Wilhelms-Universit�t Bonn. 249 pp.
Samathi, Chanthasit and Sander, 2019a. A review of theropod dinosaurs
from the Late Jurassic to mid-Cretaceous of southeast Asia. Annales de
Pal�ontologie. 105(3), 201-215.
Samathi, Chanthasit and Sander, 2019b. Two new basal coelurosaurian
theropod dinosaurs from the Lower Cretaceous Sao Khua Formation of
Thailand. Acta Palaeontologica Polonica. 64(2), 239-260.
Coelurosauria incertae sedis
"Aratasaurus" Say�o, Saraiva, Brum, Bantim, Andrade, Cheng, Lima, Paula Silva and Kellner, 2020
"A. museunacionali" Say�o, Saraiva, Brum, Bantim, Andrade, Cheng, Lima, Paula Silva and Kellner, 2020
Albian, Early Cretaceous
Mina Pedra Branca, Romualdo Formation of Santana Group, Brazil
Material- (MPSC R 2089) (~3 m, ~34 kg) (four year old juvenile)
distal femur, proximal tibia (~412 mm), incomplete metatarsal I,
phalanx I-1 (23 mm), pedal ungual I (10 mm), distal metatarsal II,
phalanx II-1 (62 mm), phalanx II-2 (40 mm), pedal ungual II (30 mm),
distal metatarsal III (~243 mm), phalanx III-1 (65 mm), phalanx III-2
(48 mm), phalanx III-3 (41 mm), pedal ungual III (32 mm), distal
metatarsal IV, phalanx IV-1 (42 mm), phalanx IV-2 (32 mm), phalanx IV-3
(17 mm), phalanx IV-4 (17 mm)
Diagnosis- (after Say�o et al.,
2020) tibia exhibiting a proximomedial fossa; distal condyles of
metatarsi II, III and IV symmetric mediolaterally and with subequal
width; width of metatarsi II and IV similar, presenting the dorsal
surface of the distal articulation bulbous; symmetric pes, with digits
II and IV subequal in total length.
Comments- The specimen was
discovered at least by 2016 and was later described by Say�o et al. on
July 10 2020 as a new taxon. However, this paper has no mention
of ZooBank although the taxon does have an entry on the ZooBank
website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Aratasaurus museunacionali"
Say�o et al., 2020 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically. Say�o et al. added the specimen to
Choiniere's coelurosaur analysis, recovering it as sister to Zuolong, the pair being the first branching coelurosaurs.
Reference- Say�o, Saraiva,
Brum, Bantim, Andrade, Cheng, Lima, Paula Silva and Kellner, 2020. The
first theropod dinosaur (Coelurosauria, Theropoda) from the base of the
Romualdo Formation (Albian), Araripe basin, northeast Brazil.
Scientific Reports. 10:10892.
Iliosuchidae
Iliosuchus Huene, 1932
I. incognitus Huene, 1932
= Megalosaurus incognitus (Huene, 1932) Romer, 1966
Middle Bathonian, Middle Jurassic
Stonesfield Slate, England
Holotype- (NHMUK R83) incomplete ilium
Referred- (OUM J29780) partial ilium (~93 mm) (Galton, 1976)
Comments- Foster and Chure (2000) and Galton and Molnar (2005) referred
OUM J28971 to Iliosuchus, but it was removed by Benson (2009).
This taxon is traditionally allied with Stokesosaurus and thus with tyrannosauroids
due to the vertical ilial ridge. The concave anterior edge on the pubic peduncle
is also a tyrannosauroid-like character. Yet both are actually widespread among
basal coelurosaurs. The reduced ischial peduncle is similar to coelurosaurs and Concavenator,
and the mediolaterally narrow pubic peduncle is similar to most avetheropods.
Benson and Carrano et al. (2012) considered Iliosuchus an indeterminate
avetheropod or juvenile Megalosaurus, despite not considering Megalosaurus
to be an avetheropod. Yet Iliosuchus differs from Megalosaurus
in the characters noted above (Megalosaurus' vertical ridge is much lower)
in addition to having a more anteriorly angled pubic peduncle which lacks a
concave posterior margin. It may also possess a unique combination of characters
when compared to other taxa, even if it lacks autapomorphies. It is provisionally
retained here as a valid genus of coelurosaur.
References- Phillips, 1871. Geology of Oxford and the Valley of the Thames:
Oxford at the Clarendon Press. 523 pp.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Galton, 1976. Iliosuchus, a Jurassic dinosaur from Oxfordshire and Utah.
Palaeontology. 19, 587-589.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Foster and Chure, 2000. An ilium of a juvenile Stokesosaurus (Dinosauria,
Theropoda) from the Morrison Formation (Upper Jurassic: Kimmeridgian), Meade
County, South Dakota. Brigham Young University Geology Studies. 45, 5-10.
Galton and Molnar, 2005. Tibiae of small theropod dinosaurs from Southern England.
In Carpenter (Ed.). The Carnivorous Dinosaurs. 3-22.
Benson, 2009. An assessment of variability in theropod dinosaur remains from
the Bathonian (Middle Jurassic) of Stonesfield and New Park Quarry, UK and taxonomic
implications for Megalosaurus bucklandii and Iliosuchus incognitus.
Palaeontology. 52(4), 857-877.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
undescribed coelurosaur (Kirkland, Lucas and Estep, 1998)
Early Albian, Early Cretaceous
Lorrie's Site, Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- (DMNH coll.) (small) tibia
Comments- Kirkland et al. (1998) list Coelurosauridae new genus and species
under the Middle Cedar Mountain Formation, which includes the Ruby Ranch and
Poison Strip Members. Coelurosauridae is a misspelling of Coelurosauria. Kirkland
(online 2008) stated Lorrie's "site has also produced a tibia, or shin bone of some other small theropod", which is presumably
the same record.
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.
Kirkland, online 2008. https://web.archive.org/web/20090517064058/http://scientists.dmns.org/kenCarpenter/cedar%2Dmountain%2Dproject/dinosaurs%2Dof%2Dthe%2Dcedar%2Dmountain%2Dformation/
unnamed possible Coelurosauria (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
Cenomanian-Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (NCSM 33268) incomplete lateral tooth (~20x8.39x4.5 mm)
(NCSM coll.) twenty tooth fragments, eleven bone fragments
Comments- NCSM 33268 was referred to Coelurosauria indet. by Avrahami et al. (2018) and said to plot within Coelophysoidea, Nuthetes,
or "within or adjacent to Tyrannosauroidea and Dromaeosauridae."
Avrahami et al. stated "tooth and bone fragments are too poorly
preserved to allow for confident lower level taxonomic
identifications", but these are probably coelurosaurs given their age
and provenence.
References- 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.
undescribed Coelurosauria (Metcalf and Walker, 1994)
Early Bathonian, Middle Jurassic
Chipping Norton Formation, England
Material- (GLRCM coll.; B) tooth (2.3 mm; FABL 2.6 mm)
(GLRCM coll.; G) tooth (1.9 mm; FABL 1.7 mm)
Comments- These two teeth were labeled as "dromaeosaur-like"
by Metcalf and Walker (1994).
Teeth B and G in their figure 18.7 exhibit similar morphology, so may belong
to the same taxon. Mesial serrations are present apically, while the crowns
are short and slightly recurved. B and G have DSDIs of 1.17 and 1.4 respectively.
Serrations are fairly flat and not hooked apically, but are taller than wide
on the distal carina. Serration density is 5-6/mm on B, and 12/mm on G. G may
have a constricted base, though blood grooves are not apparent on either specimen.
They resemble those referred to posterior teeth of cf. Compsognathus sp.
by Zinke (1998) except that B has a slightly lower serration count.
References- Metcalf and Walker, 1994. A new Bathonian microvertebrate
locality in the English Midlands. in Fraser and Sues (eds.). In the Shadow of
the Dinosaurs- Mesozoic Small Tetrapods, Cambridge (Cambridge University Press).
322-332.
Zinke, 1998. Small theropod teeth from the Upper Jurassic coal mine of Guimarota
(Portugal). Palaontologische Zeitschrift. 72(1/2) 179-189.
undescribed coelurosaur (Austen, Brockhurst and Honeysett, 2010)
Valanginian, Early Cretaceous
Wadhurst Clay of the Hastings Group, England
Material- (BEXHM : 2010.3) cervical centrum
Comments- Austen et al. (2010) compared this to Ornithodesmus,
but Naish and Sweetman (2011) correctly noted they do not share comparable elements.
References- Austen, Brockhurst and Honeysett, 2010. Vertebrate fauna
from Ashdown Brickworks, Bexhill, East Sussex. Wealden News. 8, 13-23.
Naish and Sweetman, 2011. A tiny maniraptoran dinosaur in the Lower Cretaceous
Hastings Group: Evidence from a new vertebrate-bearing locality in south-east
England. Cretaceous Research. 32(4), 464-471.
undescribed possible coelurosaur (Seeley, 1887)
Barremian, Early Cretaceous
Upper Weald Clay Formation of the Weald Clay Group, England
Material- incomplete pubis (Seeley, 1887)
Comments- Seeley (1887) mentions a "pubis, imperfect distally, of
a type very similar to Coelurus, from Tilgate" when discussing Aristosuchus,
though the specimen has not been located since.
Reference- Seeley, 1887. On Aristosuchus pusillus Owen, being
further notes on the fossils described by Sir R. Owen as Poikilopleuron pusillus.
Owen. Quarterly Journal of the Geological Society of London. 43, 221-228.
unnamed Coelurosauria (Lydekker, 1888)
Barremian, Early Cretaceous
Wessex Formation, England
Material- (Dinosaur Expeditions Centre coll.) two distal metatarsals (Mattsson pers. comm.,
2015)
(IWCMS 1995.208) ulna (Hutt, 2001)
(MIWG 5137; cast as NHMUK R9230) tibia (171.8 mm) (Carrano, 1998)
(MIWG 5823) vertebra (Hutt, 2001)
(MIWG 5824) vertebra (Hutt, 2001)
(NHMUK R899) partial manual ungual (Lydekker, 1888)
(NHMUK R5194) proximal femur (Galton, 1973)
(NHMUK R6424; = BMNH R6426 of Naish, 2002) proximal ischium (Naish, 2002)
Comments- Lydekker (1888) referred a manual ungual (BMNH R899) to Aristosuchus,
but Naish (2002) notes it has a low and distally positioned flexor
tubercle unlike the paratype ungual so may be from another species. It
was illustrated in Naish et al. (2001) as an indeterminate
theropod. Note the NHMUK online catalogue incorrectly indicates
this is now NHMUK R889, although the original number is still written
on the fossil, and the physical catalogue lists R899 as "Ungual
phalanx. imperfect anteriorly" and R889 as "Plastron and costal plates
of chelonian".
NHMUK R5194 was originally collected in 1917 and catalogued as Hypsilophodon, but described
by Galton (1973) and referred to Aristosuchus.
Tibia MIWG 5137 was discovered in 1976, and described by Naish (1999) as possibly being Aristosuchus as it was said to be virtually identical to Mirischia.
It has since been figured by Naish et al. (2001) as "a possible compsognathid theropod."
Carrano (1998) listed the tibia BMNH R.9230 as Calamosaurus foxi, which based on the photo in the online NHMUK catalogue is a cast of MIWG 5137 made in 1977.
Hutt (2001) listed IWCMS 1995.208, MIWG 5823 and MIWG 5824 as Aristosuchus
sp., but the specimens are undescribed.
Naish (2002) illustrated partial ischium BMNH R6426 as a possible Aristosuchus
specimen, but the NHMUK online catalogue indicates this is actually
NHMUK 6424. Unfortunately, the physical catalogue states its
history is unknown but was acquired in 1953.
Mattsson (pers. comm., 2015) informs me of two distal metatarsals discovered
in 1996 and initially identified as testudine elements, which are now on display
at the Dinosaur Expeditions Centre.
These specimens may belong to Aristosuchus, Calamosaurus, Calamospondylus
(difficult to determine as the type is lost and poorly described), Ornithodesmus,
Thecocoelurus and/or Yaverlandia and have been compared to basal
tyrannosauroids, coelurids and compsognathids. Additionally, femur MIWG 6214
(Naish, 2000), tibia NHMUK R186 (Lydekker, 1888), and the two fragmentary skeletons
exhibited as Calamosaurus at the Dinosaur Expeditions Centre (Mattsson,
pers. comm. 2015) may belong to the same taxa, though these have been recently
hypothesized to be ornithomimosaurs (Allain et al., 2014).
References- 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, London. 309 pp.
Galton, 1973. A femur of a small theropod dinosaur from the Lower Cretaceous
of England. Journal of Paleontology. 47, 996-997.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Naish, 1999. Studies on Wealden Group theropods - An investigation into the
historical taxonomy and phylogenetic affinities of new and previously neglected
specimens. Masters thesis, University of Portsmouth. 184 pp.
Naish, 2000. A small, unusual theropod (Dinosauria) femur from the Wealden Group
(Lower Cretaceous) of the Isle of Wight, England. Neues Jahrbuch f�r Geologie
und Pal�ontologie Monatshefte. 2000, 217-234.
Hutt, 2001. Catalogue of Wealden Group Dinosauria in the Museum of Isle of Wight
Geology. In Martill and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological
Association. 411-422.
Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: Theropods. In Martill
and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association.
242-309.
Naish, 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria)
Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings
of the Geologists' Association. 113, 153-163.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria,
Theropoda): An undetected record. Geologica Acta. 12(2), 127-135.
unnamed possible coelurosaur (Clark, 2005)
Late Bajocian-Early Bathonian, Middle Jurassic
Vlatos Sandstone Formation, Scotland
Material- (GLAHM 101240) (~2 m) incomplete mid caudal vertebra (31 mm)
Comments- While it was originally referred to Coelophysoidea (Clark,
2005), Brusatte and Clark (2015) found the vertebra is most similar to basal
coelurosaurs.
References- Clark, 2005. Tracking dinosaurs in Scotland. Open University
Geological Society Journal. 26, 30-35.
Brusatte and Clark, 2015. Theropod dinosaurs from the Middle Jurassic (Bajocian-Bathonian)
of Skye, Scotland. Scottish Journal of Geology. 51(2), 157-164.
unnamed possible Coelurosauria (Barco and Ruiz-Ome�aca 2001)
Tithonian-Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (LC-1) partial tooth (?x15.1x10.2 mm) (Su�er, Santisteban
and Galobart, 2005)
(MPZ01/98) partial tooth (Barco and Ruiz-Ome�aca 2001)
References- Barco and Ruiz-Ome�aca, 2001. Primeros dientes de
ter�podos (Dinosauria, Saurischia) en la Formaci�n Villar del
Arzobispo (Tit�nico-Berriasiense): Yacimientos Cuesta Lonsal y Las Cerradicas
2 (Galve, Teruel). Publicaciones del Seminario de Paleontolog�a de Zaragoza.
5, 239-246.
Su�er, Santisteban and Galobart, 2005. Nuevos restos de Theropoda del
Jur�sico Superior-Cret�cico Inferior de la Comarca de Los Serranos
(Valencia). Revista Espa�ola de Paleontologia. N. Extra X, 93-99.
undescribed coelurosaur (Gasulla, Ortega, Escaso and Sanz, 2006)
Early Aptian, Early Cretaceous
Arcillas de Morella Formation, Spain
Material- (CMP-3-743) distal tibia
Reference- Gasulla, Ortega, Escaso and Sanz, 2006. Diversidad de ter�podos
del Cret�cico Inferior (Fm. Arcillas de Morella, Aptiense) en los yacimientos
del Mas de la Parreta (Morella, Castell�n). In Fern�ndez-Mart�nez
(ed.). XXII Jornadas de Paleontolog�a de la Sociedad Espa�ola
de Paleontolog�a. Libro de res�menes, 124-125.
undescribed coelurosaur (Torices, Barroso-Barcenilla, Cambra-Moo, Perez
and Serrano, 2011)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- ungual
Comments- Torices et al. (2011) mention Coelurosauridae indet..
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.
unnamed possible coelurosaur (Lanser and Heimhofer, 2015)
Late Barremian-Early Aptian, Early Cretaceous
Balve-Beckum quarry, Germany
Material- (LWL MN Ba 14) anterior tooth (12.7x5.5x4.6 mm)
Reference- Lanser and Heimhofer, 2015 (online 2013). Evidence of theropod dinosaurs
from a Lower Cretaceous karst filling in the northern Sauerland (Rhenish Massif,
Germany). Pal�ontologische Zeitschrift. 89(1), 79-94.
possible Coelurosauria (Jurcsak, 1982)
Berriasian-Hauterivian, Early Cretaceous
Bauxite of Cornet, Romania
Material- (MTCO 16499) cervical vertebra
(MTCO 17245) caudal centrum
Comments- These were assigned to Aristosuchus sp. by Jurcsak (19820
and Jucsak and Popa (1983). They may not belong to the same taxon, and the relationship
of either with Aristosuchus is uncertain.
References- Jurcsak, 1982. Occurrences nouvelles des Sauriens mesozoiques
de Roumanie. Vertebrata Hungarica. 21, 175-184.
Jurcsak and Popa, 1983. La faune de dinosauriens du Bihor (Roumanie). In Buffetaut,
Mazin and Salmon (eds.). Actes du Symposium paleontologique Georges Cuvier.
325-335.
unnamed Coelurosauria (Knoll and Ruiz-Omenaca, 2005)
Beriassian, Early Cretaceous
KM 1983, Ksar Metlili Formation, Morocco
Material- (MNHN SA 2004/1; Maniraptora indet. Morphotype I; lost) tooth (?x8.20x4.00 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/3C; Maniraptora indet. Morphotype I; lost) tooth (2.76x1.80x.68 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/3E; Maniraptora indet. Morphotype I; lost) tooth (3.44x1.88x.80 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/3F; Maniraptora indet. Morphotype I; lost) tooth (?x?x.64 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/4B; Maniraptora indet. Morphotype I; lost) tooth (1.48x1.08x.40 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/4D; Maniraptora indet. Morphotype I; lost) tooth (2.28x1.52x.88 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/4E; Maniraptora indet. Morphotype III; lost) tooth (?x1.60x.72 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/5A; Maniraptora indet. Morphotype II; lost) anterior tooth (?x2.60x1.32 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA mcm 153; Maniraptora indet. Morphotype I; lost) partial tooth (?x1.92x1.00 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA mcm 162; Maniraptora indet. Morphotype I; lost) tooth (1.48x1.08x.52 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA mcm 166; Maniraptora indet. Morphotype I; lost) tooth (1.52x?x.56 mm) (Knoll and Ruiz-Omenaca, 2009)
Beriassian, Early Cretaceous
KM-A1, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-A1-13; Theropoda gen. et sp. indet. morphotype I) lateral
tooth (2.65x1.63x.84 mm), tooth (1.87x1.50x.82 mm (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-A2, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-A2-5; Coelurosauria gen. et sp. indet. morphotype I) lateral tooth (Lasseron, 2020)
(FSAC-KM-A2-9 [2]; Theropoda gen. et sp. indet. morphotype I) lateral tooth (5.21x2.59x1.26x mm) (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-B', Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-B'-22; Theropoda gen. et sp. indet. morphotype I) lateral tooth (2.09x1.18x1.03 mm) (Lasseron, 2020)
(FSAC-KM-B'-24; Theropoda gen. et sp. indet. morphotype V) lateral tooth (2.61x1.58x1.17 mm) (Lasseron, 2020)
(FSAC-KM-B'-26 [1]; Coelurosauria gen. et sp. indet. morphotype I) lateral tooth (2.54x1.78x1.20 mm) (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-D1, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-D1-6; Coelurosauria gen. et sp. indet. morphotype I) lateral tooth (3.31x2.31x1.21 mm) (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-D2, Ksar Metlili, Ksar Metlili Formation, Morocco
?(FSAC-KM-D2-11; Theropoda gen. et sp. indet. morphotype I) lateral tooth (Lasseron, 2020)
Comments-
The MNHN specimens were collected in 1983, 1986 and 1999, while the
FSAC specimens were collected in 2010, 2015 and 2018 (Lasseron,
2020). Lasseron noted the "dinosaur remains (Knoll, 2000; Knoll & Ruiz-Ome�aca, 2009) ... were taken out of the MNHN and lost." Note
Lasseron listed FSAC-KM-D2-11 as both a 'Theropoda gen. et sp.
indet. morphotype I' and 'Theropoda gen. et sp. indet. morphotype II'
tooth, without using a number in brackets to designate multiple teeth
falling under one specimen number. As the mesial serration
density is
listed, the measurements given belong to a morphotype II tooth.
The MNHN specimens are referred to Maniraptora by Knoll and
Ruiz-Omenaca (2009) as they lack mesial serrations, but these are
also absent in several other coelurosaurs (e.g. megaraptorans,
compsognathid-grade maniraptoromorphs). Lasseron stated his
Theropoda gen. et sp. indet. morphotype I were similar to Knoll and
Ruiz-Omenaca's Maniraptora indet. Morphotype I, while his Coelurosauria
gen. et sp. indet. morphotype I is similar to their Maniraptora indet.
Morphotype III. Knoll and Ruiz-Omenaca stated Maniraptora indet.
Morphotype I resembled Tsaagan, Hell Creek Saurornitholestes and European teeth described as Dromaeosauridae indet., while Morphotype 2 was similar to Juravenator posterior premaxillary teeth, and Morphotype III most like juvenile Saurornitholestes and Microraptor
teeth. Lasseron finds quantitative analysis classifies
FSAC-KM-D2-11 as noasaurid, although it overlaps with most groups of
small theropods in the figure. Lasseron et al. list all four
teeth under FSAC-KM-B'-26 and FSAC-KM-D1-6 as Dromaeosauridae and both
FSAC-KM-A2-9 teeth and FSAC-KM-B'-22 as Maniraptora.
References- Knoll and Ruiz-Omenaca, 2005. Theropod teeth from the Berriasian
of Anoual (Morocco). Journal of Vertebrate Paleontology. 25(3), 78A.
Knoll and Ruiz-Omenaca, 2009. Theropod teeth from the basalmost Cretaceous of
Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine.
146(4), 602-616.
Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020 (online 2019). New data on the microvertebrate fauna from
the Upper Jurassic or lowest Cretaceous of Ksar Metlili (Anoual
Syncline, eastern Morocco). Geological Magazine. 157, 367‑392.
unnamed coelurosaur (Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016)
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of the Rio Limay Subgroup, Rio Negro, Argentina
Material-(MPCA-Pv
806) (adult) incomplete anterior cervical vertebra, incomplete distal
caudal vertebra, three incomplete caudal centra, two partial manual
unguals, distal metapodial
Comments-
Motta et al. identify this as Coelurosauria gen. et sp. indet..
Characteristics include elongate cervical vertebra without ventral
groove or keel and low neural spine; neural spine on amphiplatyan
distal caudal vertebrae; proximally positioned manual ungual flexor
tubercle and no proximodorsal lip.
Reference- Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016. New theropod fauna from the Upper Cretaceous (Huincul
Formtation) of northwestern Patagonia, Argentina. In Khosla and Lucas (eds.).
Cretaceous period: Biotic diversity and biogeography. New Mexico Museum of Natural
History and Science Bulletin. 71, 231-253.
unnamed coelurosaur (Canudo, Filippi, Salgado, Garrido, Cerda, Garcia
and Otero, 2009)
Late Coniacian-Early Santonian, Late Cretaceous
Plottier Formation of the Rio Neuquen Subgroup, Neuquen, Argentina
Material- (MAU-PV-PH-447/1) tooth (~23.3 mm)
(MAU-PV-PH-447/3) tooth (21.2 mm)
(MAU-PV-PH-447/5) tooth (19.6 mm)
(MAU-PV-PH-447/8) tooth (14.14 mm)
(MAU-PV-PH-462) tooth
Comments- Canudo et al. (2009) referred these to Maniraptora based on
the absence of mesial serrations, but this is known for several
other coelurosaurs (e.g. megaraptorans, compsognathid-grade maniraptoromorphs).
Reference- Canudo, Filippi, Salgado, Garrido, Cerda, Garcia and Otero,
2009. Theropod teeth associated with a sauropod carcass in the Upper Cretaceous
(Plottier Formation) of Rinc�n de los Sauces. Actas de las IV Jornadas
Internacionales sobre Paleontolog�a de Dinosaurios y su Entorno. 321-330.
unnamed coelurosaur (Azevedo, Simbras, Furtado, Candeiro and Bergqvist,
2013)
Campanian-Maastrichtian, Late Cretaceous
Presidente Prudente Formation, Brazil
Material- (UFRJ-DG 390-R) proximal fibula
Reference- Azevedo, Simbras, Furtado, Candeiro and Bergqvist, 2013 (online 2012). First
Brazilian carcharodontosaurid and other new theropod dinosaur fossils from the
Campanian-Maastrichtian Presidente Prudente Formation, S�o Paulo State, southeastern Brazil. Cretaceous Research. 40, 131-142.
unnamed possible coelurosaur (Marshall, 1989)
Maastrichtian, Late Cretaceous
El Molino Formation, Bolivia
Material- (MHNC 3702) incomplete tooth (~18 mm)
Comments- Marshall (1989) referred this to Coelurosauria based on its
small size, but based on the completely serrated carinae and absence of wrinkles,
it may be a juvenile abelisaur instead.
Reference- Marshall, 1989. El primer diente de dinosaurio en Bolivia.
Revista T�cnica de Yacimientos Petrol�feros Fiscales Bolivia.
10(3-4), 129-130.
unnamed Coelurosauria (Rich and Vickers-Rich, 1994)
Early Aptian, Early Cretaceous
Wonthoggi Formation of the Strzelecki Group, Victoria, Australia
Material- (NMV P186353) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P186457) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P198947) tooth (21 x 9.5 x 5 mm) (Benson, Rich, Vickers-Rich and Hall,
2012)
(NMV P198958) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P199070) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P210025) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P210084) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P212859) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P221187) (juvenile) dorsal centrum (60 mm) (Kool, 1997)
(NMV P221204) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P221205) tooth (3.5 x 2 x 1 mm) (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P229111) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV P230871) tooth (Benson, Rich, Vickers-Rich and Hall, 2012)
(NMV coll.) 14 anterior teeth (mean 10 mm), 75 lateral teeth (mean 11.1 mm)
(Benson, Rich, Vickers-Rich and Hall, 2012)
Comments- Rich and Vickers-Rich (1994) reported the first eight theropod
teeth to be discovered at the Flat Rocks site, more of which were reported in
successive Dinosaur Dreaming field reports and briefly mentioned in papers describing
new taxa from the site. They have recently been described by Benson et al. (2012),
who state more detailed work is being done by Salisbury, Currie and Novas on
the more than ninety teeth known by then. While Kool (2003) noted that preliminary
study by Currie of the forty teeth known at the time indicated the presence
of four taxa of small theropod, and Rich (2004) stated study by Salisbury indicated
three taxa, Benson et al. described only one morphology as being present. Rich
and Vickers-Rich identified the teeth as dromaeosaurid, which has been followed
by most later authors who identified them past Theropoda. This was based on
the lack of mesial serrations, but this is known in many other coelurosaurs
(e.g. megaraptorans, compsognathids, most troodontids). Benson et al. labeled
them as possible megaraptorans.
NMV P221187 was first reported by Kool (1997) as a juvenile theropod. Benson
et al. referred this to Neovenatoridae instead of Tyrannosauridae based on the
ventral median keel, but this is present in Tyrannosaurus and other coelurosaurs
as well.
References- Rich and Vickers-Rich, 1994. Digs at Dinosaur Cove and Flat
Rocks 1994. Excavation Report. Dinosaur Cove 1993 - 1994 & Inverloch 1994.
10-13.
Kool, 1997. Dinosaur Dreaming 1997 Field Report. Dinosaur Dreaming 1997. Flat
Rocks Site Report. 1-2.
Kool, 2003. Dinosaur Dreaming 2003: Field report. Dinosaur Dreaming 2003 Report.
3-11.
Rich, 2004. Research update. Dinosaur Dreaming 2004 Field Report. 7-9.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from southern Australia
indicates high polar diversity and climate-driven dinosaur provinciality. PLoS
ONE. 7(5), e37122.
unnamed Coelurosauria (Rich and Vickers-Rich, 1994)
Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV 180880) incomplete pubis (Benson, Rich, Vickers-Rich and
Hall, 2012)
(NMV P186343) (small) tooth (Rich and Vickers-Rich, 1994)
(NMV P230845) incomplete femur (Benson, Rich, Vickers-Rich and Hall, 2012)
Comments- NMV P186343 was mentioned by Rich and Vickers-Rich (1994) as
a theropod tooth. Currie et al. (1996) believed it was dromaeosaurid-like. It
is possibly the specimen photographed on Pigdon's website, which is highly recurved,
with only distal serrations (2.5-3/mm). Besides basal dromaeosaurids, this is
known in many other coelurosaurs (e.g. megaraptorans, compsognathids, most
troodontids).
References- Rich and Vickers-Rich, 1994. Dig at Dinosaur Cove 1993. Excavation
Report. Dinosaur Cove 1993 - 1994 & Inverloch 1994. 1-8.
Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria)
specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia.
Alcheringa. 20(1-2), 73-79.
Pigdon, online 1997. http://home.alphalink.com.au/~dannj/austdino.htm
Rich and Vickers-Rich, 1997. Future directions for dinosaur research in Australia.
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. 275-277.
Long, 1998. Dinosaurs of Australia and New Zealand and other animals of the
Mesozoic Era. Harvard University Press. 192 pp.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from southern Australia
indicates high polar diversity and climate-driven dinosaur provinciality. PLoS
ONE. 7(5), e37122.
unnamed probable coelurosaur (Molnar, 1999)
Albian, Early Cretaceous
Griman Creek Formation, New South Wales, Australia
Material- (AM F103591) (juvenile) partial dorsal centrum
Comments- Molnar (1999) found this most closely resembles "Ichthyornis"
minusculus, an enantiornithine. This was based on the D-shaped articular
surface, which differs from Ichthyornis' circular surface. However, taxa
such as Confuciusornis and Microraptor also have D-shaped articulations,
as do some of Microvenator's centra. Unfortunately, the distribution
is hard to establish since small theropods generally don't preserve dorsal centra
in anterior or posterior view. The small size (centrum height 5.9 mm) probably
constrains it to Coelurosauria, even though it is a juvenile.
Reference- Molnar, 1999. Avian tibiotarsi from the Early Cretaceous of
Lightning Ridge, N.S.W. In Tomida, Rich and Rich (eds.). Proceedings of the Second
Gondwanan Dinosaur Symposium, National Sciences Museum Monographs. 15, 197-209.
Vayuraptor Samathi, Chanthasit and Sander, 2019
V. nongbualamphuensis Samathi, Chanthasit and Sander, 2019
Late Barremian, Early Cretaceous
Phu Wat A1, Sao Khua Formation, Thailand
Holotype- (SM-NB A1-2) (~4-4.5 m, adult) tibia (515 mm), astragalus (62.5 mm wide), calcaneum
Paratypes- ....(PRC-NB A1-3) ?pubic shaft fragment
....(PRC-NB A1-4) fibular fragment
....(PRC-NB A1-10) rib fragment
....(PRC-NB A1-11) incomplete coracoid
....(PRC-NB A1-16) ?pedal phalanx
....(PRC-NB A1-20) distal manual phalanx
Diagnosis- (after Samathi et
al., 2019) astragalus has two short horizontal grooves and two foramina
on body, and two fossae at base of ascending process; ascending process
of astragalus straight
laterally and straight and parallel medially at the base; at the middle
of the ascending process, the medial rim slopes to the tip laterally;
vertical ridge starting from tip and disappearing just above the middle
of the ascending process; extremely high and narrow ascending process
of astragalus, with ratio of the ascending process height/ascending
process width 1.66.
Comments- This was discovered
in August 1988, but not mentioned until Samathi and Chanthasit (2015),
who discussed it as a new megaraptoran. Note it gives the wrong
province (Khon Kaen), which instead corresponds to the locality of Phuwiangvenator.
Samathi (2016) mentions it in a later abstract as "a new megaraptoran
(PW A1-2)", and it is one of two specimens noted by Samathi and
Chanthasit (2017) as "basal members of Megaraptora" using both Carrano
et al.'s and Novas et al.'s tetanurine analyses. Vayuraptor
was described fully by Samathi et al. (2019), who added it to Novas et
al.'s analysis to now recover it as a coelurosaur in a trichotomy with
megaraptorans and tyrannoraptorans. Notably, they recover a
similar position for the co-occuring Siamotyrannus but find they cannot be compared (sharing only non-descript rib and pubic fragments) except for size, with the adult Vayuraptor being ~60% the size of adult Siamotyrannus.
References- Samathi and
Chanthasit, 2015. New megaraptoran (Dinosauria: Theropoda) from the
Early Cretaceous Sao Khua Formation of Thailand. 2nd International
Symposium on Asian Dinosaurs. 69.
Samathi, 2016. Theropod dinosaurs from Thailand and southeast Asia: A
review with newly found specimens. Young Natural History Scientists
Meeting. [pp]
Samathi and Chanthasit, 2017. Two new basal Megaraptora
(Dinosauria: Theropoda) from the Early Cretaceous of Thailand with
comments on the phylogenetic position of Siamotyrannus and Datanglong. Journal of Vertebrate Paleontology. Program and Abstracts, 188.
Samathi, Chanthasit and Sander, 2019. Two new basal coelurosaurian
theropod dinosaurs from the Lower Cretaceous Sao Khua Formation of
Thailand. Acta Palaeontologica Polonica. 64(2), 239-260.
"Kagasaurus" Hisa, 1988
Hauterivian, Early Cretaceous
Kuwajima Formation of the Itoshiro Subgroup of the Tetori Group, Japan
Material- (FPM 85050-1; Kaga-ryu) (~6 m) partial anterior tooth (>19.5
mm) (Manabe et al., 1989)
? tooth (Azuma, 1991)
Comments- The first tooth was discovered in 1985 and referred to Carnosauria.
This was illustrated and described in detail by Manabe et al. (1989), who assigned
it to Carnosauria fam. indet.. This was based on comparison to a supposedly
carnosaurian tooth (NSMP17178-17180) from Lufeng. Between 1985 and 1990 an additional
tooth was discovered, referred to Megalosauridae indet (Azuma, 1991). Hisa (1988)
referred to at least one of the teeth as "Kagasaurus", which is a
nomen nudum because it wasn't associated with a description. Manabe et al. state
FPM 85050-1 has the nickname Kaga-ryu, while Azuma calls both teeth Kaga-ryu.
Dong et al. (1990) regard the teeth as Megalosauridae indet.. Whether both teeth
are referrable to the same taxon is unknown, as the second has yet to be described.
FPM 85050-1 preserves on the the basal two-thirds of a tooth with a FABL 10.6
mm and a basal width of 5.6 mm. The lingual face is flat and the labial one
convex, indicating this is probably a premaxillary or anterior dentary tooth.
The mesial carina lacks serrations, while the distal carina has 17 serrations
per 5 mm. The serrations are rounded and angled slightly apically.
The flat lingual face is present in Saltriovenator, most abelisauroids, allosaurians and coelurosaurs (except Bicentenaria, compsognathid-grade maniraptoromorphs, Caudipteryx,
birds and some paravians with serrationless teeth). at least some
carnosaurs and most coelurosaurs (except those taxa which lack
carinae). Abelisaurids and allosaurs always have mesial serrations,
often extending to the base of the crown. Some non-maniraptoriform
coelurosaurs and dromaeosaurids have anterior teeth which lack only
mesial serrations, making "Kagasaurus" likely to be a member of one of
these groups.
References- Hisa, 1988. Utan Scientific Magazine. 4(24).
Manabe, Hasegawa and Azuma, 1989. Two new dinosaur footprints from the Early
Cretaceous Tetori Group of Japan. Gillette and Lockley (eds.). Dinosaur Tracks
and Traces. Cambridge University Press, Cambridge. 309-312.
Dong, Hasegawa and Azuma, 1990. The Age of Dinosaurs in Japan and China. Fukui,
Japan: Fukui Prefectural Museum. 65 pp.
Azuma, 1991. Early Cretaceous Dinosaur Fauna from the Tetori Group, central
Japan. Research on Dinosaurs from the Tetori Group (1). Professor S. Miura Memorial
Volume, 55-69.
"Vitakridrinda" Malkani, 2003
"V. sulaimani" Malkani, 2003
Maastrichtian, Late Cretaceous
Vitakri Formation, Pakistan
Material- ?(GSI/MSM-1049-16) distal caudal centrum (110 mm) (Malkani, 2020b)
(GSP/MSM-59-19; intended syntype of "Vitakridrinda sulaimani") proximal femur (~600-800 mm) (Malkani, 2004)
....(GSP/MSM-60-19; intended syntype of "Vitakridrinda sulaimani") proximal femur (Malkani, 2004)
Other diagnoses- Malkani (2006a) lists several characters for "Vitakridrinda"
in his diagnosis. Those involving the snout GSP/MSM-155-19 ("an rostrum") now refer to the baurusuchid Induszalim (here considered a junior synonym of Pabwehshi),
while those of the supposed braincase GSP/MSM-61-19 ("thick and long
basioccipital condyle articulated with posterior braincase") now apply
to a specimen referred by Malkani to his titanosaur "Gspsaurus".
Malkani's characters "laterally compressed tooth, having low crown and
extremely low ratio of crown height and rostro-caudal width" refer to a
block containing supposed teeth GSP/MSM-61-19 which do not appear to be
dinosaurian teeth regardless of what they actually are. The syntype
femora are diagnosed as having- "greater trochanter, eroded inturned
head with joint (only joint are found in both left and right proximal
femur), and hollow and thick peripheral bone of cross section". The
former is plesiomorphic for archosaurs, the second plesiomorphic for
dinosaurs, and the last typical of non-avialan theropods.
Another character listed by Malkani is "amphicoelous vertebrae", but
this is based on referred specimens (several of which he referred to
"Vitakrisaurus" in 2020b which seem to be sauropod) and is plesiomorphic for archosaurs.
Comments- Malkani's GSP specimen numbers are designed so that the last number is
the locality and the first is the specimen-specific number from that
locality, so are listed in order of locality first. Malkani first mentioned "Vitakridrinda" in his 2003 description of
"Brohisaurus", stating only "Vitakridrinda Sulaimani Abelisaurid Theropod (Malkani,2004a)." The only publication in the bibliography solely by Malkani is
his saurischian biodiversity paper, listed as in progress, but not
actually published until 2006. As the 2003 mention of "Vitakridrinda"
lacks a description or definition (ICZN Article 13.1.1; note the
reference to Malkani, 2004a doesn't count under 13.1.2 since it was not
published yet), was not indicated to be a new taxon (16.1), and did not
have a type specimen indicated (16.4), it was a nomen nudum at the
time. The first description of "Vitakridrinda" is generally claimed to
be the 2004 "Saurischian Dinosaurs from Late Cretaceous of Pakistan",
as referenced in Malkani 2006a and the Paleobiology Database. Yet ICZN
Article 9.9 lists "abstracts of articles, papers, posters, texts of
lectures, and similar material when issued primarily to participants at
meetings, symposia, colloquia or congresses" as not being published
work, so the paper doesn't count. In addition, it still violates 16.4
in not indicating a type specimen, saying only "One new genus and species of Abelisaurids Theropod dinosaur Vitakridrinda sulaimani
is diagnosed on the basis of rostrum, thick basioccipital condyle
articulated with posterior braincase; a pair of proximal femora with
its greater trochanter and partial head joint; and hollow and thick
peripheral bone of femoral cross section; and amphicoelous vertebrae."
Note while Malkani's "Saurischian
dinosaurs from the Late Cretaceous Pab Formation of Pakistan" has been
listed as being published in 2005 (Malkani, 2006a) or "2004d" and in
review (Malkani, 2006b), it was not published until 2015. Two 2006
publications of Malkani's almost describe
"Vitakridrinda" sufficiently, with 2006b being published in December,
while 2006a was published in April. However, a continual issue with
Malkani's taxa is that they almost always credit their authorship to
prior invalid publications instead of proposing them as new taxa, which
is necessary for all names after 1999- ICZN Article 16.1 is "Every new
name published after 1999, including new replacement names (nomina
nova), must be explicitly indicated as intentionally new." Thus until
Malkani publishes a description of "Vitakridrinda sulaimani" as gen et
sp. nov. instead of Malkani, 2006, and not in a conference abstract,
the name will never be valid.
The material was probably mostly discovered in 2001, as Malkani
discovered 2700 bones and bone fragments from the area at that time.
The femora GSP/MSM-59-19 and GSP/MSM-60-19 were found associated in one
mass and seemingly belong to one individual. The supposed braincase GSP/MSM-62-19 was found
100 meters away in the same horizon, while the snout GSP/MSM-155-19 and supposed tooth block GSP/MSM-61-19 were
found later (though early enough to be mentioned in 2004) and 50 meters
from the braincase. Malkani believed they all belong to the same
individual based on supposed lack of transportation and similar size,
but the area of separation is so large that this lacks merit. While
Malkani (2006a, b) referred to them all as the holotype, they may more properly be
syntypes, and their uncertain derivation from one animal would make
selection of a lectotype desirable. Indeed Malkani (2014a) states
"Previously the specimen GSP/MSM-155-19 is assigned to Vitakridrinda sulaimani (theropod dinosaur) but due to secondary palate nature it is being established as Induszalim bala -new genus and species of very large mesoeucrocodile" and "Its braincase and basioccipital can be assigned to Maojandino alami (because of the yellow brown matrix covered on supposed braincase is same as on associated vertebrae and limb elements of Maojandino alami)
if character supports after specimen preparation." Malkani (2019
online, 2020a) later referred the supposed braincase to his sauropod
taxon "Gspsaurus pakistani" instead, which he synonymized "Maojandino"
with.
Identification of material-
Given the designation of the snout as the holotype of a new taxon and
Malkani's referral of the supposed braincase to another taxon, this
leaves the supposed tooth section and femora as potential holotypes.
The latter are suggested here as the femora are definitely theropodan
while the tooth block is more controversial. Early photos of the
femora were either too small to be useful (e.g. Malkani, 2006a: fig.
5-8) or in a generally unhelpful distal cross section view (e.g.
Malkani, 2006a: fig. 14a). The thickness/diameter ratio of ~0.20 is
typical of non-avialan theropods. Malkani (2020b) finally published a
high resolution photo in medial view (fig. 1) which shows the shafts
are broken a few centimeters distal to the head and that the medial
portion of the head is broken off in both as well. The anterior
trochanter is proximally separate from the shaft unlike crocodyliforms,
sauropods and ankylosaurs. Contrary to the condition in abelisaurids (Carnotaurus, Ekrixinatosaurus, Rahiolisaurus, Tarascosaurus, Xenotarsosaurus)
the anterior trochanter is proximally placed so that it diverges from
the shaft at the midpoint of the femoral head. Among Late Cretaceous
Gondwanan dinosaurs this also differs from hadrosaurids, noasaurids and
carcharodontosaurids, but is similar to megaraptorans and
unenlagiines. While at least the right femur would seem more similar
to unenlagiines (e.g. Unenlagia comahuensis) than megaraptorans (e.g. Australovenator)
based on the anteroposteriorly narrow anterior trochanter, this is may
be due to the oblique perspective of Malkani's figure and the material
is referred to Coelurosauria incertae sedis. A more precise
phylogenetic position is not suggested here pending details visible in
additional perspectives such as greater trochanter anteroposterior
width, anterior trochanter sectional shape and presence of a posterior
trochanter. Its size is larger than most maniraptoromorphs but is
comparable to the unenlagiine Austroraptor.
Based on Malkani's (2006a) description, figure 14b of GSP/MSM-62-19
(mislabeled GSP/MSM-61-19 in early papers) is supposed to be an
anterior view of paroccipital processes and basipterygoid processes,
which is similar in rough outline to Abelisaurus (assuming the supposed basipterygoid
processes are basal tubera instead) except for the decurved and much
taller paroccipital processes. The latter features match titanosaur
braincases better, though in that case everything beneath the occipital
condyle area would be broken off. In either case, it's only the shape
that is similar, and since there are no obvious surface features or
natural edges, it could just as easily be part of an ilium, vertebra,
etc.. Figures 15a and b do have the rough shape of an occipital
condyle, but no apparent foramen magnum above it. The possibility it
could merely be the shape of the other side of the sediment nodule
should not be excluded, especially as the "anterior" view shows a gray
bone shape in the plane of a yellow nodule while the "posterior" and
"lateral" views show a yellow nodule shaped into what are supposed to
be bone structures. As noted above, Malkani (2014) refers this to his
sauropod "Maojandino alami" which he later synonymizes with another of
his sauropods "Gspsaurus pakistani" (Malkani, 2020a), but it is
provisionally placed as Titanosauria indet. here pending study of
Malkani's sauropod taxa.
Regarding the snout GSP/MSM-155-19, Wilson (pers. comm.,
2014) correctly observed the posterior view (fig. 2C in Malkani, 2010) is very
similar to the snout cross section of Pabwehshi. As noted above, Malkani (2014a) makes this snout the holotype of his new baurusuchid Induszalim bala, here provisionally retained as a junior synonym of Pabwehshi pakistanensis
as the supposedly distinct characters are due to misinterpreting
elements and the section being more anterior in position (see entry for
details).
The final supposed holotype specimen is block GSP/MSM-61-19 (mislabeled
GSP/MSM-62-19 in early papers), initially reported as including the
triangular coronal section of a tooth (e.g. Malkani, 2006a). Malkani
(2020) later included a drawing showing at least thirteen additional
much smaller structures labeled as teeth and another two labeled as
cranial bones. Malkani compares it to abelisaurids, but while the
first premaxillary tooth of Majungasaurus
is roughly similar in shape with a flat lingual side, it differs
greatly in having enamel of roughly equal thickness around the entire
tooth. The structure in GSP/MSM-61-19 instead has "walls" that
steadily decrease in thickness to tapered points "mesially" and
"distally" from a maximum thickness at the "labial" corner. This is
unlike any theropod tooth sections, so the Vitakri structure is
something else. The additional supposed teeth are at most a third the
size of the main supposed tooth and are of varying shapes, while the
only visible supposed cranial element (described as "eye peripheral
bone/lacrimal and other trirays star like bones") is equal in size to
some of these larger supposed teeth. Thus if the major structure were
a tooth, these would be far too small to be additional teeth or
circumorbital elements like a lacrimal or postorbital. Given the
preservation style in Vitakri taxa like Induszalim and Vitakrisaurus,
it is likely this block shows cross sections of often hollow cranial
and/or axial elements. It is tentatively referred to Archosauria
indet. here.
The paratype vertebrae are from other localities and do not overlap the
type material, so cannot be convincingly referred to the same taxon.
Four of them (GSP/MSM-53-2, 54-2, 55-2 and 57-3) were later referred to
Vitakrisaurus
by Malkani (2020) and are here placed in Titanosauria based on their
size and anteriorly placed neural arches (in the caudals).
GSP/MSM-56-1 was originally called a caudal vertebra (Malkani, 2006a)
and later an anterior dorsal (Malkani, 2020). The lack of a large
diapophysis suggests this may be an axis instead, tentatively assigned
to Titanosauria based on the broad, rounded and posteriorly sloped
neural spine. This would make Malkani's supposed posterior side
anterior, and note his labeled hyposphenes don't match the narrow
structure ventral to more horizontal postzygapophyseal surfaces seen in
abelisaurids. GSP/MSM-58-15 was identified as a caudal and later
(2020) as a proximal caudal, which seems correct. The size and
anteriorly placed neural arch resemble titanosaurs. Note the supposed
transverse process is a broken posterodorsal corner while the supposed
chevron facets are also broken surfaces. The material referred by
Malkani (2009) is far too fragmentary to be justified, including cross
sections of long bones, fragments of vertebral centra, and material
that Malkani himself lists as possibly non-fossil and coprolites. These
and most of the several additional fragmentary specimens referred by
Malkani (2020) are here placed in Archosauria indet.. One exception is
distal caudal centrum GSI/MSM-1049-16, which is elongate (elongation
index ~2.5 unlike crocodyliforms or large ornithischians), amphicoelous
and has "parallel longitudinal ridges/fibrous/laminar structures
alternated by long grooves trending antero-posteriorly located on all
sides except the anterior and posterior concave articular surfaces."
These latter may be the same longitudinal grooves and ridges present in
many paravians like unenlagiines, so could belong to "Vitakridrinda" if
the femoral similarities hold up. The supposed pes GSP/MSM-303-2 (here
identified as a series of vertebrae in section) was referred to Vitakridrinda by Malkani (2009), but
later made the holotype of supposed abelisauroid Vitakrisaurus by Malkani (2010b). The latter is placed in Archosauria incertae sedis here.
References-
Malkani, 2003. First Jurassic dinosaur fossils found from Kirthar range, Khuzdar
District, Balochistan, Pakistan. Geological Bulletin University of Peshawar.
36, 73-83.
Malkani, 2004. Saurischian dinosaurs from Late Cretaceous of Pakistan. in Hussain
and Akbar (eds.). Abstract volume of Fifth Pakistan Geological Congress, Islamabad,
Pakistan. 71-73.
Malkani, 2006a. Biodiversity of saurischian dinosaurs from the Latest Cretaceous
park of Pakistan. Journal of Applied and Emerging Sciences. 1(3), 108-140.
Malkani, 2006b. First rostrum of carnivorous Vitakridrinda (abelisaurid
theropod dinosaur) found from the latest Cretaceous Dinosaur Beds (Vitakri)
Member of Pab Formation, Alam Kali Kakor locality of Vitakri area, Barkhan District,
Balochistan, Pakistan. Sindh University Research Journal (Science Series). 38(2),
5-24.
Malkani, 2009. New Balochisaurus (Balochisauridae, Titanosauria, Sauropoda)
and Vitakridrinda (Theropoda) remains from Pakistan. Sindh University
Research Journal (Science Series). 41(2), 65-92.
Malkani, 2010a. Vitakridrinda (Vitakrisauridae, Theropoda) from the Latest
Cretaceous of Pakistan. Journal of Earth Science. 21(Special Issue 3), 204-212.
Malkani, 2010b. Stratigraphy and mineral potential of Sulaiman (Middle Indus)
basin, Pakistan. Sindh University Research Journal (Science Series). 42(2),
39-66.
Malkani, 2011. Vitakridrinda and Vitakrisaurus of Vitakrisauridae
Theropoda from Pakistan. Proceedings of the 6th Symposium of IGCP 507 on Paleoclimates
of the Cretaceous in Asia and their global correlation. Beijing, China. 59-66.
Malkani, 2014. Theropod dinosaurs and mesoeucrocodiles from the
terminal Cretaceous of Pakistan. The second International Symposium of
International Geoscience Programme (IGCP) Project 608, abstract volume.
169-172.
Malkani, 2015. Dinosaurs, mesoeucrocodiles, pterosaurs, new fauna and
flora from Pakistan. Geological Survey of Pakistan, Information
Release. 823, 1-32.
Malkani, 2019 online. Revision, discussion and diagnostic features of
valid titanosaurs (Sauropoda, Dinosauria) from Indo-Pakistan landmass.
[no longer uploaded] DOI: 10.13140/RG.2.2.25076.81287
Malkani, 2020a. First skull of medium sized titanosaur in Indo-Pakistan
subcontinent found from the Latest Maastrichtian Vitakri Formation of
Pakistan; Associated cranial and postcranial skeletons of Gspsaurus pakistani (Poripuchia, stocky Titanosauria, Sauropoda) from Pakistan and India. Open Journal of Geology. 9, 631-634.
Malkani, 2020b. Theropods, mesoeucrocodiles and pterosaurs found from
the latest Maastrichtian Vitakri Formation of Balochistan, Pakistan;
Description with large photographs and comparison with coeval taxa from
Indo-Pakistan subcontinent. Open Journal of Geology. 10, 510-551.
Xinjiangovenator Rauhut
and Xu, 2005
X. parvus Rauhut and Xu, 2005
Early Cretaceous
Lianmugin Formation of Tugulu Group, Xinjiang, China
Holotype- (IVPP V 4024-2) (2.5-4.2 m) tibia (312 mm including tarsal),
fibula, astragalus, calcaneum
Diagnosis- (after Rauhut and Xu, 2005) fibular condyle of tibia extending
farther posteriorly than lateral side of proximal end of this bone; fibula with
longitudinal groove on anterior side of proximal end.
Comments- The holotype was found in the same horizon (but a different
site) as Phaedrolosaurus, and was originally referred to it.
The tibia is not fused with the fibula and astragalocalcaneum, contra Dong (1973).
Phylogenetic relationships- Rauhut and Xu (2005) ran Xinjiangovenator
in an analysis that resulted with it being placed sister to Bagaraatan
inside Paraves. They assigned it to Maniraptora incertae sedis.
The broad ascending process is shared with megaraptorans plus
tyrannoraptorans, while the anterior transverse astragalar groove is
generally found in megaraptorans and basal tyrannosauroids within that
group, though rarely in maniraptoriforms as well.
References- Dong, 1973. Dinosaurs from Wuerho. In Reports of paleontological expedition to Sinkiang
(II), pterosaurian fauna from Wuerho, Sinkiang. Memoirs of the Institute of
Vertebrate Paleontology and Paleoanthropology Academia Sinica. 11, 45-52.
Rauhut and Xu, 2005. The small theropod dinosaurs Tugulusaurus and Phaedrolosaurus
from the Early Cretaceous of Xinjiang, China. Journal of Vertebrate Paleontology.
25(1), 107-118.
Megaraptora Benson, Carrano and Brusatte,
2009 online
Definition- (Megaraptor namunhuaiquii <- Baryonyx walkeri,
Chilantaisaurus tashuikouensis, Neovenator salerii, Carcharodontosaurus saharicus,
Allosaurus fragilis, Tyrannosaurus rex, Passer domesticus) (Novas, Agnolin,
Ezcurra, Porfiri and Canale, 2013)
Other definitions- (Megaraptor namunhuaiquii <- Chilantaisaurus
tashuikouensis, Neovenator salerii, Carcharodontosaurus saharicus, Allosaurus
fragilis) (Benson, Carrano and Brusatte, 2010)
Comments-
Benson et al.'s paper was originally released online on October 14 2009
but not officially published until January 2010. Since
Megaraptora is not a family-level clade however, it is not bound by the
rules of the ICZN and thus the 2009 date is here considered valid.
The taxa included in this clade have had a controversial history. Hucknull et
al. (2009) noted similarities between Australovenator, Fukuiraptor
and "Allosaurus" "robustus". I noted early on (online,
2008) that Aerosteon and Orkoraptor were extremely similar. Benson
et al. (2010) were the first to propose a relationship between all of these
taxa though, along with Megaraptor. They placed them in their new clade
Megaraptora, which they found to be sister to Chilantaisaurus and Neovenator
within Carcharodontosauridae (their Carcharodontosauria). Novas et al.
(2013) later reevaluated Benson et al.'s characters and used a different dataset
to place megaraptorans in Tyrannosauroidea, more derived than Dilong
and proceratosaurids, sister to Xiongguanlong
and tyrannosaurids. While Cau (2018) recovers megaraptorans in this
position too, only two steps move them to just outside Tyrannoraptora
as basal coelurosaurs. This compromise position is used here
pending more extensive future analyses.
Unfortunately, the original definition of Megaraptora assumes a carnosaurian
relationship, so does not include any coelurosaur external specifiers and would
be synonymous with Coelurosauria in my and Novas et al.'s (2013) phylogenies.
To avoid this, Novas et al. added Passer domesticus and Tyrannosaurus
rex to the definition.
References- Mortimer, online 2008. https://web.archive.org/web/20100822223105/http://scienceblogs.com/tetrapodzoology/2008/10/unhappy_with_aerosteon.php#comment-1144175
Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliot, 2009. New Mid-Cretaceous
(Latest Albian) dinosaurs from Winton, Queensland, Australia. PLoS ONE. 4(7),
e6190.
Benson, Carrano and Brusatte, 2010 (2009 online). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25. DOI: 10.4435/BSPI.2018.01
unnamed possible megaraptoran (O'Connor et al., 2006)
Albian, Early Cretaceous
TZ-07, Namba Member of the Galula Formation, Tanzania
Material- (TNM 03041) incomplete mid caudal vertebra (80 mm), partial mid caudal vertebra
Comments- Discovered between
2002 and 2005, this material was described as theropod by O'Connor et
al. (2006) and referred to the Unit I of the Red Sandstone Group at the
time. Roberts et al. (2010) subsequently revised the stratigraphic
nomenclature, naming Unit I the Galula Formation.
The caudals are amphicoelous, with a narrow ventral median groove, and
centra 153% taller than wide. Particularly interesting is that
"distinct cranial centrodiapophyseal and prezygodiapophyseal laminae
demarcate a substantial infraprezygapophyseal fossa on the neural
arch", which are rarely developed so well in such elongated theropod
caudals (centrum length 174% of height). Notably absent in
abelisaurids, spinosaurids and carcharodontosaurids, a similar
situation is present in Maip.
This could thus be the first recognized African megaraptoran, which
should be expected in the continent given their distribution in
Australia and South America.
References- O'Connor, Gottfried, Stevens, Roberts, Ngasala, Kapilima and
Chami, 2006. A new vertebrate fauna from the Cretaceous Red Sandstone Group,
Rukwa Rift Basin, southwestern Tanzania. Journal of African Earth Sciences.
44, 277-288.
Roberts, O'Connor, Stevens, Gottfried, Jinnah, Ngasala, Choh and
Armstrong, 2010 (online 2009). Sedimentology and depositional
environments of the Red Sandstone Group, Rukwa Rift Basin, southwestern
Tanzania: New insight into Cretaceous and Paleogene terrestrial
ecosystems and tectonics in sub-equatorial Africa. Journal of African
Earth Sciences. 57, 179-212.
Chilantaisaurus Hu,
1964
C. tashuikouensis Hu, 1964
Turonian, Late Cretaceous
Tashuikou, Ulanhsuhi Formation, Inner Mongolia, China
Lectotype- (IVPP V2884.1) humerus (580 mm)
Paralectotypes- ....(IVPP V2884.2) manual ungual II (250 mm straight, 260
mm along curve)
....(IVPP V2884.3) fragmentary ilium
....(IVPP V2884.4) femora (1.19 m)
....(IVPP V2884.5) tibiae (954 mm)
....(IVPP V2884.6) partial fibula
....(IVPP V2884.7) metatarsal II (415 mm), metatarsals III (460 mm), incomplete
metatarsals IV
Diagnosis- (after Benson and Xu, 2008) subrectangular, anteromedially
curving deltopectoral crest that protrudes almost as far anteriorly as it is
long proximodistally and bears a pitted scar on its anterior surface; obliquely
oriented ulnar condyle.
Other diagnoses- Hu's (1964)
original diagnosis was- "Humerus massive and elongate. Ungual strongly
curved. Fourth trochanter of femur less developed. Tibia shorter than
femur. Three metatarsals, short and not compactly united."
Comments- This was discovered in 1960. White et al. (2012) noted "morphological features [which] suggest the Chilantaisaurus
ungual belongs to the second digit." Hu (1964) believed three other
associated elements to be carnosaurian and
"most probably belong to the same species." Benson and Xu (2008)
referred the tooth IVPP V2884.8 to Theropoda indet. (here Averostra
indet.), the mid caudal vertebra IVPP V2884.X to be Sauropoda indet.
(originally in Rauhut, 2003) and a distal caudal centrum to be
Dinosauria indet. (here Eusaurischia indet.). Rauhut mentions a
third vertebra that "shows the same depressions underneath the
transverse process as found in 'C.' maortuensis [Shaochilong]", but Benson and Xu note this is actually catalogued as IVPP V2564.6, not V2884, and it is here provisionally referred to Shaochilong.
Relationships- Originally placed in Megalosauridae sensu lato by Hu (1964),
both Paul (1988) and Molnar et al. (1990) considered it part of a paraphyletic
Allosauridae more closely related to tyrannosaurids than Allosaurus and
Acrocanthosaurus based on its posteriorly reduced metatarsal III. Chilantaisaurus
was later identified as a megalosauroid by Chure (2000) and Rauhut (2003) based
on its straight humerus and elongate supposed manual ungual I. The latter author found
it to be the sister taxon of Spinosauridae based on the form of the tibial surface
that articulates with the astragalar ascending process, being a rounded medially
limited ridge in both Chilantaisaurus and Cristatusaurus. Benson
and Xu (2008) found that Chilantaisaurus had some characters suggestive
of avetheropod affinities (m. cuppedicus fossa; proximally wedge-shaped metatarsal
III), and shared a prominent ulnar epicondyle with allosauroids, and a weakly
hooked preacetabular process and reduced fourth trochanter with coelurosaurs.
Yet they also noted the anteriorly flat distal humerus and large humerofemoral
ratio are unlike allosauroids. They noted that Coelurus also has a rounded
medially limited ridge on its distal tibia, and that some avetheropods have
straight humeri and an elongate manual ungual I too. The low astragalar ascending
process is unlike coelurosaurs and most carnosaurs however. Most recently, Benson
et al. (2010) and Carrano et al. (2012) found it to be a megaraptoran, while
Novas et al. (2013) found it to be unresolved in Avetheropoda less derived than
Tyrannoraptora (including megaraptorans) + Compsognathidae, and Porfiri et al.
(2014) recovered it as the most basal coelurosaur.
When constrained in Carrano et al.'s matrix, it takes only one extra step to
place it in Carcharodontosauridae, showing either position is about as likely.
It takes 7 more steps to make it a spinosaurid, so this is less probable.
References- Hu, 1964. Carnosaurian remains from Alashan, Inner Mongolia.
Vertebrata PalAsiatica. 8, 42-63.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. Berkeley: University of California Press. 169-209.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. Ph.D. thesis. Columbia University. 964 pp.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 213 pp.
Benson and Xu, 2008. The anatomy and systematic position of the theropod dinosaur
Chilantaisaurus tashuikouensis Hu, 1964 from the Early Cretaceous of
Alanshan, People’s Republic of China. Geological Magazine. 145(6), 778-789.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
White, Cook, Hocknull, Sloan, Sinapius and Elliott, 2012. New forearm elements
discovered of holotype specimen Australovenator wintonensis from Winton,
Queensland, Australia. PLoS ONE. 7(6), e39364.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Porfiri, Novas, Clavo, Agnolin, Ezcurra and Cerda, 2014. Juvenile specimen of
Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation.
Cretaceous Research. 51, 35-55.
Siats Zanno and Makovicky, 2013
S. meekerorum Zanno and Makovicky, 2013
Cenomanian-Early Turonian, Late Cretaceous
Mussentuchit Member of Cedar Mountain Formation, Utah, US
Holotype- (FMNH PR 2716) (subadult; ~3.9 tons) teeth(?), possible cervical
fragments, fourth(?) dorsal centrum (167.2 mm), two partial anterior dorsal
neural arches, incomplete fifth dorsal vertebra (165.9 mm), thirteenth(?) dorsal
centrum (179.5 mm), several dorsal fragments, sacral(?) centrum, first caudal
neural arch, second caudal neural arch, third caudal neural arch, five distal
caudal vertebrae (~122.6, 134.3, ~121.4, 117.5, 111 mm), five fragmentary caudal
vertebrae, mid caudal chevron, partial ilium, proximal ischia, partial fibula,
phalanx II-1 (~170.4 mm), distal metatarsal III, phalanx III-2 (~136.3 mm),
phalanx IV-3 (60 mm), distal metatarsal II or IV
Paratype- (FMNH PR 3059) mid caudal neural arch, chevron, pedal phalanx,
fragments
Diagnosis- (after Zanno and Makovicky, 2013) anteroposteriorly expanded
centrodiapophyseal laminae yet lacking well developed infradiapophyseal fossae
on proximal caudals; anteroposterior elongation of anterior dorsal centra; abbreviated,
transversely broad neural spines on dorsal vertebrae (neural spine height ~50%
maximum height of centrum); transversely flattened, axially concave ventral
surface yielding subtriangular cross-section on distal caudal vertebrae; transversely
concave acetabular rim of iliac pubic peduncle; truncated lateral brevis shelf
with notched posterior end; brevis fossa with subparallel mediolateral margins;
supraacetabular crest truncated above midpoint of acetabular rim.
Comments- Zanno and Makovicky (2013) include this in a modified version
of Carrano et al.'s tetanurine matrix and find it to be a megaraptoran more
derived than Chilantaisaurus, in a polytomy with Fukuiraptor+Australovenator
and Megaraptor+Aerosteon. Porfiri et al. (2014) listed numerous
characters dissimilar to other megaraptorans, though they did not include it
in an analysis or propose a precise alternative reletionship, merely stating
"Siats lacks clear derived characters linking it with Megaraptora,
and even Coelurosauria." Resolution will require an analysis including
all of the proposed characters.
References- Zanno, Makovicky and Gates, 2012. A new giant carcharodontosaurian
allosauroid from the Lower Cretaceous Cedar Mountain Formation of Central Utah.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 199.
Zanno and Makovicky, 2013. Neovenatorid theropods are apex predators in the
Late Cretaceous of North America. Nature Communications. 4, 2827.
Porfiri, Novas, Clavo, Agnolin, Ezcurra and Cerda, 2014. Juvenile specimen of
Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation.
Cretaceous Research. 51, 35-55.
unnamed possible megaraptoran (Krumenacker and Scofield, 2017)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2251/49872) (juvenile) anterior dorsal centrum (49.9 mm)
Reference- Krumenacker, Simon, Scofield and Varricchio, 2017 (online 2016). Theropod
dinosaurs from the Albian-Cenomanian Wayan Formation of eastern Idaho. Historical
Biology. 29(2), 170-186.
Phuwiangvenator
Samathi, Chanthasit and Sander, 2019
P. yaemniyomi
Samathi, Chanthasit and Sander, 2019
Late Barremian, Early Cretaceous
Phu Wiang 9B, Sao Khua Formation, Thailand
Holotype-(SM-PW9B)
(~6 m) mid or posterior dorsal centrum, fused first to third sacral
centra (89, 73, 92 mm), phalanx I-1 (107 mm), proximal manual ungual I,
distal metacarpal II, proximal manual ungual II, phalanx III-1 (50 mm),
phalanx III-2, phalanx III-3 (49 mm), incomplete manual ungual III,
tibiae (615 mm; one partial), incomplete astragalus (94 mm wide),
calcaneum, metatarsal I, pedal ungual I, proximal metatarsal II,
phalanx II-1, phalanx II-2, incomplete pedal ungual II, proximal
metatarsal III, phalanx III-2, phalanx III-3, proximal pedal ungual
III, proximal metatarsal IV, phalanx IV-1, phalanx IV-4, pedal ungual
IV (68 mm)
Paratype- ....(SM-PW9A) atlantal intercentrum, incomplete astragalus (90.5 mm wide), calcaneum
Diagnosis- (after Samathi et
al., 2019) short sulci on the sacral vertebrae ventrally along the
anterior and posterior parts of the centra; anterior rim of metatarsal
IV slopes proximolaterally to distomedially.
Comments- The holotype was
discovered in 1993, with the referred specimens found later.
Samathi et al. write "The referred elements, found about 300 m away
from the holotype, seem to belong to
the same animal as the holotype, based on the size, matching
articulation (e.g., the right astragalocalcaneum fits perfectly to the
right tibia) and shared phylogenetic affinity (i.e., the right
astragalocalcaneum and the left astragalocalcaneum)." Samathi
(2016) first mentioned the holotype as "a new, as-yet-undescribed
theropod (PW9B) possibly a neovenatorid or a coelurosaur", and it is
one of two specimens noted by Samathi and Chanthasit (2017)
as "basal members of Megaraptora" using both Carrano et al.'s and Novas
et al.'s tetanurine analyses. Samathi et al. (2019) fully
described and named Phuwiangvenator as a new megaraptorian. They added it to Novas et al.'s analysis to find it as a megaraptoran basal to Fukuiraptor plus Megaraptoridae.
References- Samathi, 2016. Theropod dinosaurs from Thailand and southeast Asia: A
review with newly found specimens. Young Natural History Scientists
Meeting. [pp]
Samathi and Chanthasit, 2017. Two new basal Megaraptora
(Dinosauria: Theropoda) from the Early Cretaceous of Thailand with
comments on the phylogenetic position of Siamotyrannus and Datanglong. Journal of Vertebrate Paleontology. Program and Abstracts, 188.
Samathi, Chanthasit and Sander, 2019. Two new basal coelurosaurian
theropod dinosaurs from the Lower Cretaceous Sao Khua Formation of
Thailand. Acta Palaeontologica Polonica. 64(2), 239-260.
Aoniraptor Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento,
Brisson Egli and Novas, 2016
A. libertatum Motta, Aranciaga Rolando, Rozadilla, Agnolin,
Chimento, Brisson Egli and Novas, 2016
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of the Rio Limay Subgroup, Rio Negro, Argentina
Holotype- (MPCA-Pv 804/1-25)
(~6 m) incomplete last sacral vertebra, incomplete first caudal vertebra, second
caudal centrum, third caudal centrum, two proximal neural arches, five proximal-mid
caudal vertebrae, three partial proximal-mid caudal neural arches, five distal-mid
caudal vertebrae, distal caudal vertebra, three proximal chevrons, two mid chevrons
Diagnosis-
(after Motta et al., 2016) proximal-mid caudal
vertebrae with fan-shaped prezygapophyses lacking discernible articular surface;
blunt and thick process on lateral surface of proximal-mid caudal prezygapophyses;
distal-mid caudals with pair of non-articular flat surfaces on the posterodorsal
corner of the centrum.
Comments- Motta et al. (2016)
described this as a new taxon of non-megaraptorid megaraptoran,
proposed to be most closely related to Deltadromeus and
Bahariasaurus although lacking a phylogenetic analysis. Based on caudal similarities, many experts (e.g.
Cau, online 2016) synonymized it with the contemporaneous theropod
Gualicho described from the same formation two weeks earlier.
However, Aranciaga Rolando et al. (2020) argued Aoniraptor
is distinct from Gualicho
based on- "lateral pleurocoels in the centrum, pneumatic fossae related
to the transverse process or the base of the neural spine, or pneumatic
foramina within the pre- and postspinal fossae," "caudal centra with an
articular surface in contour, being dorsoventrally taller that
transversely wide (vs. the subcircular-shaped articular surface in Gualicho;
Figure 11b); not flared articular surfaces, resulting in flat lateral
surfaces of centrum (vs. flared articular surfaces with notably
longitudinally concave surfaces of centrum in Gualicho; Figure 11b); strongly transversely convex ventral surface of centrum (vs. flat in Gualicho), and absence of transversely sub-horizontal shelf connecting both postzygapophyses (vs. present and prominent in Gualicho;
Figure 11b″)", transverse processes placed within neural arch, mid
caudal prezygapophyses "short and strongly dorsally oriented" with
shorter articular surfaces, "postzygapophyses of mid-caudals are
notably reduced to a very small articular surface located in the
posterior corner of the neural arch", prespinal fossae "well-developed,
deep, notably longer than wide and sub-quadrangular in contour" and
"well-developed and deep" postspinal fossae. However, pleurocoels
are only present in Aoniraptor on its first caudal (not those preserved more distally in Gualicho),
the tranverse flaring of articular ends is not that different, and
differences in transverse ventral convexity cannot be confirmed with
published data. Despite these caveats the other differences are
quite real and Aranciaga Rolando et al. (2022) found that using Novas'
tetanurine analysis "17 extra steps are required to move Gualicho as the sister taxa to Aoniraptor", with the latter emerging as a megaraptoran more derived than Phuwiangovenator but outside the Murusraptor+Maip clade.
References-
Cau, online 2016. http://theropoda.blogspot.com/2016/07/nuovi-resti-di-aoniraptor-ehm-benvenuto.html
Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson Egli and Novas,
2016. New theropod fauna from the Upper Cretaceous (Huincul Formtation) of northwestern
Patagonia, Argentina. In Khosla and Lucas (eds.). Cretaceous period: Biotic
diversity and biogeography. New Mexico Museum of Natural History and Science
Bulletin. 71, 231-253.
Aranciaga Rolando, Marsa and Novas, 2020. Histology and pneumaticity of Aoniraptor libertatem (Dinosauria, Theropoda), an enigmatic mid-sized megaraptoran from Patagonia. Journal of Anatomy. 237(4), 741-756.
Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2022. A
large Megaraptoridae (Theropoda: Coelurosauria) from Upper Cretaceous
(Maastrichtian) of Patagonia, Argentina. Scientific Reports. 12:6318.
Megaraptora sensu Benson, Carrano and Brusatte, 2010 (online 2009)
Definition- (Megaraptor namunhuaiquii <- Chilantaisaurus tashuikouensis,
Neovenator salerii, Carcharodontosaurus saharicus, Allosaurus fragilis)
undescribed possible Megaraptora (Chokchaloemwong, Azuma, Shibata and Jintasakul, 2015)
Aptian, Early Cretaceous
Khok Kruat Formation, Thailand
Material- teeth
Comments- These were suggested to be close to Fukuiraptor.
Reference- Chokchaloemwong,
Azuma, Shibata and Jintasakul, 2015. The carcharodontosaurid teeth from
the Lower Cretaceous Khok Kruat Formation of Nakhon Ratchasima,
Thailand. 2nd International Symposium on Asian Dinosaurs. 32.
Fukuiraptor Azuma and
Currie, 2000
?= "Tsuchikurasaurus" dinosaur.net.cn, online 1998
F. kitadaniensis Azuma and Currie, 2000
Middle-Late Aptian, Early Cretaceous
Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Holotype- (FPDM 9712201-9712228) (~4.2 m) (subadult) maxillary fragment,
dentary fragment, premaxillary tooth (>17 mm), two maxillary teeth (26, 31.3mm),
dentary tooth (34 mm), dorsal centrum (77.5 mm), three proximal dorsal ribs,
distal caudal vertebra (26.7 mm), humerus (230 mm), ulna (211 mm), manual ungual
I (121 mm straight, 154 mm on curve), phalanx II-1 (64.9 mm), manual ungual
II (107.5 mm straight, 150 mm on curve), several manual phalanges, two pubic
fragments, two ischial fragments, femur (507 mm), proximal tibia, astragalus
(85.5 mm), metatarsal I (~70 mm), phalanx I-1 (67 mm), metatarsal II (297.5
mm), metatarsal III (297.5 mm), phalanx III-1 (99.2 mm), phalanx III-2 (77.4
mm), phalanx IV-2 (38 mm)
....(FPDM 96082443) humerus (242 mm)
....(FPDM 97080206) distal fibula
Paratypes- (FPDM 9712229) maxillary fragment, tooth
(FPDM 9712230) dentary fragment
(FPDM 9712231) tooth
(FPDM 9712232) tooth
(FPDM 9712233) dentary tooth (18.8 mm)
(FPDM 9712234) maxillary tooth (12.6 mm)
(FPDM 9712235) maxillary tooth (25 mm)
(FPDM 9712236) dentary tooth (18.5 mm)
(FPDM 9712237) tooth
(FPDM 9712238) tooth
(FPDM 9712239) tooth (>18 mm)
Referred- (FPDM-V96080810) maxillary tooth (50 mm) (Currie and Azuma,
2006)
(FPDM-V96081134) tooth (Currie and Azuma, 2006)
?(FPDM-V970730003) (~1.10 m) (juvenile) incomplete femur (Currie and Azuma,
2006)
(FPDM-V97080208) maxillary tooth (Currie and Azuma, 2006)
?(FPDM-V97080937) (~1.10 m) (juvenile) femur (Currie and Azuma, 2006)
?(FPDM-V9708102884) partial femur (Currie and Azuma, 2006)
(FPDM-V97081128) dentary tooth (33.4 mm)(Currie and Azuma, 2006)
(FPDM-V97081201) (~1.71 m) (juvenile) femur (196 mm) (Currie and Azuma, 2006)
(FPDM-V970813046) (~925 mm) (juvenile) femur (116.3 mm) (Currie and Azuma, 2006)
?(FPDM-V97081330) (~1.08 m) (juvenile) femur (134.9 mm) (Currie and Azuma, 2006)
(FPDM-V970821039) (~978 mm) (juvenile) femur (122.7 mm) (Currie and Azuma, 2006)
(FPDM-V97082330) maxillary tooth (17 mm) (Currie and Azuma, 2006)
(FPDM-V97082367) maxillary tooth (?23 mm) (Currie and Azuma, 2006)
?(FPDM-V97082553) humerus (Currie and Azuma, 2006)
(FPDM-V97082574) maxillary tooth (33 mm) (Currie and Azuma, 2006)
(FPDM-V97082728) maxillary tooth (>41 mm) (Currie and Azuma, 2006)
?(FPDM-V97120001) (~1.10 m) (juvenile) proximal femur (Currie and Azuma, 2006)
?(FPDM 9712240) fifth cervical centrum (58 mm) (Azuma and Currie, 2000)
....(FPDM 9712241) fifth cervical neural arch (Azuma and Currie, 2000)
?(FPDM 9712242) dorsal neural arch (Azuma and Currie, 2000)
?(FPDM 9712243) coracoid (58 mm deep) (Azuma and Currie, 2000)
(FPDM-V97122BNA3) (~1.65 m) (juvenile) femur (200 mm) (Currie and Azuma, 2006)
(FPDM-V97122BNA12) (~2.02 m) (juvenile) femur (244 mm) (Currie and Azuma, 2006)
(FPDM-V980721002) dentary tooth (18 mm) (Currie and Azuma, 2006)
(FPDM-V98072302) (~1.07 m) (juvenile) femur (134.2 mm) (Currie and Azuma, 2006)
(FPDM-V980724112) dentary tooth (Currie and Azuma, 2006)
(FPDM-V980801101) tooth (Currie and Azuma, 2006)
(FPDM-V980803001) premaxillary tooth (Currie and Azuma, 2006)
(FPDM-V980803120) maxillary tooth (>24 mm) (Currie and Azuma, 2006)
(FPDM-V980803123) tooth (Currie and Azuma, 2006)
(FPDM-V980804135) maxillary tooth (>17.6 mm) (Currie and Azuma, 2006)
(FPDM-V980804144) tooth (Currie and Azuma, 2006)
(FPDM-V980805018) (~735 mm) (juvenile) femur (92.2 mm) (Currie and Azuma, 2006)
(FPDM-V980805101) maxillary tooth (>33 mm) (Currie and Azuma, 2006)
(FPDM-V980806009) tooth (>27 mm) (Currie and Azuma, 2006)
(FPDM-V980810141) maxillary tooth (34 mm) (Currie and Azuma, 2006)
?(FPDM-V98081028) (~1.19 m) (juvenile) partial femur (Currie and Azuma, 2006)
(FPDM-V980813008) maxillary tooth (23 mm) (Currie and Azuma, 2006)
?(FPDM-V980813017) (~1.05 m) (juvenile) femur (Currie and Azuma, 2006)
(FPDM-V980815020) dentary tooth (>27.5 mm) (Currie and Azuma, 2006)
(FPDM-V980815176) dentary tooth (>25 mm) (Currie and Azuma, 2006)
(FPDM-V98081540) maxillary tooth (54.8 mm) (Currie and Azuma, 2006)
(FPDM-V980819055) maxillary tooth (>32 mm) (Currie and Azuma, 2006)
(FPDM-V980819173) tooth (Currie and Azuma, 2006)
(FPDM-V981200001) dentary tooth (>39 mm) (Currie and Azuma, 2006)
?(FPDM-V98120001) (~1.19 m) (juvenile) partial femur (Currie and Azuma, 2006)
?(FPDM-V98120002) (~1.42 m) (juvenile) partial femur (Currie and Azuma, 2006)
(FPDM-V981200012) dentary tooth (6 mm) (Currie and Azuma, 2006)
?(FPDM-V9812638) (~1.07 m) (juvenile) partial femur (Currie and Azuma, 2006)
?(FPDM-V99090901) (~1.02 m) (juvenile) distal femur (Currie and Azuma, 2006)
?(Tsuchikura-ryu) tooth (Azuma, 1991)
Diagnosis- (after Azuma and Currie, 2000) narrow dentary (~30% of depth)
(also in Eotyrannus); teeth with oblique blood grooves (also in Megalosaurus
and tyrannosaurids); ulnohumeral ratio >90%.
Other diagnoses- Of the other diagnostic characters listed by Azuma and
Currie (2000), fused interdental plates are also present in Fukuiraptor
and Eotyrannus. Larger hands with better developed unguals than Allosaurus
and a tall astragalar ascending process are primitive for coelurosaurs. The
supposedly broader than long pubic peduncle of the ilium is based on what is
probably an ornithopod pubis (see below).
Comments- The supposed ilium is more probably a Fukuisaurus pubis
(Jansma, pers. comm. 2004).
The first discovered element of Fukuiraptor may be a tooth from the type
quarry nicknamed Tsuchikura-ryu by Azuma (1991) and referred to Megalosauridae.
Currie and Azuma (2006) note 89% of the teeth from that quarry are from Fukuiraptor,
whose teeth do possess the generalized carnosaur/megalosaur morphology. While
several other Japanese nicknames have been inappropriately transformed into
nomina nuda in the published literature, "Tsuchikurasaurus" is so
far restricted to the internet, specifically due to the IVPP's dinosaur.net
site.
In 1991, jaw fragments were found in the quarry and identified as dromaeosaurid
based on their fused interdental plates. This was followed by the discovery
of a manual ungual I, astragalus and metatarsal III in 1993. Azuma and Currie
(1995) described these remains in an abstract as those of a giant dromaeosaurid,
which was associated in the paleontological community with the name "Kitadanisaurus"
through the late 1990's. Azuma and Currie (2000) later described the material
in more detail, along with more elements that made up a partial skeleton. Their
new taxon Fukuiraptor was identified as a basal carnosaur instead of
a dromaeosaurid, though dromaeosaurid material is known from the quarry (including
the original "Kitadanisaurus" tooth). Thus "Kitadanisaurus"
is not a synonym of Fukuiraptor, contra Olshevsky (DML 2000).
Most of the elements listed under 'holotype' were found associated in one small
area of the Kitadani quarry. The left humerus was given the separate call number
FPMN 96082443. The paratype maxillary fragment, dentary fragment, nine teeth,
cervical centrum and neural arch which fits it, dorsal neural arch and coracoid
were found in the same level, but in different areas of the quarry. They are
all the right size to belong to the holotype, but this can not be proven. At
least fourteen individuals are preserved in the type quarry, based on femoral
number. The more similar-sized pairs of femora possibly belong to single individuals
(99090901 and 980813017; 9812638 and 97080937; 970730003 and 97120001; 98081028
and 98120001). The provisionally referred femora are similar to Fukuiraptor
and not obviously dromaeosaurid. Several other specimens (humerus, manual phalanx
I-1, three manual unguals, three tibiae, pedal phalanx III-2) were found in
the quarry. Some are not referrable to Fukuiraptor (a straight manual
ungual and humerus), but others may be. Novas et al. (2013) suggested only the
more recurved teeth with mesial serrations limited to the apex were referrable
to Fukuiraptor, with the others being carcharodontosaurid. Yet there are intermediate
morphologies (e.g. FPDM-V97082728), so that Currie and Azuma's identification
of teeth with limited mesial serrations as anterior teeth (vs. posterior teeth
with extensive mesial serrations) seems likely.
Azuma and Currie (2000) found Fukuiraptor to be a basal carnosaur in
their phylogenetic analysis, as did Holtz (2001) and Holtz et al. (2004). Hucknull
et al. (2009) found it to be a non-carcharodontosaurid carnosaur more derived
than Sinraptor, while Benson (2010) placed it as the sister group of
Avetheropoda. Benson et al. (2010) added more characters and taxa to Benson's
earlier analysis and found Fukuiraptor was in their new clade Megaraptora
within Carcharodontosauridae, sister to Australovenator.
Longrich (2001) in placed it as a very basal coelurosaur, presaging
Novas et al. (2013) and derivatives that recover Megaraptora even
deeper within Coelurosauria as tyrannosauroids.
References- Azuma, 1991. Early Cretaceous Dinosaur Fauna from the Tetori
Group, central Japan. Research on Dinosaurs from the Tetori Group (1). Professor
S. Miura Memorial Volume, 55-69.
Azuma and Currie, 1995. A new giant dromaeosaurid from Japan. Journal of Vertebrate
Paleontology. 15(3), 17A.
dinosaur.net.cn, online 1998. https://web.archive.org/web/20031228215825/http://www.dinosaur.net.cn/museum/Tsuchikurasaurus.htm
Azuma and Currie, 2000. A new carnosaur (Dinosauria: Theropoda) from the Lower
Cretaceous of Japan. Canadian Journal of Earth Sciences. 37(12), 1735-1753.
Olshevsky, DML 2000. https://web.archive.org/web/20191030133242/http://dml.cmnh.org/2000Dec/msg00399.html
Holtz, 2001. Pedigree of the tyrant kings: New information on the origin and
evolution of the Tyrannosauridae. Journal of Vertebrate Paleontology. 21(3),
62A-63A.
Longrich, 2001. Secondarily flightless maniraptoran theropods? Journal of Vertebrate
Paleontology. 21(3), 74A.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska.
The Dinosauria Second Edition. University of California Press. 861 pp.
Currie and Azuma, 2006. New specimens, including a growth series of Fukuiraptor
(Dinosauria, Theropoda) from the Lower Cretaceous Kitadani Quarry of Japan.
Journal of the Paleontological Society of Korea. 22(1), 173-193.
Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliot, 2009. New Mid-Cretaceous
(Latest Albian) dinosaurs from Winton, Queensland, Australia. PLoS ONE. 4(7),
e6190.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. Zoological Journal of the Linnean
Society. 158(4), 882-935.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
F. sp. indet. (Molnar, Obata, Tanimoto and Matsukawa, 2009)
Barremian Formation, Early Cretaceous
Lower Member of the Sebeyashi Formation, Japan
Material- (NDC-P0001) lateral tooth (34.8 mm)
Comments- Molnar et al. (2009) referred this tooth to Fukuiraptor
aff. kitadaniensis.
Reference- Molnar, Obata, Tanimoto and Matsukawa, 2009. A tooth of Fukuiraptor
aff. F. kitadaniensis from the Lower Cretaceous Sebayashi Formation,
Sanchu Cretaceous, Japan. Bulletin of Tokyo Gakugei University, Division of
Natural Sciences. 61, 105-117.
F. sp. indet. (Chure, Manabe, Tanimoto and Tomida, 1999)
Late Cenomanian-Early Turonian, Late Cretaceous
Jobu Formation of Mifune Group, Japan
Material- (MDM 341) tooth (53 mm)
Comments- Originally referred to Carcharodontosauridae due to its enamel
wrinkles, these are shared by Fukuiraptor, which is similarly known from
Japan. Currie and Azuma (2006) found the width/FABL ratio and posterior serration
size matched Fukuiraptor more closely than carcharodontosaurids.
Reference- Chure, Manabe, Tanimoto and Tomida, 1999. An unusual theropod
tooth from the Mifune Group (Late Cenomanian to Early Turonian), Kumamoto, Japan.
in Tomida, Rich, and Vickers-Rich (eds.). Proceedings of the Second Gondwanan
Dinosaur Symposium. National Science Museum (Tokyo) Monographs. 15, 291-296.
Currie and Azuma, 2006. New specimens, including a growth series of Fukuiraptor
(Dinosauria, Theropoda) from the Lower Cretaceous Kitadani Quarry of
Japan. Journal of the Paleontological Society of Korea. 22(1), 173-193.
Megaraptoridae Novas, Agnolin, Ezcurra,
Porfiri and Canale, 2013
Diagnosis- (Megaraptor namunhuaiquii <- Fukuiraptor kitadaniensis,
Chilantaisaurus tashuikouensis, Baryonyx walkeri, Neovenator
salerii, Carcharodontosaurus saharicus, Allosaurus fragilis,
Tyrannosaurus rex, Passer domesticus) (Novas, Agnolin, Ezcurra,
Porfiri and Canale, 2013)
Reference- Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution
of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia.
Cretaceous Research. 45, 174-215.
unnamed megaraptorid (Martinez, Lamanna, Smith, Casal and Luna, 1999)
Middle Cenomanian-Turonian, Late Cretaceous
Lower Member of Bajo Barreal Formation, Chubut, Argentina
Material- ?(Ameghino coll.) distal caudal centrum (115 mm) (Huene, 1929)
(UNPSJB-PV 944) (subadult) ~third/fourth dorsal vertebra (~64 mm), two
dorsal ribs (one proximal), incomplete ~fifteenth caudal vertebra,
incomplete ~nineteenth caudal vertebra, incomplete ~twenty-fifth caudal
vertebra, proximal manual ungual I (~278 mm), partial phalanx II-2,
manual ungual II fragment, proximal manual ungual III, femoral
fragment, fibular fragment, distal metatarsal II, two fragmentary pedal
phalanges
(UNPSJB-PV 958) (~5 m; ~1 ton) partial proximal caudal vertebra, incomplete
mid caudal vertebra (64 mm), incomplete manual ungual I (~350 mm),
incomplete manual ungual III, fragmentary femur, partial tibia,
incomplete fibula, distal metatarsal I, incomplete metatarsal II (~400
mm), proximal phalanx II-1, phalanx II-2, incomplete phalanx III-2, two
fragmentary pedal phalanges, fragments
?(UNPSJB-PV 988) tooth (15.1x10.6x5.7 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
?(UNPSJB-PV 989) tooth (11.6x7.4x4.1 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
?(UNPSJB-PV 990) tooth (?x12.4x6.5 mm) (Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009)
Diagnosis- (after Lamanna et al., 2020) proximodorsal depression on manual ungual I.
Comments- UNPSJB-PV 944 was
found in 1989 and UNPSJB-PV 958 in 1998. They were first announced by
Martinez et al. (1999) as a theropod of uncertain affinities, with
manual ungual I of UNPSJB-PV 958 confused for pedal ungual II as was
the case for Megaraptor's holotype. The ungual was said to be similar to Megaraptor, but metatarsal II "much more massive than metatarsal III of Megaraptor, suggesting heavier construction of the pes." By 2004, Lamanna et al. had referred these specimens to Megaraptor in an SVP abstract, while Lamanna (2004) described them in detail in his thesis as Megaraptor
sp. indet.. Lamanna et al. (2020) puiblished a detailed description of
them as Megaraptoridae gen. et sp. indet., noting "comparisons of
elements that overlap between UNPSJB-PV 944 and UNPSJB-PV 958 (middle
caudal vertebrae, manual unguals I and III, metatarsal II) strongly
suggest that both specimens pertain to the same species" but that "due
to their highly fragmentary nature, we refrain from assigning UNPSJB-PV
944 and UNPSJBPV 958 to an existing megaraptorid genus or species or
erecting a new taxon to receive them." The dorsal vertebra of 944 was
reidentified as a third or fourth instead of first, and caudal
vertebrae of 958 were newly identified. Added to Novas' tetanurine
analysis, Lamanna et al. recovered the composite 944/958 OTU as a
megaraptorid basal to named taxa but more derived than cf. Rapator LRF 100-106.
Huene (1929) assigned a caudal centrum to an otherwise unknown group of
theropods, derived in having caudal pleurocoels. Mendez et al. (2012) noted
strong resemblences to their new Brazilian megaraptoran caudal MPMA 08-003-94,
showing Huene was correct.
Casal et al. (2009) describe three teeth as Dromaeosauridae, strongly
recurved and lacking mesial serrations. Ibiricu et al. (2020) later
referred these to Megaraptora indet..
References- Huene, 1929. Los Saurisquios y Ornitisquios del Cret�ceo
Argentino. Anales del Museo de La Plata (Serie 2). 3, 1-194.
Martinez, Lamanna, Smith, Casal and Luna, 1999. New Cretaceous
theropod material from Patagonia. Journal of Vertebrate Paleontology. 19(3),
62A.
Lamanna, Mart�nez, Luna, Casal, Ibiricu and Ivany, 2004. Specimens of
the problematic large theropod dinosaur Megaraptor from the Late Cretaceous
of central Patagonia. Journal of Vertebrate Paleontology. 24(3), 252A.
Lamanna, 2004. Late Cretaceous dinosaurs and crocodiliforms from Egypt and Argentina.
PhD Thesis. University of Pennsylvania. 305 pp.
Casal, Candeiro, Martinez, Ivany and Ibiricu, 2009. Theropod teeth
(Dinosauria: Saurischia) from the Bajo Barreal Formation, Upper
Cretaceous, Chubut Province, Argentina. Geobios. 42, 553-560.
Mendez, Novas and Iori, 2012. First record of Megaraptora (Theropoda, Neovenatoridae)
from Brazil. Comptes Rendus Palevol. 11, 251-256.
Ibiricu, Casal, Martinez, Alvarez and Poropat, 2020 (online 2019). New
materials and an overview of Cretaceous vertebrates from the Chubut
Group of the Golfo San Jorge Basin, central Patagonia, Argentina.
Journal of South American Earth Sciences. 98, 102460.
Lamanna, Casal, Martinez and Ibiricu, 2020. Megaraptorid (Theropoda:
Tetanurae) partial skeletons from the Upper Cretaceous Bajo Barreal
Formation of central Patagonia, argentina: Implications for the
evolution of large body size in Gondwanan megaraptorans. Annals of
Carnegie Museum. 86(3), 255-294.
unnamed possible megaraptorid (Paulina-Carabajal and Coria, 2015)
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of Rio Neuquen Subgroup, Neuquen, Argentina
Material- (MCF-PVPH 320) frontal (Paulina-Carabajal and Coria, 2015)
Comments- MCF-PVPH 320 was described by Paulina-Carabajal and Coria (2015) as
Allosauroidea gen. et sp. indet. and emerged as a metriacanthosaurid
when entered into Cau's Sauroniops
matrix. Paulina-Carabajal and Currie (2017) stated it "shares several features with Murusraptor, suggesting that it is also a megaraptorid, although it appears to be a different taxon than Megaraptor."
"These characters include a wide supratemporal fossa that covers more
than the 50% of the length of the frontal (as in tyrannosaurids), the
presence of a shallow pit and an alar projection on the dorsal surface
of the postorbital process of the frontal, a short orbital vault, the
presence of a triangular wedge of the parietals separating the frontals
posteriorly (in Murusraptor the wedge is anteroventral, whereas in MCF-PVPH the wedge is anterodorsal, as in Piatnitzkysaurus),
a short and robust olfactory tract impression, and the greater relative
size of the olfactory bulb impression." Note if this is a megaraptorid
some material currently assigned to Megaraptor may belong to it.
References- Paulina-Carabajal and Coria, 2015. An unusual theropod frontal
from the Upper Cretaceous of north Patagonia. Alcheringa. 39(4), 514-518.
Paulina-Carabajal and Currie, 2017. The braincase of the theropod dinosaur Murusraptor:
Osteology, neuroanatomy and comments on the paleobiological
implications of certain endocranial features. Ameghiniana. 54, 617-640.
Megaraptoridae indet. (Casal, Ibiricu, Martinez, Luna, Gonzalez
Svoboda and Ivany, 2015)
Coniacian-Maastrichtian, Late Cretaceous
Lago Colhue Huapi Formation, Chubut, Argentina
Material- (UNPSJB-PV 1028) incomplete manual ungual II (Casal, Mart�nez, Luna and Ibiricu, 2016)
(UNPSJB-PV 1046) manual ungual I (240 mm) (Casal, Mart�nez, Luna and Ibiricu, 2016)
....(UNPSJB-PV 1066) incomplete metatarsal III (Casal, Mart�nez, Luna and Ibiricu, 2016)
(UNPSJB-PV 1102) manual ungual I (~225 mm) (Ibiricu, Casal, Martinez, Alvarez and Poropat, 2020)
(UNPSJB-PV 1104) (medium-sized; juvenile) maxilla, partial braincase,
mandibles, dorsal ribs, chevron, scapula, humerus, radius, ulna,
ungual, femur, distal tibia, metatarsus, pedal phalanges, elements (Casal, Ibiricu, Alvarez, Luna and Martinez, 2019)
Comments- Initially, Casal et al. (2015) reported (translated) "theropods are rare, represented so far only by megaraptorids." Casal et al. (2016) figured UNPSJB-PV 1028 and 1046, mentioning 1066 as (translated) "a
possible metatarsal III also of this taxon", incorrectly calling these
'megaraptoriforms' when there is no clade 'Megaraptoriformes'. This
material and additional ungual UNPSJB-PV 1102 was
described in detail by Ibiricu et al. (2020) as Megaraptoridae indet..
Note Lamanna et al. (2019) incorrectly cite "an incomplete but
associated skeleton consisting of a tooth, a manual ungual I, and a
hind limb element, either a tibia or a metatarsal III (UNPSJB-PV 1066;
Casal et al. 2016, 2018; GAC pers. obs. 2018)" as separate from UNPSJB-PV 1046, perhaps confused with abelisauroid specimen UNPSJB-PV 1067 also including a tooth and tibia.
Casal et al. (2019) announced a new skeleton (translated) "assigned
to the clade Megaraptoridae due to the presence of the
anteroposteriorly expanded and transversely compressed olecranon
process in the ulna, and the ungual phalanx with a developed ventral
keel that connects the flexor tubercle." Lamanna et al. (2020)
revealed the specimen number.
References- Casal, Ibiricu, Martinez, Luna, Gonzalez Svoboda and Ivany, 2015.
El registro fosil de la Formacion Lago Colhue Huapi (Coniaciano-Maastrichtiano),
Grupo Chubut, Argentina. XXIX Jornadas Argentinas de Paleontolog�a de
Vertebrados, resumenes. Ameghiniana. 52(4) suplemento, 10-11.
Casal, Mart�nez, Luna and Ibiricu, 2016. Ordenamiento y caracterizaci�n
faun�stica del Cret�cico Superior del Grupo Chubut, Cuenca del Golfo
San Jorge, Argentina. Revista Brasileira de Paleontologia. 19, 53-70.
Casal, Ibiricu, Alvarez, Luna and Martinez, 2019. Nuevos materiales de
Megaraptoridae del Maastrichtiano de la Formaci�n Lago Colhu� Huapi,
Grupo Chubut, Patagonia Argentina. Reuni�n de Comunicaciones de la
Asociaci�n Paleontol�gica Argentina 2019. R69.
Lamanna, Casal, Ibiricu and Mart�nez, 2019. A new peirosaurid
crocodyliform from the Upper Cretaceous Lago Colhu� Huapi Formation of
central Patagonia, Argentina. Annals of Carnegie Museum. 85, 193-211.
Ibiricu, Casal, Martinez, Alvarez and Poropat, 2020 (online 2019). New
materials and an overview of Cretaceous vertebrates from the Chubut
Group of the Golfo San Jorge Basin, central Patagonia, Argentina.
Journal of South American Earth Sciences. 98, 102460.
Lamanna, Casal, Martinez and Ibiricu, 2020. Megaraptorid (Theropoda:
Tetanurae) partial skeletons from the Upper Cretaceous Bajo Barreal
Formation of central Patagonia, argentina: Implications for the
evolution of large body size in Gondwanan megaraptorans. Annals of
Carnegie Museum. 86(3), 255-294.
undescribed Megaraptoridae (Coria and Arcucci, 2004)
Campanian, Late Cretaceous
Anacleto Formation, Rio Colorado Subgroup, Mendoza, Argentina
(MPCM-PV 3109-3117, 5120-5123) ulna, manual ungual I, manual ungual II,
three manual phalanges, pedal ungual I, metatarsals II, metatarsals
III, phalanx IV-1, pedal ungual IV (Novas, Agnolin, Ezcurra, Porfiri
and Canale, 2013)
Campanian, Late Cretaceous
Anacleto Formation, Rio Colorado Subgroup, Neuquen, Argentina
Material- (MCF-PVPH-416) pubic fragment (Coria and Arcucci, 2004)
Comments- Coria and Arcucci
(2004) described MCF-PVPH-416 as Theropoda indet., suggesting it was a
basal tetanurine with similarities to Giganotosaurus.
Baiano and Coria (2018) reanalyzed the material, finding "in light of
current knowledge, it would be possible to fine-tune the identification
of these elements as Megaraptora indet."
Novas et al. (2013) stated "some associated metatarsals and phalanges
were found, including ... a single right manual ungual of the first
digit (MCNA-PV-3112)..." Mendez et al. (2019) briefly described the
material in an abstract, noting "this ulna is more robust and the
olecranon process is less posteriorly projected than that of Australovenator and Megaraptor" and "Mt-III exhibits a marked extensor fossa, which is deeper than in the Mt-III of Australovenator." Size-wise, "metatarsals are 25% longer than those of Australovenator."
References- Coria and Arcucci,
2004. Nuevos dinosaurios ter�podos de Auca Mahuevo, provincia del
Neuqu�n (Cretácico tard�o, Argentina). Ameghiniana. 41, 597-603.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Baiano and Coria, 2018. Revisiting theropod material from the Late
Cretaceous nesting site Auca Mahuevo and the possible record of a giant
megaraptoran. XXXII Jornadas Argentinas de Paleontolog�a de Vertebrados
y VII Jornadas T�cnicas de Paleontolog�a de Vertebrados, Libro de
Res�menes). R7.
M�ndez, Gianechini, Canale and D�az-Mart�nez, 2019. A new megaraptorid
specimen (Theropoda, Coelurosauria) from Ca�ad�n Amarillo (Anacleto
Formation, Campanian,
Upper Cretaceous), Mendoza Province, Argentina. 33.as Jornadas Argentinas de Paleontolog�a de Vertebrados. 60-61.
unnamed Megaraptoridae (Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Briss�n-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, �lvarez-Herrera, Gentil, Bogan, Chimento,
Garc�a-Mars�, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019)
Late Campanian-Early Maastrichtian, Late Cretaceous
Chorrillo Formation, Santa Cruz, Argentina
Material- (MACN-Pv 19066) tooth (19.0x10.7x6.8 mm)
?(MPM 21546) (~3 m, juvenile?) posterior dorsal centrum (27.2 mm)
(MPM-PV-22864) four teeth (Moyano-Paz et al., 2022)
(MPM-PV-22865) nine teeth (Moyano-Paz et al., 2022)
Comments- Discovered between
January and March 2019, Novas et al. assign MPM 21546 to
Megaraptoridae based on paired dorsal pleurocoels separated by a
septum. However, Aranciaga Rolando et al. (2022) say "comparisons
of the isolated dorsal centrum (MPM 21,546) with unenlagiids, such as Unenlagia,
better suggests that this bone might pertain to Unenlagiidae rather
than to Megaraptoridae." MACN-Pv 19066 was found in 1981 and was
said to
resemble megaraptorids in lacking mesial serrations, but differs from
named taxa in lacking an 8-shaped basal section and having more dense
serrations (5/mm).
MPM-PV-22864 and 22865 were discovered in March 2020 and referred to Megaraptoridae by Moyano-Paz et al. (2022), closer to Megaraptor than Australovenator based on the lack of mesial serrations.
References- Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Briss�n-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, �lvarez-Herrera, Gentil, Bogan, Chimento,
Garc�a-Mars�, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019. Paleontological discoveries in the Chorrillo
Formation (upper Campanian-lower Maastrichtian, Upper Cretaceous),
Santa Cruz Province, Patagonia, Argentina. Revista del Museo Argentino
de Ciencias Naturales. 21(2), 217-293.
Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2022. A
large Megaraptoridae (Theropoda: Coelurosauria) from Upper Cretaceous
(Maastrichtian) of Patagonia, Argentina. Scientific Reports. 12:6318.
Moyano-Paz, Rozadilla, Agnolin, Vera, Coronel, Varela, Gomez-Dacal,
Aranciaga-Rolando, D'Angelo, Perez-Loinaze, Richiano, Chimento, Motta,
Sterli, Manabe, Tsuihiji, Isasi, Poire and Novas, 2022. The uppermost
Cretaceous continental deposits at the southern end of Patagonia, the
Chorrillo Formation case study (Austral-Magallanes Basin):
Sedimentology, fossil content and regional implications. Cretaceous
Research. 130, 105059.
unnamed megaraptorid (Frey and Martill, 1995)
Albian, Early Cretaceous
Romualdo Formation of Santana Group, Brazil
Material- (SMNS 58023) (juvenile) third sacral vertebra (63.2 mm), fourth
sacral vertebra (62.5 mm), fifth sacral vertebra (71.7 mm), first caudal vertebra, ilial fragment
Comments- Frey and Martill
(19956) described this as a possible 'oviraptorosaurid'
(oviraptorosaurian), and while this was doubted by several later
authors, Aranciaga Rolando et al. (2018) were the first to reidentify
the taxon as a megaraptoran using Carrano's and Novas's tetanurine
analyses.
Reference- Frey and Martill, 1995. A possible oviraptorosaurid theropod
from the Santana Formation (Lower Cretaceous, Albian?) of Brazil. Neues Jahrbuch
Fur Geologie und Palaeontologie. 7, 397-412.
Aranciaga Rolando, Brisson Egli, Sales, Martinelli, Canale and Ezcurra,
2018 (online 2017). A supposed Gondwanan oviraptorosaur from the Albian of Brazil
represents the oldest South American megaraptoran. Cretaceous Research.
84, 107-119.
unnamed megaraptorid (Sales, Martinelli, Francischini, Rubert, Marconato, Soares and Schultz, 2018)
Santonian-Maastrichtian, Late Cretaceous
Morro do Cambambe, Bauru or Parecis Group, Brazil
Material- (UFRGS-PV-032-K) mid caudal centrum (61 mm)
Comments- This was discovered in the 1990s. Sales et al. (2017) identified it as a megaraptoran which "closely resembles that of Aerosteon instead of other megaraptorans."
Reference- Sales, Martinelli,
Francischini, Rubert, Marconato, Soares and Schultz, 2018 (online 2017). New dinosaur
remains and the tetrapod fauna from the Upper Cretaceous of Mato Grosso
State, central Brazil. Historical Biology. 30(5), 661-676.
unnamed megaraptorid (Souza, Kuhn and Hirooka, 2011)
Santonian-Maastrichtian, Late Cretaceous
Cachoeira do Bom Jardim Formation, Brazil
Material- (CD-CRP-127) dorsal centrum
Comments- Souza et al. (2011) figured this as as Theropoda indet., but Sales et al. (2017) referred it to Megaraptora.
References- Souza, Kuhn and
Hirooka, 2011. Saur�podes e ter�podes da Chapada dos Guimar�es, Mato
Grosso, Brasil. In Carvalho, Srivastava, Stroschschoen and Lana (eds.).
Paleontologia: Cen�rios de Vida. 4th ed. Interci�ncia. 655-662.
Sales, Martinelli, Francischini, Rubert, Marconato, Soares and Schultz,
2018 (online 2017). New dinosaur remains and the tetrapod fauna from the Upper
Cretaceous of Mato Grosso State, central Brazil. Historical Biology.
30(5), 661-676.
unnamed megaraptorid (Martinelli, Borges Ribeiro, M�ndez, Neto, Cavellani, Felix, da Fonseca Ferraz and Antunes Teixeira, 2013)
Campanian?, Late Cretaceous
Uberaba Formation, Bauru Group, Brazil
Material- (CPPLIP 1324) mid caudal centrum (80 mm)
Comments- This was found in 2011. Martinelli et al. (2013) refer it to Megaraptora and note a greater similarity to Aerosteon than Sao Jose do Rio Preto Formation centrum MPMA 08-003-94.
Reference- Martinelli, Borges
Ribeiro, M�ndez, Neto, Cavellani, Felix, da Fonseca Ferraz and Antunes
Teixeira, 2013. Insight on the theropod fauna from the Uberaba
Formation (Bauru Group), Minas Gerais State: New megaraptoran specimen
from the Late Cretaceous of Brazil. Rivista Italiana di Paleontologia e
Stratigrafia. 119, 205-214.
unnamed megaraptorid (Mendez, Novas and Iori, 2012)
Maastrichtian, Late Cretaceous
Sao Jose do Rio Preto Formation, Brazil
Material- (MPMA 08-003-94) distal caudal centrum (118 mm)
Reference- Mendez, Novas and Iori, 2012. First record of Megaraptora
(Theropoda, Neovenatoridae) from Brazil. Comptes Rendus Palevol. 11, 251-256.
unnamed Megaraptoridae (Rich, 1998)
Late Aptian-Early Albian, Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P186076) ulna (192.6 mm) (Rich, 1998)
(NMV P221081) (juvenile) partial ~sixth cervical vertebra
(37 mm, excluding anterior ball) (Barrett, Benson, Rich and Vickers-Rich,
2010)
(NMV P239459) tooth (11.5x6.2x3.8 mm) (Poropat, White, Vickers-Rich and Rich, 2019)
(NMV P239464) manual ungual I (166 mm, 201 mm on curve) (Poropat, White, Vickers-Rich and Rich, 2019)
(NMV P252264) tooth (15.8x8.1x~4 mm) (Poropat, White, Vickers-Rich and Rich, 2019)
(NMV P252715A) manual ungual III (60.5 mm, 73 mm on curve) (Poropat, White, Vickers-Rich and Rich, 2019)
(NMV P253701) (juvenile?) astragalus (41.6 mm trans) (Poropat, White, Vickers-Rich and Rich, 2019)
Comments- Pigdon (DML 1998) noted a supposedly dromaeosaurid ulna had
been reported in the 1998 Flat Rocks Site Report, mentioned by Rich and Rich.
Pidgon states (pers comm., 2007) that Rich and Rich had forgotten about suggesting
a dromaeosaurid identity and that it may be the ulna photographed in Rich and
Vickers-Rich (2003) and identified merely as theropod. Salisbury et al. (2007)
stated it compared favorably to Megaraptor, which was expanded on in
Smith et al.'s (2008) description and phylogenetic analysis. They referred it
to cf. Megaraptor based on two characters- proximocaudally expanded blade-like
olecranon process that extends distally as a caudal olecranon crest; pronounced
lateral tuberosity that is continuous distally with a distinct lateral crest.
Benson et al. (2010) claimed they were potentially more widespread within megaraptorans,
referring the ulna to Neovenatoridae indet.. The proximocaudally expanding olecranon
is indeed shared with Australovenator (but not Fukuiraptor), and
the lateral tuberosity is even larger in Australovenator (unpreserved
in Fukuiraptor). However, even if the blade-like olecranon is due to
crushing as they say, the posterior olecranon crest is absent in Australovenator
and Fukuiraptor, and the lateral crest emerging from the lateral tuberosity
is absent in Australovenator at least. Thus pending further study, NMV
V186076 is closer to Megaraptor than to other megaraptorans known from
ulnae, but how it compares to Aerosteon or Orkoraptor is unknown.
Although referred to Spinosauridae in its original description, Novas
et al. (2013) questioned this and only retained NMV P221081 as
Averostra or Tetanurae indet.. Poropat et al. (2019) referred it
to
?Megaraptoridae gen. et sp. indet. based on "the presence of two
pneumatic foramina on the left side (centrum + parapophysis)."
Discovered between 2013 and 2017, NMV P239459, P239464, P252264,
P252715A and P253701 were "found isolated, and all were sourced from a
fluvial deposit that also contains isolated bones pertaining to other
vertebrate groups" and so "no two specimens can be confidently
attributed to a single theropod individual." All were assigned to
Megaraptoridae cf. Australovenator wintonensis, but that species from the slightly later Winton Formation was not stated to be more similar in any characters than e.g. Megaraptor.
References- Pigdon, DML 1998. https://web.archive.org/web/20201110054703/http://dml.cmnh.org/1998Sep/msg00454.html
Rich, 1998. Research Update. Dinosaur Dreaming 1998 Annual
Report. Monash University. [pp]
Rich and Vickers-Rich, 2003. Protoceratopsian? ulnae from the Early Cretaceous
of Australia. Records of the Queen Victoria Museum. 113, 12 pp.
Salisbury, Agnolin, Ezcurra and Pais, 2007. A critical reassessment of the Cretaceous
non-avian dinosaur faunas of Australia and New Zealand. Journal of Vertebrate
Paleontology. 27(3), 138A.
Smith, Makovicky, Agnolin, Ezcurra and Salisbury, 2008. A Megaraptor-like
theropod (Dinosauria: Tetanurae) from Australia; Support for faunal exchange
between eastern and western Gondwana in the Mid-Cretaceous. Journal of Vertebrate
Paleontology. 28(3), 145A.
Smith, Makovicky, Agnolin, Ezcurra, Pais and Salisbury, 2008. A Megaraptor-like
theropod (Dinosauria: Tetanurae) in Australia: Support for faunal
exchange across eastern and western Gondwana in the Mid-Cretaceous.
Proceedings of the Royal Society B. 275(1647), 2085-2093.
Barrett, Benson, Rich and Vickers-Rich, 2010. A definitive
spinosaurid theropod from the Lower Cretaceous of Australia and its implications
for Gondwanan paleobiogeography. Journal of Vertebrate Paleontology. Program
and Abstracts 2010, 57A.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Barrett, Benson, Rich and Vickers-Rich, 2011. First spinosaurid dinosaur from
Australia and the cosmopolitanism of Cretaceous dinosaur faunas. Biology Letters.
7(6), 933-936.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from southern Australia
indicates high polar diversity and climate-driven dinosaur provinciality. PLoS
ONE. 7(5), e37122.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Poropat, White, Vickers-Rich and Rich, 2019. New megaraptorid
(Dinosauria: Theropoda) remains from the Lower Cretaceous Eumeralla
Formation of Cape Otway, Victoria, Australia. Journal of Vertebrate
Paleontology. 39, e1666273.
"Allosaurus" "robustus"
Chure, 2000 vide Glut, 2003
Early Aptian, Early Cretaceous
Wonthoggi Formation of Strzelecki Group, Victoria, Australia
Material- (NMV P150070; Allosaurus "robustus"; Cape Peterson allosaur) (~4.8 m) astragalus (~108 mm wide) (Molnar, Flannery and Rich, 1981)
?(NMV P186153) (~8-9 m) partial manual ungual ?I (Benson, Rich, Vickers-Rich and Hall, 2012)
?(NMV P208096) mid caudal centrum (36 mm) (Benson, Rich, Vickers-Rich and Hall, 2012)
Diagnosis- (suggested) astragalar ascending process tall as in other
coelurosaurs; astragalar ascending process is primitively restricted to the
lateral body compared to Fukuiraptor, Australovenator and other
coelurosaurs except Coelurus. Differs from Coelurus in plesiomorphically
having a bulbous medial condyle in anterior view.
Other diagnoses- Of the characters listed by Molnar et al. (1980), the
astragalus does not seem more robust than Fukuiraptor. The absence of
a pit on the posterior base of the astragalar ascending process is primitive.
Many other taxa such as Torvosaurus, Sinraptor, Fukuiraptor,
Australovenator, Coelurus and Appalachiosaurus have the
vertical groove on the posterior face of the ascending process.
Comments- Chure (2000) is the first person to publish the name Allosaurus
"robustus", previously confined to a museum label. Names in theses
aren't usually listed in this website, and this one is only because it was later
published by Glut (2003). Glut's work includes a caveat to the effect that it
is not available to establish new taxonomy however, so the name remains unofficial.
Molnar et al. (1981) initially described this specimen as Allosaurus
sp., which was disputed by Welles (1983) who argued it resembled 'ornithomimoid'
tarsi more. Molnar et al. (1983) countered Welles, though no other supposed
allosaurid genera were compared in either of Molnar et al.'s works. In addition,
Welles' 'ornithomimoid' type refers to a grade of tarsus encompassing Maniraptoriformes,
and not ornithomimosaurs in particular. Chure (1998) again disputed an allosaurid
relationship in an abstract, though he acknowledged it could be allosauroid.
This was elaborated on in Chure's (2000) thesis, where he assigns it to Avetheropoda,
but not Allosauridae. The resolution of "robustus"' identity occured
when Azuma and Currie (2000) described their new supposed basal carnosaur Fukuiraptor,
which has an extremely similar astragalus. Subsequently, Hocknull et al. (2009)
found their new supposed basal carcharodontosaurid Australovenator was
extremely similar to Fukuiraptor and "robustus" as well, referring
the latter to Australovenator sp.. Benson et al. (2010) agreed these
three taxa were closely related, creating the clade Megaraptora for them and
viewing "robustus" as an indeterminate member. Comparing all three
taxa, the ascending process reaches further laterally and angles more laterally
in Fukuiraptor and "robustus" than in Australovenator.
The ascending process is pointed and 20% taller in Fukuiraptor than Australovenator
or "robustus". It reaches further medially in Fukuiraptor and
Australovenator compared to "robustus". The ventomedial angle
of the astragalar body is intermediate in Fukuiraptor between Australovenator
and "robustus". In both Fukuiraptor and "robustus",
the transverse condylar groove angles dorsomedially, whereas it angles ventromedially
in Australovenator. Fukuiraptor and Australovenator have
a shorter straight lateroventral edge to the astragalar body. Thus there seems
no reason to believe "robustus" is closer to Australovenator
than to Fukuiraptor, besides provenance. As Megaraptora has recently
been placed in both Allosauroidea and Coelurosauria by different authors, Molnar's
and Welles' early arguments were each prescient and Chure's compromise accurate.
Agnolin et al. (2005) argued "robustus" was probably an abelisauroid,
which was officially published by Agnolin et al. (2010). They argued several
characters support this. They claim the ascending process lacks apical tapering,
but both "robustus" and Australovenator have a roughly parallel
basal portion and a tapered apical portion created by a medal slope. The difference
is one of degree, where the slope in "robustus" is longer. A vertical
groove on the posterior ascending process' surface probably corresponds to the
ridge found in the middle of the process' articular surface in some abelisauroid
tibiae. The ridge has since been identified in numerous tetanurines including
the megaraptoran Aerosteon, and both Australovenator and Fukuiraptor
exhibit the groove. The anterior ridge along the lateral ascending process'
edge is caused by the fibula articulating along that edge. This was originally
thought to be an Allosaurus character by Molnar et al., and said to be
present in Xenotarsosaurus by Agnolin et al., but is found in several
other theropods including Fukuiraptor as well. Finally, their 2010 paper
argued "robustus" has a broad posterior ascending process peaking
near the middle of the astragalar body, but according to Molnar et al.'s original
stereophotos, this is a misinterpretation. Instead, the labeled "plap"
is merely the posterior wall of the normal ascending process, and the lower
medially placed posterior ascending process is like that in Fukuiraptor
and Australovenator. I conclude there is no reason to doubt "robustus"'
megaraptoran placement, and similar arguments were made by Fitzgerald et al.
(2012).
References- Molnar, Flannery and Rich, 1981. An allosaurid theropod dinosaur
from the Early Cretaceous of Victoria, Australia. Alcheringa. 5, 141-146.
Welles, 1983. Allosaurus (Saurischia, Theropoda) not yet in Australia.
Journal of Paleontology. 57, 196.
Molnar, Flannery and Rich, 1985. Aussie Allosaurus after all. Journal
of Paleontology. 59, 1511-1513.
Chure, 1998. A reassessment of the Australian Allosaurus and its implications
for the Australian refugium concept. Journal of Vertebrate Paleontology. 18(3),
34A.
Azuma and Currie, 2000. A new carnosaur (Dinosauria: Theropoda) from the Lower
Cretaceous of Japan. Canadian Journal of Earth Sciences. 37(12), 1735-1753.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. Ph.D. dissertation, Columbia University, 1-964.
Glut, 2003. Dinosaurs - The Encyclopedia - Supplement 3. McFarland Press, Jefferson,
NC.
Agnolin, Ezcurra and Pais, 2005. Systematic reinterpretation of the pigmy Allosaurus
from the Lower Cretaceous of Victoria (Australia). Ameghiniana. 42, 13R.
Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliot, 2009. New Mid-Cretaceous
(Latest Albian) dinosaurs from Winton, Queensland, Australia. PLoS ONE. 4(7),
e6190.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna
from southern Australia indicates high polar diversity and climate-driven dinosaur
provinciality. PLoS ONE. 7(5), e37122.
Fitzgerald, Carrano, Holland, Wagstaff, Pickering, Rich and Vickers-Rich, 2012.
First ceratosaurian dinosaur from Australia. Naturwissenschaften. 99, 397-405.
"Allosaurus"
sibiricus Riabinin, 1915
= Antrodemus sibiricus (Riabinin, 1915) Huene, 1932
= Chilantaisaurus? sibiricus (Riabinin, 1915) Molnar, Kurzanov and Dong,
1990
Berraisian-Hauterivian, Early Cretaceous
Tignin Formation or Turgin Formation or Zugmar Formation, Chitinskaya Oblast,
Russia
Holotype- (PIN coll.) distal metatarsal II (~300-400 mm)
Late Barremian-Mid Aptian, Early Cretaceous
Mogoito Member of Murtoi Formation, Buryatia, Russia
Referred- ? bone (Ivanov, 1940)
Diagnosis- (suggested) (combination of) metatarsal II lateral condyle
with ventral surface width ~46% of condylar depth; metatarsal II medial condyle
dorsal surface ~73% as wide as surface of lateral condyle and angled 35 degrees
from lateral edge of lateral condyle.
Comments- Note the description was actually published in 1915, though
the volume was intended for 1914. The holotype was discovered in 1912 and deposited
in what was then the Geological Museum of the Russian Academy of Sciences in
Petrograd (Tolmachoff, 1924) (now St. Petersburg), which has since moved to
Moscow. Riabinin (1915; partially translated in Chure, 2000) named it Allosaurus
(?) sibiricus based on what he identified as a distal metatarsal IV. Huene
(1932) said only that it did not permit exact characterization and probably
belonged to an allosaurid (he renamed it Antrodemus? sibiricus as he
thought that was a senior synonym of Allosaurus). Molnar et al. (1981)
felt sibiricus resembled Ceratosaurus so closely that they "hesitate
to accept it as an allosaurid". Molnar et al. (1990) on the other hand
stated it was "almost identical with that of C. tashuikouensis in
form and proportions of the distal condyle", so questionably referred it
to Chilantaisaurus. Nessov (1995) agreed the species were similar and
that sibiricus may belong in Chilantaisaurus. He noted stratigraphic
data gave three possible formations the specimen was discovered in. Chure (2000)
incorrectly said Riabinin did not illustrate the material, as he apparently
had only a small portion of the original document. This error was repeated by
Benson and Xu (2008). Chure excluded it from Allosauridae because he believed
the distal outline was rectangular, but it is actually trapezoidal in both Allosaurus
and sibiricus. Both Benson and Xu and Carrano et al. (2012) incorrectly
credited Holtz et al. (2004) as being responsible for assigning it to Chilantaisaurus,
though Carrano et al. correctly identified it as a second metatarsal. They believed
it "too fragmentary to be assigned to a known taxon or identified as a
distinct form" and noted similarity to Allosaurus, Neovenator,
Torvosaurus and Afrovenator (though no metatarsal II has been
reported for the latter).
Ivanov (1940) reported a bone referred to Allosaurus sibiricus from the
Mortoi Formation, though without more data this referral is uncertain.
The holotype is ~70% the size of Chilantaisaurus tashuikouensis in distal
width and depth, which would make it ~286 mm if similarly stout. If from an
elongate metatarsus like Australovenator's though, it would be ~398 mm
long. It differs from Allosaurus in having a more medially oriented dorsal
curve to the lateral condyle, having a larger and more medially flaring medial
condyle, having a lateral condyle which is recessed ventrally, and lacking the
lateral flare on the ventral edge of Allosaurus' lateral condyle. Chilantaisaurus'
medial condyle flares slightly more than Allosaurus' and it has a ventrally
recessed lateral condyle, but it has the small medial condyle and a lateral
flare like Allosaurus and has the dorsal curve oriented even further
laterally. Of course, Riabinin and Molnar et al. were comparing sibiricus
with the fourth metatarsals of Ceratosaurus, Allosaurus and Chilantaisaurus,
not the second metatarsals (both fourth metatarsals differ from sibiricus
in having a ventrally pointed medial condyle and lacking a ventrally inset medial
condyle, while Allosaurus' element is much narrower, and Chilantaisaurus'
has a larger dorsolateral bulge; Molnar et al. were correct that Chilantaisaurus'
is more similar, but it is not almost identical). Taxa with similarly medially
flaring medial condyles are Ceratosaurus, Torvosaurus, Sinraptor,
Acrocanthosaurus, Fukuiraptor, Australovenator, Megaraptor
and Harpymimus, though only Australovenator's and Acrocanthosaurus'
are close in size. Taxa with a ventrally inset lateral margin are Rajasaurus,
Megalosaurus, Chilantaisaurus, Fukuiraptor, Australovenator,
Megaraptor, Appalachiosaurus and Alxasaurus. Overall, it
is most similar to Australovenator, differing in being 9% broader compared
to depth, a broader lateral condyle ventrally, and having a more medially oriented
dorsal surface of the lateral condyle. Next most similar is Megaraptor,
which it differs from in having a broader lateral condyle ventrally with ventral
surface angled more laterally, and a less rounded dorsal surface of the lateral
condyle. The amount of ventral inset of the lateral condyle and dorsal exposure
of the medial condyle is in between these two taxa. This suggests sibiricus
may be a megaraptoran, which is congruent with its age, size and location. As
it is intermediate in two variables, more similar to Megaraptor in depth,
more similar to Australovenator in the orientation of the lateral condyle's
ventral surface, and differs from both in the lateral condyle's ventral width,
placing it in any named genus is not possible. As it does differ from all known
theropod metatarsals, it is not a nomen dubium, contra Rauhut (2003), Holtz
et al. and Carrano et al..
References- Riabinin, 1915. Zamtka o dinozavry ise Zabaykalya [A note
on a dinosaur from the trans-Baikal region]. Trudy Geologichyeskago Muszeyah
Imeni Petra Velikago Imperatorskoy Academiy Nauk. 8(5), 133-140.
Tolmachoff, 1924. On dinosaurs in northern Asia. American Journal of Science.
5(7), 489-490.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Ivanov, 1940. [On the age of the coal-bearing deposits of Transbaikalia]. Sovietskaya
Geologiya. 11, 45-54.
Molnar, Flannery and Rich, 1981. An allosaurid theropod dinosaur from the early
Cretaceous of Victoria, Australia. Alcheringa. 5, 141-146.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria. Berkeley: University of California Press. 169-209.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii [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. 156 pp.
Chure, 2000. A new species of Allosaurus from the Morrison Formation
of Dinosaur National Monument (Utah-Colorado) and a revision of the theropod
family Allosauridae. Ph.D. thesis. Columbia University. 964 pp.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 213 pp.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Benson and Xu, 2008. The anatomy and systematic position of the theropod dinosaur
Chilantaisaurus tashuikouensis Hu, 1964 from the Early Cretaceous of
Alanshan, People’s Republic of China. Geological Magazine. 145(6), 778-789.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Rapator Huene, 1932
R. ornitholestoides Huene, 1932
Early Cenomanian, Late Cretaceous
Griman Creek Formation, New South Wales, Australia
Holotype- (NHMUK R3718) (~5 m) metacarpal I (70 mm)
Referred- ?(AM F112816) incomplete mid caudal centrum (Brougham, Smith and Bell, 2019)
?(LRF 100-106) (~6 m) rib fragments, gastralial fragments,
proximal ulna, proximal manual ungual ?I, ilial fragment, ?fibular fragments,
incomplete metatarsal III (350 mm), fragments (Bell, Cau, Fanti and Smith, 2016)
Diagnosis- (after Hocknull et al., 2009) differs from Australovenator
in- more concave proximal articular surface.
(after White et al., 2013) differs from Australovenator in- less tapered
proximomedial process; more laterally concave distal shaft; medial condyle projects
less medially due to less ventromedial angling; medial condyle angled less transversely;
deeper trochlea both distally and ventrally; dorsal longitudinal ridge proximal
to medial condyle; medial condyle transversely compressed in ventral view; more
distally restricted metacarpal II facet; medial condyle only slightly concave
medially in distal view; no transverse ridge on medial half of proximal surface;
dorsal edge of proximal surface more concave.
(after Bell et al., 2016) proximal end of metatarsal III strongly asymmetrical
in medial/lateral view with trapezoidal anterior process extending further distally
along the shaft than the posterior process giving a ball-peen hammer-shaped
profile; contact for metatarsal II on metatarsal III divided into anterior and
posterior halves by shallow, longitudinal groove.
Differs from Australovenator in- more robust anterior process on ulna;
more gracile manual ungual I with sharply defined median ridge on proximal articular
surface; prominent, broad groove between articular facet and flexor tubercle
on manual ungual I (also in Megaraptor); metatarsal III with well-developed
lateral ridge on proximal shaft; distal articular surface of metatarsal III
as wide as it is long.
Comments- Originally identified as a compsognathid metacarpal I similar
to Ornitholestes and Oviraptor (Huene, 1932), Molnar (1992) suggested
it was an abelisaurid based on biogeography. Headden (DML, 2000) later noticed
similarities between it and alvarezsaurid phalanx I-1, which was the conclusion
published by Holtz et al. (2004). Salisbury et al. (2007) stated it may belong
to a Nqwebasaurus-like basal coelurosaur, presumably as a metacarpal
I once more. Agnolin et al. (2010) confirmed it was a first metacarpal but found
it to be most similar to Megaraptor and Australovenator, especially
the latter. They incorrectly called it a nomen dubium, despite saying it and
Australovenator differ from Megaraptor in having a more dorsoventrally
developed mediodistal condyle and a metacarpal II facet lying in almost the
same plane as the lateral margin of the shaft, and that it differs from Australovenator
in several other features. Agnolin et al. made it a nomen dubium because of
its fragmentary condition (irrelevant), the absence of autapomorphies (irrelevant
given the unique combination of characters) and absence of clear differences
with Australovenator (which could only make it a senior synonym of the
latter, as no other taxon was presented as having no/subtle differences from
Rapator). White et al. (2013) explicitly compared the two genera based
on a more complete metacarpal I of Australovenator and digital scans
of each, finding Rapator differs in numerous details. The authors viewed
these as "sufficient differences to warrant assignment to separate genera",
but this seems premature without analyzing variation in other taxa. The genera
are kept separate here due to the combination of morphological and stratigraphic
difference.
Bell et al. (2016) described a fragmentary postcranium from the same formation
as Rapator. They referred to this as Megaraptoridae indet., though listed
several characters distinguishing it from Australovenator and other megaraptorids.
Given the locality and lack of megaraptoran diversity in other formations, this
specimen is here provisionally referred to Rapator although it cannot
be directly compared since metacarpal I is unpreserved.
Brougham et al. (2019) described caudal AM F112816 as Megaraptora indet. based on its pneumaticity.
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre
Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1),
361 pp.
Molnar, 1992. Paleozoogeographic relationships of Australian Mesozoic tetrapods.
In Chatterjee and Hotton (eds.). New Concepts in Global Tectonics. Texas Technical
Press, USA. 259-265.
Headden, DML 2000. https://web.archive.org/web/20201116202321/http://dml.cmnh.org/2000Mar/msg00555.html
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 861 pp.
Salisbury, Agnolin, Ezcurra and Pias, 2007. A critical reassessment of the Creaceous
non-avian dinosaur faunas of Australia and New Zealand. Journal of Vertebrate
Paleontology. 27(3), 138A.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
White, Falkingham, Cook, Hocknull and Elliott, 2013. Morphological comparisons
of metacarpal I for Australovenator wintonensis and Rapator ornitholestoides:
Implications for their taxonomic relationships. Alcheringa. 37(4), 435-441.
Bell, Cau, Fanti and Smith, 2016 (online 2015). A large-clawed theropod (Dinosauria: Tetanurae)
from the Lower Cretaceous of Australia and the Gondwanan origin of megaraptorid
theropods. Gondwana Research. 36, 473-487.
Brougham, Smith and Bell, 2019. New theropod (Tetanurae: Avetheropoda)
material from the 'mid'-Cretaceous Griman Greek Formation at Lightning
Ridge, New South Wales, Australia. Royal Society Open Science. 6,
180826.
Australovenator Hocknull,
White, Tischler, Cook, Calleja, Sloan and Elliot, 2009
A. wintonensis Hocknull, White, Tischler, Cook, Calleja, Sloan
and Elliot, 2009
Cenomanian, Late Cretaceous
Winton Formation, Queensland, Australia
Holotype- (AODF 604) (~5.5 m) dentaries (342.6 mm), proximal first dorsal
rib, proximal second or third dorsal rib, proximal seventh or eighth dorsal
rib, dorsal rib shaft fragments, nine gastralial fragments, humeri (307.35,
303.35 mm), radii (one incomplete; 215.37, 211.28 mm), ulnae (265.58, 267.22
mm), radiale, distal carpal I, metacarpals I (78.38, 79.91 mm), phalanges I-1
(118.21, 111.1 mm), incomplete manual ungual I (150.95 mm straight, 190 mm on
curve), metacarpal II (138.15 mm), phalanx II-1 (84.9 mm), phalanx II-2 (86.51
mm), incomplete manual ungual II, phalanx III-1 (73.11 mm), phalanges III-3
(39.44, 41.6 mm), manual ungual III (75.12 mm straight), partial ilium, femur
(578 mm), tibiae (569, 564 mm), fibulae (538 mm; one incomplete), astragalus
(105 mm wide), metatarsal I (66 mm), pedal unguals I (one incomplete; 66 mm),
metatarsal II (284 mm), incomplete phalanx II-1 (~106 mm), incomplete phalanx
II-2 (~64 mm), pedal ungual II (84mm), metatarsal III (322 mm), incomplete phalanx
III-1 (~115 mm), phalanx III-2 (106 mm), phalanges III-3 (one partial; 73 mm),
pedal ungual III (95 mm), incomplete metatarsal IV, phalanx IV-1 (82 mm), incomplete
phalanx IV-2 (49 mm), phalanx IV-3 (46 mm), phalanx IV-4 (33 mm), pedal ungual
IV (77 mm)
Paratypes- (AODF 822) anterior tooth (14x6.8x6 mm)
(AODF 823) lateral tooth (13.6x9.35.5 mm)
(AODF 824) lateral tooth (12.8x8.4x4.3 mm)
(AODF 825) lateral tooth (18.4x10x6.2 mm)
(AODF 826) lateral tooth (16.5x9.5x6.5 mm)
(AODF 827) tooth (19.8x9.7x? mm)
(AODF 828) lateral tooth (14.4x9.4x5.8 mm)
(AODF 829) lateral tooth (16.4x10.4x5.7 mm)
(AODF 830) tooth (16.3x11.7x? mm)
(AODF 831) anterior tooth (12.5x7.3x4.8 mm)
Referred- (AODF 664) anterior tooth (9x5.3x4.1 mm) (White, Bell, Cook,
Poropat and Elliott, 2015)
(AODF 819) lateral tooth (17.8x8.3x10.2 mm) (White, Bell, Cook, Poropat and
Elliott, 2015)
(AODF 820) lateral tooth (16.5x9.7x7.2 mm) (White, Bell, Cook, Poropat and Elliott,
2015)
?(AODF 967) partial caudal centrum (White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020)
....(AODF 968) partial caudal centrum (White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020)
....(AODF 972) distal pedal phalanx II-1 (White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020)
....(AODF 977) proximal metatarsal II (White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020)
....(AODF 978) distal metatarsal II (White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020)
....(AODF 979) distal metatarsal IV (White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020)
....(AODF coll.) fragments (White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020)
Diagnosis- (after Hocknull et al., 2009) at least eighteen [nineteen
based on White et al., 2015] dentary teeth (also in Compsognathus); dorsal
ribs with pneumatic cavities (also in Aerosteon); olecranon process inflated
in proximal view; round and discontinuous lateral tuberosity on ulna; distal
extensor groove deep and narrow (also in Bagaraatan and Xiongguanlong);
ventral process on anterior edge of lateral tibial condyle; proximal articular
surface of fibula bevelled to be higher anteriorly (also in Scipionyx).
Other diagnoses- Hocknull et al. (2009) also included several characters
which are primitive for coelurosaurs- gracile dentary; dentary with subparallel
dorsal and ventral margins; rounded dentary symphysis, chin absent on dentary;
primary row of dentary neurovascular foramina not decurved posteriorly; gastralia
unfused; gastralia distally tapered; ulna straight; femoral flexor groove lacks
cruciate ridge; lateral malleolus of tibia extends distal of medial malleolus;
medial astragalar condyle transversely expanded; astragalus with tall ascending
process; anterior groove across astragalar condyles; groove at base of astragalar
ascending process; metatarsals elongate and gracile. Others are also present
in Fukuiraptor- fused interdental plates; anterolateral groove on ulnar
shaft; anterior trochanter extends proximally to near proximal greater trochanter
edge; anterolateral process projects from antero proximal margin of astragalar
lateral condyle. The general trend of a quadrangular first dentary alveolus,
followed by several cicular alveoli and then transversely compressed alveoli
is common in theropods. A dorsally directed femoral head is also present in
Chilantaisaurus, Gasosaurus, Bagaraatan and tyrannosauroids.
An anteromedially directed femoral head is also present in Megaraptor, Tugulusaurus
and Xiongguanlong.
Comments- White et al. (2012) redescribed the forelimb including many
new elements, finding a supposed distal manual ungual II to be an incomplete
pedal ungual I, supposed manual phalanx II-2 is III-1, and supposed right metacarpal
II is from the left side. Similarly, White et al. (2013) redescribed the hindlimb
elements, finding metatarsal I is from the left side, and that some pedal phalanges
were incorrectly identified. White et al. (2015) described a newly described
dentary of the holotype, believed the isolated teeth found with the holotype
were not referrable to that individual, and referred additional teeth from other
localities.
Discovered in 2018,
White et al. (2020) stated "the close proximity and size congruence of
the specimens recovered from AODL 261 suggests that they pertain to a
single individual." The authors felt "the identification of the AODL
261 material as megaraptorid lies principally on the presence of
pleurocoels on the two incomplete caudal centra" and that "The distal
end of metatarsal II (AODF 978) also bears some resemblance to that of
a specimen assigned to Megaraptor sp. (UNPSJB-PV 944) and to a lesser extent Australovenator." It is tentatively listed under Australovenator here due to provenance.
Hocknull et al. (2009) included Australovenator in the carnosaur analysis
of Brusatte and Sereno (2008) and found it emerges as the basalmost carcharodontosaurid.
However, this did not include the astragalar characters they note are shared
with Fukuiraptor,
nor any coelurosaurs except Compsognathidae. Benson et al. (2010) using
a larger dataset found it to be a megaraptoran carcharodontosaurid, but
again included few coelurosaurs. While this position within
Megaraptora is now well established, the placement of that clade is
still controversial.
References- Brusatte and Sereno, 2008. Phylogeny of Allosauroidea (Dinosauria:
Theropoda): Comparative analysis and resolution. Journal of Systematic Palaeontology.
6(2), 155-182.
Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliot, 2009. New Mid-Cretaceous
(Latest Albian) dinosaurs from Winton, Queensland, Australia. PLoS ONE. 4(7),
e6190.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
White, Cook, Hocknull, Sloan, Sinapius and Elliott, 2012. New forearm elements
discovered of holotype specimen Australovenator wintonensis from Winton,
Queensland, Australia. PLoS ONE. 7(6), e39364.
White, Benson, Tischler, Hocknull, Cook, Barnes, Poropat, Wooldridge, Sloan,
Sinapius and Elliot, 2013. New Australovenator hind limb elements pertaining
to the holotype reveal the most complete neovenatorid leg. PLoS ONE. 8(7), e68649.
White, Bell, Cook, Poropat and Elliott, 2015. The dentary of Australovenator
wintonensis (Theropoda, Megaraptoridae); implications for megaraptorid dentition.
PeerJ. 3:e1512.
White, Bell,
Poropat, Pentland, Rigby, Cook, Sloan and Elliott, 2020. New theropod
remains and implications for megaraptorid diversity in the Winton
Formation (lower Upper Cretaceous), Queensland, Australia. Royal
Society Open Science. 7, 191462.
Megaraptor Novas, 1998
= "Megaraptor" Shreeve, 1997
Diagnosis- (after Calvo et al., 2004) manual unguals I and II enlarged
and highly transversely compressed.
(after Smith et al., 2008) proximocaudally expanded blade-like olecranon process
that extends distally as a caudal olecranon crest; pronounced lateral tuberosity
that is continuous distally with a distinct lateral crest.
M. namunhuaiquii Novas, 1998
= "Megaraptor namuhualquii" Shreeve, 1997
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of the Rio Neuquen Subgroup, Neuquen, Argentina
Holotype- (MCF-PVPH 79) ulna (332 mm), phalanx I-1 (188 mm), manual ungual
I (339 mm), distal metatarsal III (~450 mm)
....(MCF-PVPH 80) two partial dorsal ribs, several gastralia, four
proximal caudal vertebrae (~95, ~100 mm), distal tibial fragment (Coria
and Currie, 2016)
Referred- (MUCPv 341) (7 year old adult) ?sixth cervical vertebra (~84
mm), two proximal caudal vertebrae, three chevrons, incomplete scapula, coracoid,
radius (~369 mm), ulna, semilunate carpal, distal carpal II?, metacarpal I (106 mm), phalanx
I-1 (182 mm), manual ungual I (350 mm), metacarpal II (170 mm), phalanx II-1
(108 mm), phalanx II-2 (104 mm), manual ungual II (235 mm), metacarpal III (119
mm), phalanx III-1 (56 m), phalanx III-2 (41 mm), phalanx III-3 (56 mm), manual
ungual III (65 mm), metacarpal IV (40 mm), pubis (480 mm) (Calvo
et al., 2004)
(smaller individual) metatarsal IV (~290 mm) (Calvo
et al., 2004)
(MUCPv 412) distal ulna (Porfiri, Calvo and Santos, 2007)
(MUCPv 413) proximal manual phalanx I-1 (Porfiri, Calvo and Santos, 2007)
(MUCPv 595) (6 year old juvenile) incomplete premaxillae, maxillae (one incomplete),
nasals, frontal, partial braincase, axis, third cervical vertebra, fourth cervical
vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical
vertebra, eighth cervical vertebra (~49 mm), ninth cervical vertebra, tenth
cervical vertebra, first dorsal vertebra, second dorsal vertebra, third dorsal
vertebra, fourth dorsal vertebra, fifth dorsal vertebra, sixth dorsal vertebra,
seventh dorsal vertebra, eighth dorsal vertebra, tenth dorsal vertebra, eleventh
dorsal vertebra, twelfth dorsal vertebra, dorsal ribs, eight gastralia, third
sacral vertebra, fourth sacral vertebra, fifth sacral vertebra, four proximal
caudal vertebrae, scapulae, coracoid, incomplete humeri (~232 mm), partial pubes
(Porfiri, Novas, Clavo, Agnolin, Ezcurra and Cerda, 2014)
?(MUCPv 723) tooth (49x13x? mm) (Coria and Currie, 2016)
(MUCPv 1353) (12 year old adult, ~1 ton) specimen including forelimb elements and pubis
(Porfiri, Novas, Clavo, Agnolin, Ezcurra and Cerda, 2014)
?(MUCPv coll.) teeth (Poblete and Calvo, 2003)
Diagnosis- (after Novas, 1998; Lamanna, 2004; Calvo et al., 2004) cervical
vertebrae with elongate elliptical pleurocoels; blade-like olecranon process
of ulna; ulna stout and triangular in distal view; manual phalanx I-1 subquadrangular
in proximal view, with dorsal portion wider than ventral portion; proximodorsal
depression on manual ungual I absent; lateral vascular groove in manual ungual
III absent; metatarsal III with deep and wide extensor ligament pit; distal
end of metatarsal IV narrower than shaft.
(after Aranciaga Rolando et al., 2015) metacarpal I with tapered medial condyle;
manual ungual I approaching length of ulna; manual digit II phalanges with ventromedial
flanges.
Comments-
Coria and Currie (2016) state "there are a number of unprepared and
undescribed bones (MCF-PVPH-80) that were collected in association with
MCF-PVPH-079, including four caudal vertebrae, two ribs (one almost
complete), and gastralia. This cluster of fossils was found far enough
from the material described as the holotype of Megaraptor namunhuaiquii
that the collectors could not be sure whether or not they belonged to
the same specimen (Novas, personal communication 2004)."
Additionally, "one almost complete dorsal rib would have been found
associated with MCF-PVPH 79 (Novas, personal communication 2006)" and
all of these materials plus "a medial fragment of the distal end of the
right tibia" are described as MCF-PVPH-079.
Lammana et al. (2020) state "the complete right metatarsal IV assigned
to MUCPv 341 by Calvo et al. (2004) is substantially shorter than would
be expected for this animal. Given that this metatarsal was
recovered at least 3.5 m away from all other bones that comprise this
specimen (see Calvo et al. 2004:567), we suspect that it may pertain to
a different theropod individual. More specifically, based on its
possession of several probable megaraptorid synapomorphies" it "may
indicate the presence of a fourth, smaller individual in the quarry
(Porfiri pers. comm. to MCL)."
The teeth mentioned by Poblete and Calvo (2003) are labiolingually
compressed teeth with strongly posteriorly inclined crowns which are
sometimes proportionally short. They lack mesial serrations but have
distal ones. A labial carina is present, continuing apically to the tip
of the distal side. It sometimes has small serrations. Gianechini et
al. (2011) states they are similar to Bajo de la Carpa tooth MPCA 247
which has been recognized as megaraptorid.
This was first reported by Novas et al. (1996) in an abstract
as a possibly maniraptoran tetanurine. Originally thought to have an enlarged
hyperextensible pedal ungual II, a newer specimen (Calvo et al., 2002) shows this
is actually manual ungual I. Calvo et al. (2004) suggested Megaraptor
was a non-avetheropod tetanurine, though without an analysis to back it up.
Lamanna (2004) entered it in Rauhut's analysis and found it to be an allosauroid
carnosaur. Smith et al. (2007) recovered Megaraptor as a carcharodontosaurine
in their analysis, then later (2008) as the sister taxon of Spinosauridae. Benson
(2010) recovered it as a carnosaur or sister taxon to Avetheropoda, but when
more characters and taxa were added (Benson et al., 2010), it formed part of
a larger clade within Carcharodontosauridae they named Megaraptora (closest
to Aerosteon within that clade). Carrano et al. (2012) also recover it
as a megaraptoran carcharodontosaurid, and as it takes 12 more steps to force
it into Carcharodontosaurinae and 15 more steps to force it into Megalosauroidea,
the latter two options are improbable. Most recently, Novas et al. (2013) and
Porfiri et al. (2014) have recovered Megaraptora within Tyrannosauroidea, in
part due to the Dilong-like skull of MUCPv 595.
References- Novas, Cladera and Puerta, 1996. New theropods from the Late
Cretacoues of Patagonia. Journal of Vertebrate Paleontology. 16(3), 56A.
Novas, 1998. Megaraptor namunhuaiquii, gen. et sp. nov., a large-clawed,
Late Cretaceous theropod from Patagonia. Journal of Vertebrate Paleontology.
18(1), 4-9.
Calvo, Porfiri, Veralli and Novas, 2002. Megaraptor namunhuaiquii (Novas,
1998), a new light about its phylogenetic relationships. Primer Congreso latinoamericano
de Paleontolog�a de Vertebrados. Santiago de Chile, Octubre del 2002.
p.20.
Poblete and Calvo, 2003. Upper Turonian dromaeosaurid teeth
from Futalognko quarry, Barreales Lake, Neuqu�n, Patagonia, Argentina.
Ameghiniana. 40(S), 66R.
Calvo, Porfiri, Veralli, Novas and Pobletei, 2004. Phylogenetic status of Megaraptor
namunhuaiquii Novas based on a new specimen from Neuqu�n, Patagonia,
Argentina. Ameghiniana (Rev. Asoc. Paleontol. Argent.). 41(4), 565-575.
Lamanna, 2004. Late Cretaceous dinosaurs and crocodiliforms from Egypt and Argentina.
PhD Thesis. University of Pennsylvania. 305 pp.
Porfiri, Calvo and Santos, 2007. Evidencia de gregarismo en Megaraptor
namunhuaiquii (Theropoda: Tetanurae), Patagonia, Argentina. in D�az-Mart�nez
and R�bano (eds.). 4th European Meeting on the Palaeontology and Stratigraphy
of Latin America. 323-326.
Porfiri, Santos and Calvo, 2007. New information on Megaraptor namunhuaiquii
(Theropoda: Tetanurae), Patagonia: Considerations on paleoecological aspects.
Arquivos do Museu Nacional, Rio de Janeiro. 65(4), 545-550.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti
(Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications
for early theropod evolution. Zoological Journal of the Linnean Society. 151,
377-421.
Smith, Makovicky, Agnolin, Ezcurra, Pais and Salisbury, 2008. A Megaraptor-like
theropod (Dinosauria: Tetanurae) in Australia: Support for faunal exchange across
eastern and western Gondwana in the Mid-Cretaceous. Proceedings of the Royal
Society B. 275(1647), 2085-2093.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda)
from the Bathonian of the UK and the relationships of Middle Jurassic theropods.
Zoological Journal of the Linnean Society. 158(4), 882-935.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Gianechini, Lio and Apestegu�a, 2011. Isolated archosaurian teeth from
"La Bonita" locality (Late Cretaceous, Santonian-Campanian), R�o Negro
Province, Argentina. Historia Natural, tercera serie. 1, 5-16.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Porfiri, Novas, Clavo, Agnolin, Ezcurra and Cerda, 2014. Juvenile specimen of
Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation.
Cretaceous Research. 51, 35-55.
Aranciaga Rolando, Agnolin, Brisson Edli and Novas, 2015. Osteologia del miembro
anterior de Megaraptor namunhuaiquii y sus implicancias filogeneticas. XXIX
Jornadas Argentinas de Paleontolog�a de Vertebrados, resumenes. Ameghiniana.
52(4) suplemento, 5-6.
Coria and Currie, 2016. A new megaraptoran dinosaur (Dinosauria, Theropoda,
Megaraptoridae) from the Late Cretaceous of Patagonia. PLoS ONE. 11(7), e0157973.
Lamanna, Casal, Martinez and Ibiricu, 2020. Megaraptorid (Theropoda:
Tetanurae) partial skeletons from the Upper Cretaceous Bajo Barreal
Formation of central Patagonia, argentina: Implications for the
evolution of large body size in Gondwanan megaraptorans. Annals of
Carnegie Museum. 86(3), 255-294.
Murusraptor Coria and
Currie, 2016
M. barrosaensis Coria and Currie, 2016
Coniacian, Late Cretaceous
Sierra Barrosa Formation of the Rio Neuquen Subgroup, Neuquen, Argentina
Holotype- (MCF-PVPH 411) (~7 m, ~1.5 tons) lacrimal, prefrontal, postorbital,
quadrate (149 mm), braincase, pterygoids, incomplete ectopterygoid, posterior
mandible, some of thirty-one teeth (12.3-49.8 mm), anterior cervical rib, second dorsal
neural arch, sixth dorsal neural arch, seventh dorsal neural arch, eighth dorsal
neural arch, tenth dorsal neural arch, eleventh dorsal vertebra (82 mm), twelve
dorsal ribs (first 631, seventh 710, , dorsal rib fragments, gastralia, partial
fused first (110 mm) and second sacral vertebrae, third or fourth sacral neural
arch, first caudal neural arch, two proximal caudal neural arches, proximal
chevron (238 mm), incomplete manual ungual III (~62 mm), ilium (750 mm), proximal
pubes, incomplete ischia, tibia (690 mm), calcaneum
Diagnosis- (after Coria and Currie, 2016) anterodorsal lacrimal process
longer than height of ventral process; thick, shelf-like thickening on lateral
surface of surangular ventral to groove between anterior surangular foramen
and insert for uppermost intramandibular process of the dentary; sacral ribs
hollow and tubelike; short ischia distally flattened and slightly expanded dorsoventrally.
Comments-
Discovered in 2001, Coria and Currie (2002) and Paulina-Carabajal
(2009) assign this specimen to Coelurosauria, the latter describing its
braincase in detail (eventually published as Paulina-Carabajal and
Currie, 2017). Novas et al. (2013) propose a megaraptoran affinity and
Paulina-Carabajal and Coria (2015) refer to it as a "probable
megaraptorid." The specimen was finally described in detail by Coria
and Currie (2016) as Murusraptor,
recovered as a megaraptorid in a version of Carrano et al.'s tetanurine
analysis and as a megaraptoran in Novas et al.'s tetanurine analysis.
Note the quadratojugal, palatine, hyoids and cervical vertebrae
reported in 2001 must have been misidentified as they are not present
in the final description, and that the sediments were reassigned from
the Plottier Formation to the Sierra Barrosa Formation. Poropat
et al. (2019) said regarding the dentition that "personal observation
of these teeth (by S.F.P., 2018) revealed a high degree of size and
morphological disparity - it is probable that they derive from more
than one megaraptorid individual and that non-megaraptorid theropod
teeth are also present in the sample."
References- Coria, Currie, Eberth, Garrido and Koppelhus, 2001. Nuevos
vertebrados f�siles del Cret�cico Superior de Neuqu�n.
Ameghiniana. 38, 6R-7R.
Coria and Currie, 2002. Un gran ter�podo celurosaurio en el Cret�cico
Superior de Neuqu�n. Ameghiniana. 39, 9R.
Paulina-Carabajal, 2009. El neurocr�neo de los dinosaurios Theropoda
de la Argentina: Osteolog�a y sus implicancias filogen�ticas.
PhD thesis, Universidad Nacional de La Plata. 554 pp.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Paulina-Carabajal and Coria, 2015. An unusual theropod frontal from the Upper
Cretaceous of north Patagonia. Alcheringa. 39(4), 514-518.
Coria and Currie, 2016. A new megaraptoran dinosaur (Dinosauria, Theropoda,
Megaraptoridae) from the Late Cretaceous of Patagonia. PLoS ONE. 11(7), e0157973.
Paulina-Carabajal and Currie, 2017. The braincase of the theropod dinosaur Murusraptor:
Osteology, neuroanatomy and comments on the paleobiological
implications of certain endocranial features. Ameghiniana. 54, 617-640.
Aranciaga Rolando, Novas and Agnolin, 2019. A reanalysis of Murusraptor barrosaensis
Coria & Currie (2016) affords new evidence about the phylogenetical
relationships of Megaraptora. Cretaceous Research. 99, 104-127.
Aerosteon Sereno, Martinez,
Wilson, Varricchio, Alcober and Larsson, 2009
= "Aerosteon" Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson,
2008 online
A. riocoloradensis Sereno, Martinez, Wilson, Varricchio, Alcober
and Larsson, 2009
= "Aerosteon riocoloradensis" Sereno, Martinez, Wilson, Varricchio,
Alcober and Larsson, 2008 online
Late Coniacian, Late Cretaceous
Plottier Formation of the Rio Neuquen Subgroup, Mendoza, Argentina
Holotype- (MCNA-PV-3137) (~8.5 m; subadult) prefrontal (68 mm), postorbital
(114 mm long), quadrate (163 mm), posterior pterygoid, prearticular, atlas (25 mm), third cervical vertebra (96 mm), fourth
cervical vertebra (98 mm), sixth cervical vertebra (91 mm), eighth cervical
vertebra, two cervical ribs, first dorsal vertebra (85 mm), fourth dorsal vertebra
(71 mm), fifth dorsal vertebra, sixth dorsal vertebra, seventh dorsal vertebra,
eighth dorsal vertebra (88 mm), ninth dorsal vertebra, tenth dorsal vertebra
(84 mm), eleventh dorsal vertebra (84 mm), fourteenth dorsal vertebra (102 mm),
four dorsal ribs, gastralia, first sacral vertebral fragments, second sacral
vertebra, third sacral vertebra, fourth sacral vertebra, partial fifth sacral
centrum, fifth sacral transverse process, first caudal vertebra (93 mm), mid
caudal centrum (100 mm), distal caudal centrum, furcula, scapula (570 mm), coracoid
(276 mm), ilium (768 mm), pubes (620 mm)
Referred- ?(MCNA-PV-3075) manual ungual II (Novas, Agnolin, Ezcurra,
Porfiri and Canale, 2013)
?(MCNA-PV-3138) incomplete pes including metatarsal II, metatarsal III, metatarsal
IV and phalanges (Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson,
2009)
(MCNA-PV-3139) tibia, incomplete fibula, astragalus, calcaneum (Sereno, Martinez,
Wilson, Varricchio, Alcober and Larsson, 2009)
Diagnosis- (modified after Sereno et al., 2008) prefrontal with a very
short ventral process; enlarged paraquadrate foramen located entirely within
the quadrate; large tympanic diverticulum into the quadrate shaft above the
articular condyle; anterior dorsal vertebra with very large parapophyses; dorsal
neural spines with central pneumatic space (also in Acrocanthosaurus);
posteriormost dorsal vertebra with anterodorsally inclined neural spine; posteriormost
dorsal vertebra with a pneumatic canal within the transverse process (also in
Neovenator); medial gastral elements coossified with anterior and posterior
flanges; furcula with median pneumatocoel.
(after Carrano et al., 2012) robust, cylindrical transverse processes on proximal
caudal vertebrae; fossa on lateral surface of coracoid dorsal to glenoid and
(separate) subglenoid fossa.
Comments- Aerosteon
was originally named in the online-only publication PLoS ONE on
September 30 2008, which did not satisfy the ICZN's requirement (then
Article 8.6, since ammended) that publications be printed on paper and
"contain a statement that copies (in the form in which it is published)
have been deposited in at least 5 major publicly accessible libraries
which are identified by name in the work itself." This requirement was
only met on May 21 2009 (PLoS ONE, 2009 online), making Aerosteon a nomen nudum until that time.
Although originally thought to derive from the Campanian Anacleto Formation,
Novas et al. (2013) determined it is actually from the Late Coniacian Plottier
Formation.
Benson et al. (2010) tried running the referred hindlimb MCNA-PV-3139 as a separate
OTU, and it also grouped with megaraptorans, suggesting it is properly referred
to Aerosteon.
The identity of the referred metatarsal and associated pes
(MCNA-PV-3138) is less certain. Novas et al. (2013) listed megaraptoran
manual unguals from the same locality which probably belong to the
taxon, though they considered the tooth found with the holotype to be
dubious and perhaps abelisaurid. Hendrickx et al. (2020) tested
this and recovered it as an abelisaurid using cladsitic analyses, so it
is here excluded from the holotype
This taxon was originally referred to Carcharodontosauridae by Alcober et al.
(1998), and later as a carnosaur most similar to Allosaurus (Sereno et
al., 2009). Benson et al. (2010) are the first to include Aerosteon in
a phylogenetic analysis, and found it to be a carcharodontosaurid in their new
clade Megaraptora, closest to Megaraptor
itself. While this position within Megaraptora is now well
established, the placement of that clade is still controversial.
References- Alcober, Sereno, Larsson, Martinez and Varricchio, 1998.
A Late Cretaceous carcharodontosaurid (Theropoda: Allosauroidea) from Argentina.
Journal of Vertebrate Paleontology. 18(3) 23A.
Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2009 (online 2008). Evidence for
avian intrathoracic air sacs in a new predatory dinosaur from Argentina. PLoS
ONE. 3(9), e3303.
PLoS ONE, 2009 online. Steps taken to meet the requirements of the ICZN to make new taxonomic names nomenclaturally available.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 10(2), 211-300.
Rauhut, 2012. A reappraisal of a putative record of abelisauroid theropod dinosaur
from the Middle Jurassic of England. Proceedings of the Geologists' Association.
123(5), 779-786.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Aranciaga Rolando, Brisson Egli, Rozadilla and Novas, 2015. Megaraptoran metatarsals
from the Upper Cretaceous of Mendoza, Argentina. XXIX Jornadas Argentinas de
Paleontolog�a de Vertebrados, resumenes. Ameghiniana. 52(4) suplemento,
16.
Hendrickx, Tschopp and Ezcurra, 2020 (online 2019). Taxonomic
identification of isolated theropod teeth: The case of the shed tooth
crown associated with Aerosteon (Theropoda: Megaraptora) and the dentition of Abelisauridae. Cretaceous Research. 108, 104312.
Maip
Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2022
= "Maip" Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2021 online
M. macrothorax
Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2022
= "Maip macrothorax" Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2021 online
Late Campanian-Early Maastrichtian, Late Cretaceous
Megaraptorid Site, Chorrillo Formation, Santa Cruz, Argentina
Holotype- (MPM 21545) (~8.5 m)
axis, proximal fifth cervical rib, seventh cervical rib, eighth
cervical rib, cervical rib fragments, incomplete second dorsal neural
arch, third dorsal neural arch fragment, fourth dorsal centrum, fifth
dorsal neural arch fragment, incomplete sixth dorsal neural arch,
seventh dorsal neural arch fragment, ninth dorsal vertebra,
~tenth/eleventh dorsal centrum, incomplete thirteenth dorsal centrum,
dorsal transverse
process, first dorsal rib, proximal second dorsal rib, proximal sixth
dorsal rib, dorsal rib fragments, gastralial fragments, ~fourth caudal
neural arch, ~seventh caudal neural arch, partial ~tenth caudal neural
arch, two
vertebral fragments, scapular fragments, coracoid, proximal pubis
Diagnosis- (after Aranciaga
Rolando et al., 2022) fifth-ninth dorsal vertebrae with articular
surface of parapophyses saddle-shaped; ~fourth-seventh caudal vertebrae
with accessory posterior centrodiapophyseal lamina that subdivides the
postzygapophyseal-centrodiapophyseal fossa; first dorsal rib with
honeycomb internal structure on its tubercle; prominent distal
projection on middle of coracoid; coracoid without subglenoid ridge;
coracoid without subglenoid fossa; coracoid with posterodistal margin
forming a long articular surface for sternum.
Comments- Discovered between
January and March 2019, Novas et al. (2019) described MPM 21545 as
Megaraptoridae gen. et sp. indet. 1 based on several elements
(thirteenth dorsal centrum, dorsal transverse
process, dorsal rib fragments, ~tenth caudal neural arch, two
vertebral fragments, proximal pubis). They assigned it to
Megaraptoridae based on paired dorsal pleurocoels separated by septum
and anterior and posterior centrodiapophyseal laminae on
the proximal caudal. Aranciaga Rolando et al. described
additional remains of the specimen and proposed naming it "Maip
macrothorax", originally online on December 22 2021 as a preprint under
consideration at Scientific Reports (although
publically available on ResearchSquare) and thus a nomen nudum not
intended as a
permanent scientific record (ICZN Article 8.1.1). The official
paper was published in Scientific Reports on April 26 2022. Note
while Aranciaga Rolando et al. (2022) refer to a distal metatarsal II
described by Novas et al.,
the latter mention no such element although it is possible Aranciaga
Rolando et al. meant the supposed dorsal transverse process end that
they
otherwise don't mention. The presence of exposed camellae shows
this
is clearly an axial element however, as Mesozoic theropod metatarsals
are not pneumatic. Aranciaga Rolando et al. used Novas'
tetanurine analysis to recover Maip as a megaraptorid in a polytomy
with Aerosteon, Orkoraptor and Tratayenia.
References- Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Briss�n-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, �lvarez-Herrera, Gentil, Bogan, Chimento,
Garc�a-Mars�, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019. Paleontological discoveries in the Chorrillo
Formation (upper Campanian-lower Maastrichtian, Upper Cretaceous),
Santa Cruz Province, Patagonia, Argentina. Revista del Museo Argentino
de Ciencias Naturales. 21(2), 217-293.
Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2021 online.
The biggest Megaraptoridae (Theropoda: Coelurosauria) of South America.
https://www.researchsquare.com/article/rs-1152394/v1 DOI: 10.21203/rs.3.rs-1152394/v1
Aranciaga Rolando, Motta, Agnolin, Manabe, Tsuihiji and Novas, 2022. A
large Megaraptoridae (Theropoda: Coelurosauria) from Upper Cretaceous
(Maastrichtian) of Patagonia, Argentina. Scientific Reports. 12:6318.
Orkoraptor Novas, Ezcurra and
Lecuona, 2008
O. burkei Novas, Ezcurra and Lecuona, 2008
Middle Campanian, Late Cretaceous
Cerro Fortaleza Formation, Santa Cruz, Argentina
Holotype- (MPM-Pv 3457) (~7 m) postorbital, quadratojugal (82 mm), coronoid(?),
eight teeth, atlantal intercentrum, fragmentary atlantal neurapophysis, eight
fragmentary ribs, two proximal (~3-4) caudal vertebrae (90 mm), three incomplete
chevrons, proximal tibia (~700 mm)
Paratype- (MPM-Pv 3458) three teeth
Referred- ?(MPM-Pv 10004) incomplete fibula (Ramirez and Bazcko, 2009)
(MPM-Pv coll.) several teeth (Lacovara et al., 2014)
Diagnosis- (modified after Novas et al., 2008) teeth with unserrated
and transversely wide mesial margins; teeth with a median depression flanked
by two longitudinal and narrow furrows on the lingual surface; quadratojugal
with a short jugal process.
Comments- This taxon was discovered in 2001 and announced in a 2004 abstract,
but not described until 2008. Varela (2011) revised the age and nomenclature
of the formation Orkoraptor was found in, which was later revised again by Novas et al. (2019). Contra Novas et al.'s (2008)
suggestion of a more distal placement for the caudals (~14-18), their central
proportions match those around caudals 3 and 4 of Neovenator. The illustrated
chevron is probably around the twentieth based on comparison to Allosaurus.
Originally, Novas et al. (2004) identified the coronoid as a nasal.
Ramirez and Bazcko (2009) described MPM-PV 10004 briefly in an abstract, stating
its posteriorly closed proximomedial fibular fossa and moderately developed
iliofibularis tubercle were similar to tetanurines. It was said to be less robust
than carcharodontosaurids, and most similar to megaraptorans and basal coelurosaurs
like tyrannosauroids, which given the recent referral of megaraptorans to Coelurosauria
may both be true. Finally, they noted it might be referrable to Orkoraptor
from the same formation, and indeed its size (incomplete but >639 mm) matches
that genus' estimated tibial length. Thus it is provisionally referred here.
Novas et al. (2004) originally suggested relationships with spinosaurids, Megaraptor
and undescribed megaraptoran MCF-PVPH 411. Once described, Novas et al. (2008)
found it to be a coelurosaur most probably related to compsognathids or dromaeosaurids
in their version of the TWG analysis. The postorbital is almost identical to
Aerosteon, a supposed carnosaur that also has proximal caudal pleurocoels.
Benson et al. (2010) found it to be a member of their newly formed carcharodontosaurid
clade Megaraptora, though with an uncertain position within that group. Recent
analyses by Novas et al. (2013) and Porfiri et al. (2014) which place Megaraptora
inside Tyrannosauroidea could work with all of these hypotheses.
References- Novas, Lecuona, Calvo and Porfiri, 2004. Un teropodo del
Cretacico Superior de la Provincia de Santa Cruz. Ameghiniana. 41(4), 59R.
Novas, Ezcurra and Lecuona, 2008. Orkoraptor burkei nov. gen. et sp.,
a large theropod from the Maastrichtian Pari Aike Formation, southern Patagonia,
Argentina. Cretaceous Research. 29, 468-480.
Ramirez and Bazcko, 2009. Material de Theropoda (Dinosauria) de la Formaci�n
Pari Aike (Cret�cico Superior), Santa Cruz, Argentina. Ameghiniana. 46(4S), 91R.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Varela, 2011. Sedimentolog�a y modelos deposicionales de la Formaci�n
Mata Amarilla, Cret�cico de la cuenca austral, Argentina. PhD thesis,
Universidad Nacional de La Plata. 287 pp.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Lacovara, Lamanna, Ibiricu, Poole, Schroeter, Ullmann, Voegele, Boles, Carter,
Fowler, Egerton, Moyer, Coughenour, Schein, Harris, Martinez and Novas, 2014.
A gigantic, exceptionally complete titanosaurian sauropod dinosaur from southern
Patagonia, Argentina. Scientific Reports. 4, 6196.
Porfiri, Novas, Clavo, Agnolin, Ezcurra and Cerda, 2014. Juvenile specimen of
Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation.
Cretaceous Research. 51, 35-55.
Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Briss�n-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, �lvarez-Herrera, Gentil, Bogan, Chimento,
Garc�a-Mars�, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019. Paleontological discoveries in the Chorrillo
Formation (upper Campanian-lower Maastrichtian, Upper Cretaceous),
Santa Cruz Province, Patagonia, Argentina. Revista del Museo Argentino
de Ciencias Naturales. 21(2), 217-293.
Tratayenia Porfiri, Juarez Valieri, Santos and Lamanna, 2018
T. rosalesi Porfiri, Juarez Valieri, Santos and Lamanna, 2018
Santonian, Late Cretaceous
Bajo de la Carpa Formation, Rio Colorado Subgroup, Neuquen, Argentina
Holotype- (MUCPv 1162)
(~7 m, subadult) incomplete sixth dorsal neural arch, partial seventh dorsal
vertebra, incomplete eighth dorsal vertebra (80 mm), ninth dorsal
vertebra (70 mm), tenth dorsal vertebra (80 mm), incomplete twelfth
dorsal vertebra (90 mm), thirteenth dorsal vertebra (100 mm), two
dorsal rib fragments, incomplete sacrum, partial ilium, distal pubic
fragment, partial ischium
Referred- ?(MAU-Pv-LI-548) two vertebrae (Mendez, Filippi and Garrido, 2015)
?(MCF-PVPH-399) distal metatarsal II (Coria and Arcucci, 2004)
?(MCF-PVPH-418) posterior dorsal centrum (110 mm) (Coria and Arcucci, 2004)
?(MCF-PVPH-654) (~9 m) tibia (~800 mm) (Coria, Currie, Paulina Carabajal, Garrido and Koppelhus, 2004)
Santonian, Late Cretaceous
Bajo de la Carpa Formation, Rio Colorado Subgroup, Rio Negro, Argentina
?(MPCA 247) tooth (~21.69x14.53x7.8 mm) (Gianechini, Lio and Apestegu�a, 2011)
Diagnosis- (after Porfiri et
al., 2018) middle and posterior dorsal vertebrae with slender, parallel
prezygodiapophyseal and paradiapophyseal laminae, both of which persist
throughout the series; ventral margin of prezygapophysis of middle and
posterior dorsal vertebrae subhorizontal in lateral view, meeting
anterior margin of prezygapophysis at a nearly right angle; middle
dorsal vertebrae with paired, ventrally diverging laminae extending
from the ventrolateral base of the anterior edge of the neural spine to
the base of the prezygapophysis, having the form of an inverted 'Y' in
anterior view; neural spines of posteriormost three sacral vertebrae
nearly twice as long anteroposteriorly as that of the first sacral
vertebra.
Comments- The holotype was discovered
in 2006 and initially reported by Porfiri et al. (2008) as "a new large
basal tetanuran taxon, possibly an allosauroid related to the
Carcharodontosauridae." Novas
et al. (2013) suggested it may be megaraptoran based on the "complex pneumatic
dorsal and sacral vertebrae with tall neural spines inclined forward." Porfiri et al. (2018) described this as a
megaraptorid using Novas et al.'s tetanurine analysis.
Coria and Arcucci (2004) described MCF-PVPH-399 and 418 as Theropoda
indet., suggesting they were basal tetanurines but noting a a lack of
carcahrodontosaurid apomorphies.
Baiano and Coria (2018) reanalyzed the material, finding "in light of
current knowledge, it would be possible to fine-tune the identification
of these elements as Megaraptora indet."
Coria et al. (2004) briefly mention MCF-PVPH-654 as referrable to (translated) "
a lineage of tetanurine theropods, for now comparable in size with the
carcharodontosaurids." It was later asssigned to ?Megaraptoridae
indet. by Lammana et al. (2020) who also reveal the specimen number.
Gianechini et al. (2011) described a tooth as closely resembling Orkoraptor
in "the absence of a mesial carina, the abrupt curvature of the apical
portion of the crown, the presence of wear facets, the distal denticle
density, and the figure-eight shaped basal section. However, MPCA 247
differs from Orkoraptor in the presence of wrinkles..."
Mendez et al. (2015) mention (translated) "two vertebral remains (MAU-Pv-LI-548) were also recovered from a member of the clade Megaraptora."
References- Coria and Arcucci, 2004. Nuevos dinosaurios ter�podos de Auca Mahuevo,
provincia del Neuqu�n (Cretácico tard�o, Argentina). Ameghiniana. 41,
597-603.
Coria, Currie, Paulina Carabajal, Garrido and Koppelhus, 2004. �Un
nuevo linaje de ter�podos en el Cretácico Tard�o de Argentina? XX
Jornadas Argentinas de Paleontolog�a de Vertebrados. 42R.
Porfiri, Calvo,
Ju�rez Valieri and Santos, 2008. A new large theropod dinosaur from the
Bajo de la Carpa Formation (Late Cretaceous) of Neuqu�n, Patagonia.
Actas III Congresso Latinoamericano de Paleontolog�a de Vertebrados.
R202.
Gianechini, Lio and Apestegu�a, 2011. Isolated archosaurian teeth from
"La Bonita" locality (Late Cretaceous, Santonian-Campanian), R�o Negro
Province, Argentina. Historia Natural, tercera serie. 1, 5-16.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Mendez, Filippi and Garrido, 2015. Nuevos hallazgos de dinosaurios teropodos
provenientes del sitto la invernada (Formacion Fajo de la Carpa), Rincon de
los Sauces, Neuquen. XXIX Jornadas Argentinas de Paleontolog�a de Vertebrados,
resumenes. Ameghiniana. 52(4) suplemento, 28-29.
Baiano and Coria, 2018. Revisiting theropod material from the Late
Cretaceous nesting site Auca Mahuevo and the possible record of a giant
megaraptoran. XXXII Jornadas Argentinas de Paleontolog�a de Vertebrados
y VII Jornadas T�cnicas de Paleontolog�a de Vertebrados, Libro de
Res�menes). R7.
Porfiri, Juarez Valieri, Santos and Lamanna, 2018. A new megaraptoran
theropod dinosaur from the Upper Cretaceous Bajo de la Carpa Formation
of northwestern Patagonia. Cretaceous Research. 89, 302-319.
Lamanna, Casal, Martinez and Ibiricu, 2020. Megaraptorid (Theropoda:
Tetanurae) partial skeletons from the Upper Cretaceous Bajo Barreal
Formation of central Patagonia, argentina: Implications for the
evolution of large body size in Gondwanan megaraptorans. Annals of
Carnegie Museum. 86(3), 255-294.
Tyrannoraptora Sereno, 1999
Definition- (Tyrannosaurus rex + Passer domesticus) (Holtz, Molnar and Currie, 2004; modified from Sereno, 1999)
References- Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Aristosuchia Seeley, 1892
Aristosuchus Seeley, 1887b
A. pusillus (Owen, 1876) Seeley, 1887b
= Poekilopleuron pusillus Owen, 1876
= Poekilopleuron minor Owen vide Cope, 1878
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (NHMUK R178) (~2 mm) first sacral vertebra (25 mm), second sacral
vertebra (29 mm), third sacral vertebra (24 mm), fourth sacral vertebra (23
mm), fifth sacral vertebra (21 mm), distal pubes
Paratypes- ....(NHMUK R178a) dorsal vertebra (21 mm)
....(NHMUK R178b) two incomplete distal caudal vertebrae (28 mm)
....(NHMUK R179) incomplete manual ungual
Comments- The type material was discovered in 1866 (Naish, 2002; Blows,
1983), though not described by Owen until a decade later as a new species of
Poekilopleuron. As Fox (in Blows, 1983) wrote the bones "were found
in the compass of a dinner plate", the paratypes probably belong to the
same individual as the holotype. Cope (1878) wrote "Owen has recently described
an English species of Laelaps under the name Poecilopleuron minor",
which might be a mistake as pusillus means 'very small'.
Lydekker (1888) referred an additional manual ungual (NHMUK R899) to the taxon,
but Naish (2002) notes it has a low and distally positioned flexor tubercle
unlike the paratype ungual so may be from another species. Galton (1973) referred
both the tibia NHMUK R186 and proximal femur NHMUK R5194 to Aristosuchus,
but neither can be compared to the type material and are more properly Coelurosauria
incertae sedis (though the tibia has been referred to Ornithomimosauria by Allain
et al., 2014). Naish (1999) described the tibia MIWG 5137 as possibly being
Aristosuchus. Hutt (2001) listed IWCMS 1995.208, MIWG 5823 and MIWG 5824
as Aristosuchus sp., but the specimens are undescribed though the vertebrae
may be comparable to the type specimens. Naish (2002) illustrated partial ischium NHMUK R6426 as a possible Aristosuchus specimen. Any of these specimens
may be referrable to Aristosuchus, which may also be synonymous with
Calamosaurus and/or Calamospondylus.
Jurcsak (1982) referred a cervical vertebra and caudal centrum from the Cornet
Bauxite of Romania to Aristosuchus sp., but there is no rationale for
this and the specimens cannot be compared to the type.
Traditionally considered a coelurid (Paul, 1988) or a compsognathid
(Naish et al., 2001), Hartman et al. (2019) were the first to analyze Aristosuchus quantitatively and found it emerges as a tyrannoraptoran excluded from Maniraptoriformes and tyrannosauroids as close to Tyrannosaurus as Juratyrant.
References- Owen, 1876. Monograph on the fossil Reptilia of the Wealden and Purbeck Formations. Supplement No. VII. Crocodilia (Poikilopleuron) and Dinosauria? (Chondrosteosaurus). [Wealden.] Palaeontographical Society
Monograph. 30, 1-7.
Cope, 1878. The affinities of the Dinosauria. The American Naturalist. 12, 57-58.
Seeley, 1887a. On Aristosuchus pusillus Ow., being further notes on the
fossils described by Sir R. Owen as Poikilopleuron pusillus, Ow. Geological
Magazine. 47, 234-235.
Seeley, 1887b. On Aristosuchus pusillus (Owen), being further notes on
the fossils described by Sir R. Owen as Poikilopleuron pusillus, Owen.
Quarterly Journal of the Geological Society of London. 43, 221-228.
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, London. 309 pp.
Seeley, 1892. Contribution to a knowledge of the Saurischia of Europe
and Africa. Quarterly Journal of the Geological Society of London. 48,
188-191.
Galton, 1973. A femur of a small theropod dinosaur from the Lower Cretaceous
of England. Journal of Paleontology. 47, 996-1001.
Jurcsak, 1982. Occurrences nouvelles des Sauriens mesozoiques de Roumanie. Vertebrata
Hungarica. 21, 175-184.
Blows, 1983. William Fox (1813-1881), a neglected dinosaur collector of the
Isle of Wight. Archives of Natural History. 11, 299-313.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
464 pp.
Naish, 1999. Studies on Wealden Group theropods - An investigation into the
historical taxonomy and phylogenetic affinities of new and previously neglected
specimens. Masters thesis, University of Portsmouth. 184 pp.
Hutt, 2001. Catalogue of Wealden Group Dinosauria in the Museum of Isle of Wight
Geology. In Martill and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological
Association. 411-422.
Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: Theropods. In Martill
and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association.
242-309.
Naish, 2002. The historical taxonomy of the Lower Cretaceous theropods (Dinosauria)
Calamospondylus and Aristosuchus from the Isle of Wight. Proceedings
of the Geologists' Association. 113, 153-163.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Allain, Vullo, Le Loeuff and Tournepiche, 2014. European ornithomimosaurs (Dinosauria,
Theropoda): An undetected record. Geologica Acta. 12(2), 127-135.
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
Bagaraatan Osmolska, 1996
B. ostromi Osmolska, 1996
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (ZPAL MgD-I/108) (~3 m; adult) (mandible ~230 mm) anterior
dentary, posterior mandible, sacral neural spine, twenty-five caudal vertebrae
(80, 65, 44, 37 mm), several chevrons, partial ilia, proximal pubis, proximal
ischium, proximal and distal femur (~315 mm), tibia (365 mm, 380 mm with tarsus),
fibula (~350 mm), astragalocalcaneum, pedal phalanx II-2 (37 mm), pedal phalanx
IV-1 (34 mm)
Diagnosis- antarticular present; lateral longitudinal ridge present on
proximal caudal prezygopophyses; two large fossae on the lateral postacetabular
surface; anterior and greater trochanters with minimal separation; accessory
trochanter; posterior trochanter present; tibiofibular crest powerfully developed;
tibia broader mediolaterally than long anteroposteriorly in proximal view; tibia,
fibula, astragalus and calcaneum fused.
Comments- The holotype was discovered in 1970 and initially announced
by Gradzinski and Jerzykiewicz (1972) as a "coeluroid dinosaur", but
not described until 1996 by Osmolska. She placed it in Avetheropoda and noted
resemblences to supposed Iren Dabasu avimimid femur PIN 2549-100, which has
been subsequently identified as troodontid. Csiki and Grigorescu (1998) remarked
on similarities between it, several European maniraptoriforms (Elopteryx,
Heptasteornis, Bradycneme) and supposed ceratosaur distal femur FGGUB R.351
which is now thought to be a hadrosaurid metatarsal. Holtz (2000) placed it
outside Tyrannoraptora, but more derived than Proceratosaurus, Ornitholestes,
Coelurus and Scipionyx. Longrich (2001) placed it in the Maniraptora
in his unpublished analysis, in a trichotomy with alvarezsaurids and avepectorans.
Coria
et al. (2002) refer it to the Troodontidae without discussion. Rauhut (2003)
found it to be a maniraptoran in a trichotomy with enigmosaurs and paravians.
Holtz (2004) resolves it as the basalmost tyrannosauroid. Carr (2005) found
it to be the sister taxon of Bistahieversor, both being sister to Tyrannosauridae,
based on cranial characters. When the hindlimb characters in Carr (2005) are
combined with these (personal observation), Bagaraatan is resolved as
an albertosaurine sister to Appalachiosaurus. Finally, Brusatte (2013)
strongly suspects Bagaraatan is a chimaera of tyrannosauroid and other
coelurosaurian elements, which will be the subject of a paper written by Makovicky,
himself and others. Based on general morphology and the above results, it may
end up that the mandible, vertebrae and pelvis are tyrannosauroid while the
hindlimb is troodontid.
References- Gradzinski and Jerzykiewicz, 1972. Additional geographical
and geological data from the Polish-Mongolian paleontological expedition. Palaeontologica
Polonica. 27, 17-30.
Osmolska, 1996. An unusual theropod dinosaur from the Late Cretaceous Nemegt
Formation of Mongolia. Acta Palaeontologica Polonica. 41, 1-38.
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.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. Gaia 15. 5-61.
Longrich, 2001. Secondarily flightless maniraptoran theropods? Journal of Vertebrate
Paleontology. 21(3), 74A.
Coria, Chiappe and Dingus, 2002. A new close relative of Carnotaurus sastrei
Bonaparte 1985 (Theropoda: Abelisauridae) from the Late Cretaceous of Patagonia.
Journal of Vertebrate Paleontology. 22, 460-465.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska. The Dinosauria
Second Edition. University of California Press. 861 pp.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special
reference to North American forms. PhD thesis, University of Toronto. 1170 pp.
Brusatte, 2013. The phylogeny of basal coelurosaurian theropods (Archosauria:
Dinosauria) and patterns of morphological evolution during the dinosaur-bird
transition. PhD thesis, Columbia University. 944 pp.
Beipiaognathus Hu, Wang and Huang,
2016
B. jii Hu, Wang and Huang, 2016
Barremian-Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Holotype- (AGB4997) (chimaera) (1.6 m) skull (~190 mm), mandible, eleven
cervical vertebrae, cervical ribs, thirteen dorsal vertebrae, dorsal ribs, gastralia,
thirty-nine caudal vertebrae, chevrons, scapula, coracoid, three humeri (109
mm), radii (98 mm), ulnae (100 mm), six carpals, metacarpals I (14 mm), phalanges
I-1 (45 mm), manual unguals I (37 mm), metacarpals II (59 mm), phalanges II-1
(49 mm), phalanges II-2 (49 mm), manual unguals II (40 mm), metacarpals III
(54 mm), phalanges III-1 (35 mm), phalanges III-2 (37 mm), phalanges III-3 (25
mm), manual unguals III, manual claw sheaths, ilium, pubis, femora (184 mm),
tibiae (230 mm), fibulae (223 mm), proximal tarsals, metatarsals I (22 mm),
phalanges I-1 (18 mm), pedal unguals I (14 mm), metatarsals II (120 mm), phalanges
II-1 (29 mm), phalanges II-2 (23 mm), pedal unguals II (18 mm), metatarsals
III (130 mm), phalanges III-1 (37 mm), phalanges III-2 (32 mm), phalanges III-3
(28 mm), pedal unguals III (25 mm), metatarsals IV (123 mm), phalanges IV-1
(25 mm), phalanges IV-2 (20 mm), phalanges IV-3 (16 mm), phalanges IV-4 (18
mm), pedal unguals IV (20 mm), pedal claw sheaths
Diagnosis- (after Hu et al., 2016; note the chimaerical nature makes
these doubtful) teeth unrecurved and unserrated; tail short with no more than
40 caudal vertebrae; forelimb long (fl/hl ratio 55%) due to long ulna (u/h ratio
92%); metacarpal I short and rectangular, ~24% length of metacarpal II; phalanx
II-1 is most robust and longest in manus; pedal digit III is longest, followed
by IV then II.
Comments- This specimen is clearly a chimaera (first suggested by Cau,
online 2016; published by Hartman et al., 2019) as three humeri are present. In addition, elements are generally
placed to look articulated even when anatomically incorrect- e.g. the scapulocoracoid
placed as a booted ischium, unguals I and II on the rightmost manus which are
each made from separate proximal and distal portions so that the flexor tubercle
is placed dorsally, pedal phalanx III-2 in both feet is upside down despite
being articulated. In the skull, it seems likely the frontal is actually a rectangular
bone ventral to the parietal, the premaxilla is actually what's labeled as the
dentary, and that the true dentary is what's labeled as the premaxilla and maxilla.
But given the nature of the specimen and teeth placed posterior to the parietal,
the composition and theropod identity of the supposed skull elements is in doubt.
The pubis is placed backward, and the phalanges are different lengths in each
manus and pes, with manual phalanges III-1 and III-2 having obscure articulations.
This makes the identification and association of any phalanges doubtful, so
that e.g. the microraptorian-like manual unguals II or apparently elongate pedal
phalanges IV-4 are unlikely to be correctly interpreted characters of whichever
basal coelurosaurs the scapulocoracoid, humerus and antebrachium, pubis and
metatarsus belong to. Of those elements, the humerus is very similar to Ornitholestes,
while the pubis resembles Coelurus. Beipiaognathus is thus placed
near to these taxa in the cladogram here, though additional study will be necessary
to the attribution of each element and which if any deserve lectotype status.
References- Cau, online 2016. http://theropoda.blogspot.com/2016/08/lo-status-paleontologico-di.html
Hu, Wang and Huang, 2016. A new species of compsognathid from the Early Cretaceous
Yixian Formation of western Liaoning, China. Journal of Geology. 40(2), 191-196.
Kakuru Molnar and Pledge, 1980
K. kujani Molnar and Pledge, 1980
Aptian, Early Cretaceous
Maree Formation, South Australia, Australia
'Plastoholotype'- (SAM P17926) tibia (~330 mm), fibular fragments
Referred- ?(SAM P18010) pedal phalanx (44 mm) (Molnar and Pledge, 1980)
Diagnosis- (after Molnar and Pledge, 1980) astragalar facet becoming
slender dorsally to a distinct apex, not broad enough to extend across width
of tibia at any point; astragalar facet limited medially by pronounced anterior
ridge that runs dorsally from medial mediolus; medial malleolus strongly projected
medially.
Comments- This taxon has been favorably compared to Avimimus by
Paul (1988) and Molnar (pers. comm. to Norman, 1990) based on the tibia's slender
proportions, but these are seen in many other small coelurosaurs as well. As
most coelurosaur tibiae still have proximal tarsals attached, comparison is
usually limited to the shape of the lateral and medial edges and of the astragalar
ascending process. In addition to Avimimus, such varied taxa as Garudimimus
and Achillobator approach the basic outline of Kakuru's tibia.
The height of Kakuru's ascending process is characteristic of tyrannoraptorans,
though megaraptorans and noasaurids approach it. The
width to depth ratio in distal view is at least as high as abelisauroids and
tetanurines, but lower than maniraptorans or Coelurus. Most coelurosaurs'
ascending processes are more extensive medially, except for Tugulusaurus
and basal ornithomimosaurs and alvreazsauroids. The strongly pointed ascending process is only
rivaled by Shuvuuia's and Garudimimus', though Kakuru lacks
the medial notch characteristic of alvarezsaurid and basal ornithomimosaur ascending
processes. Rauhut (2005) argued for an abelisauroid identity (echoed by Salisbury
et al., 2007) based on the anterior ridge extending vertically medial to the
ascending process, as seen in Quilmesaurus, Masiakasaurus and
Velocisaurus. However, Rauhut (2012) later recognized this is present
in numerous other theropods as well, including Chuandongocoelurus, Suchomimus
and several basal coelurosaurs. Agnolin et al. (2010) misunderstood Rauhut as
claiming Kakuru had a median ridge in the ascending process' facet, which
they noted is a taphonomic artifact in that taxon. Those authors placed Kakuru
in Averostra incertae sedis, though this can be narrowed
further as they correctly point out no ceratosaur has such a high ascending
process. Kakuru is retained as Tyrannoraptora incertae sedis here.
References- Molnar and Pledge, 1980. A new theropod dinosaur from South
Australia. Alcheringa. 4, 281-287.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster.
464 pp.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". In Weishampel,
Dodson and Osmolska (eds.). The Dinosauria. University of California
Press. 280-305.
Rauhut, 2005. Post-cranial remains of ‘coelurosaurs’ (Dinosauria,
Theropoda) from the Late Jurassic of Tanzania. Geological Magazine. 142(1), 97-107.
Salisbury, Agnolin, Ezcurra and Pias, 2007. A critical reassessment of the Creaceous
non-avian dinosaur faunas of Australia and New Zealand. Journal of Vertebrate
Paleontology. 27(3), 138A.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Rauhut, 2012. A reappraisal of a putative record of abelisauroid theropod dinosaur
from the Middle Jurassic of England. Proceedings of the Geologists' Association.
123(5), 779-786.
Mirischia Naish, Martill and
Frey, 2004
M. asymmetrica Naish, Martill and Frey, 2004
Albian, Early Cretaceous
Romualdo Formation of the Santana Group, Brazil
Holotype- (SMNK 2349 PAL) (~2.1 m; subadult) posterior twelfth dorsal
vertebra, thirteenth dorsal vertebra (26 mm), twefth dorsal rib, gastralia,
anterior synsacrum, partial ilia, pubes, incomplete ischia, incomplete femora
(165 mm), proximal tibia, proximal fibula, intestine, postpubic airsac(?)
Diagnosis- (modified from Naish et al., 2004) pubic peduncle of ilium
with concave cranial surface; pubic boot with no cranial expansion and 32% total
length of pubis; pedicular fossae located craniodorsal to neural canal on caudal
dorsal vertebra; distal tips of the neural spines between 63% and 67% longer
than their bases; ventral surface of sacral centra bearing shallow median depressions
at either end; extremely thin bone walls to all known elements.
Comments- This specimen was obtained from a private collector. Brusatte
(2013) determined that contra earlier authors, both the obturator fenestrae
of the pubes and ischia were originally closed, with those of the left pubis
and right ischium only appearing open due to damage.
This taxon was originally described as a compsognathid (Martill et al., 2000;
Naish et al., 2004) and found to be in that clade in Rauhut's (2003) analysis
(though unnamed at the time), based on anteroposteriorly expanded dorsal neural
spine apices and an elongate pubic boot with reduced cranial component. However,
these characters are common among other basal coelurosaurs, including basal
tyrannosauroids. Novas et al. (2012) and Brusatte et al. (2014) also recovered
this result, but it can move to Tyrannosauroidea in the latter with only 4 steps. Naish (online, 2006) noted Mirischia is similar to tyrannosauroids
in having an anteriorly concave pubic peduncle and referred the taxon to that
clade. Dal Sasso and Maganuco (2011) found it to be the most basal coelurosaur
in their version of Senter's TWiG analysis. Hartman et al. (2019) recovered Mirischia as closest to Guanlong
and coelurids, though tyrannosauroid characters were not extensively
sampled. Placing it in Compsognathidae required 6 more
steps.
References- Martill, Frey, Sues and Cruickshank, 2000. Skeletal remains
of a small theropod dinosaur with associated soft structures from the Lower
Cretaceous Santana Formation of northeastern Brazil. Canadian Journal of Earth
Sciences. 37(6), 891-900.
Rauhut. 2003. The interrelationships and evolution of basal theropod dinosaurs.
Special Papers in Palaeontology. 69, 1-213.
Naish, Martill and Frey, 2004. Ecology, systematics and biogeographical relationships
of dinosaurs, including a new theropod, from the Santana Formation (?Albian,
Early Cretaceous) of Brazil. Historical Biology. 16(2-4), 57-70.
Naish, 2006 online. http://darrennaish.blogspot.com/2006/06/basal-tyrant-dinosaurs-and-my-pet.html
Dal Sasso and Maganuco, 2011. Scipionyx samniticus (Theropoda: Compsognathidae)
from the Lower Cretaceous of Italy: Osteology, ontogenetic assessment, phylogeny,
soft tissue anatomy, taphonomy, and palaeobiology. Memorie della Societ�
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
281 pp.
Novas, Ezcurra, Agnolin, Pol and Ortiz, 2012. New Patagonian Cretaceous theropod
sheds light about the early radiation of Coelurosauria. Revista del Museo Argentino
de Ciencias Naturales. 14(1), 57-81.
Brusatte, 2013. The phylogeny of basal coelurosaurian theropods (Archosauria:
Dinosauria) and patterns of morphological evolution during the dinosaur-bird
transition. PhD thesis, Columbia University. 944 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(10), 2386-2392.
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
Timimus Rich and Vickers-Rich, 1994
T. hermani Rich and Vickers-Rich, 1994
Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Holotype- (NMV P186303) femur (430 mm)
Comments- This taxon was only briefly described by Rich and Vickers-Rich
(1994), and their photos of the femora are not very useful for determining morphology.
It was referred to Ornithomimosauridae [sic] within Ornithomimosauria, though
without explicit supporting evidence. The authors did list two characters to
distinguish ornithomimosaur femora from carnosaurs' though- anteroposteriorly
compressed head; anterior trochanter extends proximal to greater trochanter.
At least the latter is definitely present in Timimus, and is only known
in Appalachiosaurus plus tyrannosaurids, Aniksosaurus and
maniraptorans. It is absent in ornithomimosaurs, contra Rich and Vickers-Rich,
making it unlikely Timimus belongs to this clade. As for the compressed
femoral head, this is not significantly different from Allosaurus or
Tyrannosaurus in Sinornithomimus or Gallimimus, while it
is slightly anteroposteriorly compressed in Archaeornithomimus and proximodistally
compressed in Sinornithomimus. Rich and Vickers-Rich also list one character
to distinguish "ornithomimosaurids" from elmisaurids (presumably within
Ornithomimosauria, though elmisaurids are now known to be caenagnathid oviraptorosaurs)-
proximally placed anterior trochanter base. Yet the distally placed base in
the "elmisaurid" Chirostenotes is a misinterpreted accessory trochanter.
Thus there are no valid published reasons for referring Timimus to Ornithomimosauria
or Ornithomimidae. The authors diagnose Timimus solely on the basis of
lacking an extensor groove, presumably compared to Gallimimus which has
an autapomorphic closed groove. Yet other ornithomimosaurs (Garudimimus,
Sinornithomimus, Archaeornithomimus) lack extensor grooves, which
is similar to most maniraptoriforms, and Timimus actually has a groove.
Salisbury et al. (2007) state that Timimus is paravian and shares characters
with unenlagiine deinonychosaurs, which was elaborated on by Agnolin et al.
(2010).
Benson et al. (2012) noted Agnolin et al.'s supposedly dromaeosaurid-like
characters are absent- a fourth trochanter is present, and the anterior and
greater trochanters are separated by a deep notch, and that it differs from
maniraptorans in lacking an anteroposteriorly thick greater trochanter. Instead,
they believed it to be a tyrannosauroid closer to Tyrannosaurus than Guanlong, but not as derived as Xiongguanlong due to the shallow extensor groove (and I note it is also more shallow than in Juratyrant).
The tyrannosauroid identification is based on characters
supposedly differing from ornithomimosaurs- elevated femoral head;
anteroposteriorly narrower anterior trochanter; "accessory trochanter
forms a transversely thickened region" instead of being "prominent,
triangular." However, Gallimimus also has an elevated femoral head, with the angle in both taxa being less than derived tyrannosauroids. Garudimimus has a similarly narrow anterior trochanter relative to total femoral depth. Timimus actually has a more prominent accessory trochanter than e.g. Gallimimus or Archaeornithomimus, the trochanter of e.g. Archaeornithomimus is rounded too, and those of e.g. Gallimimus and Alioramus do not differ in transverse width. Thus I don't think the data strongly favors either option and leave Timimus as Tyrannoraptora incertae sedis.
Britt (1993) mentions NMV 186303 as a dromaeosaurid dorsal vertebra. This may
be a typo for NMV 186302, an oviraptorosaur dorsal vertebra described by Currie
et al. (1996). The smaller paratype femur NMV P186323 was found to be quite different by Benson
et al. and referred to Maniraptora. Rich and Vickers-Rich also mention "a
number of vertebrae" from Victoria they refer to ornithomimosaurs, but
do not list in the hypodigm of Timimus (one may be caudal vertebra NMV
P186168 that Benson et al. refer to Ornithomimosauria). These would not be likely
to belong to Timimus now that the genus has been shown to be more basal.
Currie et al. (1996) mention the pubis NMV P186058 and vertebrae from the type
locality and the Strzelecki Group. The pubis is probably a typo for NMV P186046
from the type locality, which was described by Benson et al. (2010) as a basal
tyrannosauroid, so may actually belong to Timimus after all. The lack
of details for the vertebrae is similar to Rich and Vickers-Rich's comment.
Finally, Pigdon (online 1997) mentions a possible ornithomimosaur ungual from
the Strzelecki Group found in 1996. This may be NMV P199085, a manual ungual
described by Benson et al. (2012) as being weakly curved with a low flexor tubercle
and flattened ventral surface which is broader than the dorsal surface. The
authors referred it to Theropoda indet., and there is no evidence it belonged
to Timimus.
References- Britt, 1993. Pneumatic postcranial bones in dinosaurs and
other archosaurs. PhD thesis, University of Calgary. 383 pp.
Rich and Vickers-Rich, 1994. Neoceratopsians and ornithomimosaurs: Dinosaurs
of Gondwana origin? National Geographic Research. 10(1), 129-131.
Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria)
specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia.
Alcheringa. 20(1-2), 73-79.
Pigdon, online 1997. http://home.alphalink.com.au/~dannj/timimus.htm
Chinsamy, Rich and Vickers-Rich, 1998. Polar dinosaur histology. Journal of
Vertebrate Paleontology. 18(2), 385-390.
Salisbury, Agnolin, Ezcurra and Pias, 2007. A critical reassessment of the Creaceous
non-avian dinosaur faunas of Australia and New Zealand. Journal of Vertebrate
Paleontology. 27(3), 138A.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Benson, Barrett, Rich and Vickers-Rich, 2010. A southern tyrant reptile. Science.
327, 1613.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from southern Australia
indicates high polar diversity and climate-driven dinosaur provinciality. PLoS
ONE. 7(5), e37122.
"Tonouchisaurus"
Anonymous?, 1994
"T. mongoliensis" Anonymous?, 1994
Middle-Late Albian, Early Cretaceous
Khuren Dukh, Khuren Dukh Formation, Mongolia
Material- (IGM coll.) (~1 m) humerus, radius, ulna, manus, femur, tibia, metatarsus,
pedal phalanges
Comments-
This specimen was originally announced in a Japanese newspaper article
in 1994 (Endo, DML 1994), and was found by the Joint Japan-Mongolia
paleontological expedition in Khuren Dukh, which would mean it was
discovered between July 29 and August 9 1993 (Watabe and Suzuki, 2000)
and housed at the IGM. Holtz (DML, 1994) stated from his examination of
the article and news footage it appeared to be a coelurosaur with a
didactyl manus and a non-arctometatarsalian metatarsus (if it was in
anterior view, which is uncertain). Olshevsky (DML, 1995) reported that
Barsbold (pers. comm., November 1995) stated the taxon is a basal
tyrannosauroid with didactyl manus and non-arctometatarsalian pes,
confirming Holtz's interpretations. He also reported Barsbold said the
description was in press, though it has yet to appear more than two
decades later. Both Holtz and Olshevsky have suggested the specimen may
be a juvenile, due to its small size. Barsbold (pers. comm., 2001)
stated the manus is actually tridactyl, while the metatarsus is "almost
not pinched", perhaps indicating a subarctometatarsal morphology.
While it may still be a tyrannosauroid, the lack of a didactyl manus
removes the only known reason for this assignment, though it should be
noted basal tyrannosauroids (e.g. Dilong,
Guanlong)
have tridactyl manus and closely resemble compsognathid/coelurid grade
coelurosaurs morphologically, including in pedal anatomy. A
subarctometatarsus would suggest a tyrannoraptoran assignment at
least. Molina-Perez and Larramendi (2019) have since published
the name as a nomen nudum.
References- Anonymous?, 1994. Japanese newspaper article.
Endo, DML 1994. https://web.archive.org/web/20201110013818/http://dml.cmnh.org/1994Dec/msg00059.html
Holtz, DML 1994. https://web.archive.org/web/20201109204928/http://dml.cmnh.org/1994Dec/msg00155.html
Olshevsky, DML 1995. https://web.archive.org/web/20201110061419/http://dml.cmnh.org/1995Nov/msg00158.html
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1993. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 19-29.
Molina-Perez and Larramendi, 2019. Dinosaur Facts and Figures: The
Theropods and Other Dinosauriformes. Princeton University Press. 288 pp.
Tyrannoraptora indet. (TMP online)
Middle-Late Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
Material- (TMP 1966.025.0020) tooth
Comments- TMP 1966.025.0020 is labeled as a Chirostenotes tooth on the TMP online catalogue, which indicates it is another kind of coelurosaur as caenagnathids are toothless.
undescribed probable Tyrannoraptora (Hone and Tanke, 2015)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Material- (TMP 1980.020.0173) tooth (TMP online)
(TMP 1994.143 coll.) (small) tibia, phalanx
Comments- TMP 1980.020.0173 is labeled as a Chirostenotes tooth on the TMP online catalogue, which indicates it is another kind of coelurosaur as caenagnathids are toothless.
Reference- Hone and Tanke, 2015. Pre- and postmortem tyrannosaurid bite
marks on the remains of Daspletosaurus (Tyrannosaurinae: Theropoda) from
Dinosaur Provincial Park, Alberta, Canada. PeerJ. 3:e885.
undescribed Tyrannoraptora (Fanti and Miyashita, 2009)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Material- (UALVP 52986) tooth (Fanti, Currie and Burns, 2015)
(UALVP 52594) tooth (Fanti, Currie and Burns, 2015)
(small) ungual (Fanti and Miyashita, 2009)
References- 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.
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.
undescribed Tyrannoraptora (Eberth and Currie, 2010)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
Material- (TMP 2000.045.0048) vomer
(TMP or UALVP coll.) (unassociated) three teeth, tibia, two metatarsals, phalanx,
two unguals
References- 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.
unnamed tyrannoraptoran (Hunt and Lucas, 2006)
Kimmeridgian-Tithonian
Morrison Formation, New Mexico, US
Material- (NMMNH P-26093) femur (388 mm), tibial fragments
Comments- Hunt and Lucas (2006) referred this to Coeluridae indet., citing
similarity to Tanycolagreus.
Reference- Hunt and Lucas, 2006. A small theropod dinosaur from the Upper
Jurassic of eastern New Mexico with a checklist of small theropods from the
Morrison Formation of western North America. New Mexico Museum of Natural History
and Science Bulletin. 36, 115-118.
unnamed probable tyrannoraptoran (Langston, 1974)
Early Albian, Early Cretaceous
Paluxy Formation of the Trinity Group, Texas, US
Material- (SMU 62723) manual ungual
Comments- This was discovered with the Astrophocaudia holotype.
Langston (1974) illustrated it and referred the ungual to Ornithomimidae, correctly
distinguishing it from Microvenator and Deinonychus. The slender
shape, low curvature and proximally placed flexor tubercle seem most similar
to caenagnathids.
References- Langston, 1974. Nonmammalian Comanchean tetrapods. Geoscience
and Man. 8, 77-102.
unnamed probable Tyrannoraptora (Gallup, 1975)
Aptian-Middle Albian, Early Cretaceous
Trinity Group, Texas
Material- (FMNH PR 975) femur (365 mm)
(FMNH 2-51#1) seven teeth (4.4, 4.7, 6, 6.5, 7, 7.7, 8 mm), seven tooth fragments
( mm)
(FMNH 3-51#1) two teeth (9.5, 12 mm)
(FMNH 11s-52#1) juvenile premaxillary tooth (2 mm), two lateral teeth (5, 10.6
mm)
(FMNH 47-50) tooth (7.3 mm)
(FMNH 202-50) tooth (8.9 mm)
(FMNH Turtle Gully) two teeth (2, 11 mm), pedal phalanx IV-3 (12 mm), ungual
?IV (25 mm)
Comments- The teeth were referred to Coeluridae by Gallup (1975), are
recurved and have 29-50 serrations per 5 mm distally and an equal number mesially
when present. The femur was referred to Ornithomimus sp., but resembles
Nedcolbertia in anteroposterior aspect and robusticity. Similarly, the
pedal phalanx and ungual were referred to Ornithomimus sp. but probably
belong to more babal coelurosaurs instead.
Reference- Gallup, 1975. Lower Cretaceous dinosaurs and associated vertebrates
from north-central Texas in the Field Museum of Natural History. MS thesis,
University of Texas at Austin. 159 pp.
undescribed tyrannoraptoran (Button, Zanno and Makovicky, 2014)
Cenomanian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- partial femoral shaft, incomplete tibia, incomplete metatarsal
IV, phalanx IV-2, phalanx IV-4
Comments- Button et al. (2014) describe a coelurosaurian hindlimb discovered
in 2012 which is arctometatarsalian and gracile. They state "Metatarsal
IV most resembles Coelurus (YPM 2010) from the Upper Jurassic Morrison
Formation in general proportion, mediolateral compression of the distal aspect,
and near absence of a lateral collateral ligament pit, yet is unique in possessing
an obliquely oriented groove marking the extensor surface and a dorsally bulbous
distal condyle." Coelurus is generally found to be surrounded by
nonarctometatarsalian taxa though, which could indicate this Mussentuchit taxon
shows some coelurids convergently evolved an arctometatarsus, or that similarities
to Coelurus are homoplasious and the new taxon belongs to an arctometatarsalian
clade.
Reference- Button, Zanno and Makovicky, 2014. New coelurosaurian theropod
remains from the Upper Cretaceous Mussentuchit Member of the Cedar Mountain
Formation, Central Utah. Journal of Vertebrate Paleontology. Program and Abstracts
2014, 101.
undescribed Tyrannoraptora (Osborn, 1916)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (AMNH 974) teeth, phalanges
(AMNH 5014) twelve caudal vertebrae
(AMNH 5015) phalanx III-2, phalanx III-3
(AMNH 5019; lost) manual ungual
Comments- Osborn (1916) questionably referred this material to Ornithomimus
velox, but Russell (1972) noted none contained ornithomimid material. They
may belong to other coelurosaurs instead.
References- Osborn, 1916. Skeletal adaptation of Ornitholestes,
Struthiomimus, Tyrannosaurus. Bulletin of the American Museum
of Natural History. 35, 733-771.
Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada.
Canadian Journal of Earth Sciences. 9(4), 375-402.
undescribed Tyrannoraptora (Estes, 1964)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming
Material- (UCMP coll.) teeth
Comments- Originally referred to cf. Dryptosaurus sp. by Estes
(1964), based on provenance these are probably not Dryptosaurus and
may be Tyrannosaurus or dromaeosaurid instead.
Reference- Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance
Formation, eastern Wyoming. University of California Publications in Geological
Sciences. 49, 1-180.
unnamed Tyrannoraptora (Hunt and Lucas, 1993)
Late Santonian, Late Cretaceous
Cleary Coal Member of the Menefee Formation, New Mexico, US
Material- (NMMNH coll.) tooth fragments (Hunt and Lucas, 1993)
Comments- Hunt and Lucas (1993) stated "theropods are represented by tooth fragments"
Reference- Hunt and Lucas, 1993. Cretaceous vertebrates of New Mexico. In Lucas
and Zidek (eds.). Vertebrate Paleontology in New Mexico. New Mexico
Museum of Natural History and Science Bulletin. 2, 77-91.
unnamed Tyrannoraptora (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-30327) tooth
(NMMNH P-52508) tooth
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the
Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications
for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
unnamed tyrannoraptoran (McDonald, Wolfe and Dooley Jr, 2018)
Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
Material- (UMNH VP 28348 in part)
phalanx IV-2 (30 mm)
Comments- The supposed pedal phalanx IV-4 of Dynamoterror is far too small to be referred to the
same individual as the holotype and may be a phalanx IV-2 based on its elongation,
either ornithomimid or tyrannosauroid.
Reference- McDonald, Wolfe and Dooley Jr, 2018. A new tyrannosaurid (Dinosauria:
Theropoda) from the Upper Cretaceous Menefee Formation of New Mexico.
PeerJ. 6:e5749.
unnamed tyrannoraptoran (Williamson and Brusatte, 2014)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (NMMNH P-36545) tooth
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the
Late Cretaceous of the San Juan Basin, northwestern New Mexico and their implications
for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
unnamed tyrannoraptoran (Wang, Cau, Wang, Yu, Wu, Wang and Liu, 2023 online)
Early Aptian, Early Cretaceous
Pigeon Hill, Longjiang Formation, Inner Mongolia, China
Material- (LY 2022JZ3005) incomplete metatarsal I, incomplete metatarsals II, incomplete metatarsals III, incomplete metatarsals IV
Comments- This was discovered
in 2022. Wang et al. (2023) state "The middle shaft of metatarsal
Ⅲ is mediolaterally compressed as in several maniraptoriforms, although
not to the extreme degree seen in arctometatarsalian taxa", which
leaves Tyrannosauroidea and basal Maniraptoriformes. While the
authors propose "Metatarsal I is extremely reduced, and located close
to the distal end of metatarsal II", the proximal and distal ends of
the metatarsus are missing so that this cannot be measured. Wang
et al. "tentatively referred [this] to Coelurosauria" based on the
subarctometatarsaly.
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 tyrannoraptoran (Hu, 1963)
Late Valanginian-Early Albian, Early Cretaceous
Jehol Group, Liaoning, China
Material- (IVPP V2757) partial cervical vertebra (35 mm)
Comments- Assigned to ?Coeluridae indet. by Hu (1963), this is probably
a tyrannoraptoran due to its amphicoelous and elongate (~1.8 longer than tall)
centrum. It has a ventral keel.
Reference- Hu, 1963. [The carnivorous dinosaurian remains from Fusin,
Liaoning]. Vertebrata PalAsiatica. 7, 174-176.
Coeluridae Marsh, 1881
= Coeluria Marsh, 1881
= Coeluroidea Marsh, 1881 sensu Nopcsa, 1928
Definition- (Coelurus fragilis <- Proceratosaurus bradleyi,
Tyrannosaurus rex, Allosaurus fragilis, Compsognathus longipes, Ornithomimus
edmontonicus, Deinonychus antirrhopus) (Hendrickx, Hartman and Mateus, 2015)
Comments- Traditionally a wastebasket family for small Jurassic and Cretaceous theropods, a reduced version including Coelurus and Tanycolagreus
has been recovered in most modern analyses. While it is clearly
close to the origins of Tyrannoraptora, the most recent analyses are
ambiguous as to what branch of that clade it belongs to. Senter
et al.'s (2012) and Brusatte et al.'s (2014) TWiG analyses recover it
as the basalmost family of tyrannosauroids, but it can be moved to
basal Maniraptoromorpha in both analyses using only two extra
steps. Hartman et al. (2019) recover them as maniraptoromorphs
but do not deeply sample tyrannosauroid characters, though it only
takes 4 steps to move them to the latter clade. Cau (2018)
uniquely finds the family divided, with Tanycolagreus the basalmost tyrannosauroid and Coelurus
the basalmost maniraptoromorph. Hartman et al.'s matrix does
reject the sometimes proposed placement in Maniraptora though (e.g.
Gauthier, 1986), which requires 13 more steps to enforce.
References- Marsh, 1881. A new order of extinct Jurassic reptiles (Coeluria).
American Journal Science. 21, 339-341.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Gauthier, 1986. Saurischian monophyly and the origin of birds. Memoirs of the Californian Academy of Sciences 8, 1-55.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids
(Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the
evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body
plan culminated in rapid rates of evolution across the dinosaur-bird
transition. Current Biology. 24(20), 2386-2392.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25.
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
Zuolong Choiniere, Clark, Forster
and Xu, 2010
= "Zuolong" Choiniere, 2010
Z. salleei Choiniere, Clark, Forster and Xu, 2010
= "Zuolong salleei" Choiniere, 2010
Early Oxfordian, Late Jurassic
Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Holotype- (IVPP V15912) (~3.1 m; ~35 kg; subadult) incomplete skull (~353
mm), premaxillary tooth, angular, two lateral teeth, partial axial neural arch,
incomplete third cervical vertebra (~77 mm), incomplete fourth cervical vertebra
(~82 mm), incomplete fifth cervical vertebra (~85 mm), incomplete eighth cervical
vertebra, incomplete ninth cervical vertebra, partial tenth cervical neural
arch, two dorsal centra, two fragmentary dorsal centra (~82 mm), incomplete
sacrum, first caudal neural arch, second caudal neural arch, third caudal centrum,
incomplete fourth caudal vertebra, three incomplete mid caudal vertebrae, two
mid caudal centra, mid caudal neural arch, incomplete scapula, incomplete humerus
(~155 mm), radius (137 mm), incomplete ulna, distal phalanx I-1, incomplete
manual ungual I, partial ilium, incomplete pubes, femora (one distal; 336 mm),
partial tibia, proximal fibula, partial phalanx I-1, pedal ungual I, metatarsal
II (191.9 mm), phalanx II-1, metatarsal III (224.3 mm), partial metatarsal IV
(~201.7 mm)
Diagnosis- (after Choiniere et al., 2010) large, slit-like quadrate foramen
inclined medially at approximately 45 degrees with associated deep fossa on
the quadrate; sacral centrum 5 with an obliquely oriented posterior articular
surface that is angled anterodorsally; fovea capitis very large, occupying almost
the entire posterodorsal surface of the femoral head; distal condyle of metatarsal
III large relative to that of other metatarsals and bearing an anteromedially
projecting flange on its anteromedial margin.
Comments- This specimen was discovered in 2001 and announced in an abstract
by Clark et al. (2002) as a basal coelurosaur. It was later described in more
detail in an abstract by Choiniere et al. (2008), who used a version of the
TWG matrix and found it to be one of the most basal coelurosaurs, sister to
Tugulusaurus. Choiniere et al. (2010) named and described the taxon in
depth, although several months earlier, the description and name appeared in Choiniere's (2010) thesis.
Choiniere et al. (2010) recovered it either as a non-tyrannoraptoran coelurosaur in a trichotomy
with Tugulusaurus
or as a non-maniraptoriform maniraptoromorph. Brusatte et al. (2014)
found it to be sister to Tyrannoraptora (it appears in a polytomy in
their Figure 1 due to Tugulusaurus'
variable position), while Cau (2018) uiniquely recovered it as the most
basal tetanurine. While not shown due to non-maniraptoromorph
characters being poorly sampled, Zuolong
falls out in Coeluridae in the analysis of Hartman et al. (2019). It
is tentatively placed there on this site, pending more extensive future
analyses.
References- Clark, Xu, Forster, Wang and Andres, 2002. New small dinosaurs
from the Upper Jurassic Shishugou Formation at Wucaiwan, Xinjiang, China. Journal
of Vertebrate Paleontology. 22(3), 44A.
Choiniere, Clark, Xu and Forster, 2008. A new basal coelurosaur from the upper
Shishugou Formation (Xinjiang, People's Republic of China). Journal of Vertebrate
Paleontology. 28(3), 63A.
Choiniere, 2010. Anatomy and systematics of coelurosaurian theropods from the
Late Jurassic of Xinjiang, China, with comments on forelimb evolution in Theropoda.
PhD Thesis. George Washington University. 994 pp.
Choiniere, Clark, Forster and Xu, 2010. A basal coelurosaur (Dinosauria: Theropoda)
from the Late Jurassic (Oxfordian) of the Shishugou Formation in Wucaiwan, People's
Republic of China. Journal of Vertebrate Paleontology. 30(6), 1773-1796.
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.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25.
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
Coelurus Marsh, 1879
C. fragilis Marsh, 1879
= Coelurus agilis Marsh, 1884
= Elaphrosaurus agilis (Marsh, 1884) Russell, Beland and McIntosh, 1980
Middle-Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Utah, Salt Wash Member of the
Morrison Formation, Brushy Basin Member of the Morrison Formation?, Wyoming,
US
Syntypes- (YPM 1991) proximal caudal vertebra (35 mm), proximal caudal
vertebra, proximal caudal centrum, proximal caudal neural arch
....(YPM 1992) eight mid caudal vertebrae (33 mm), partial mid caudal centrum
....(YPM 1993) fourth cervical vertebra (~53 mm), proximal caudal neural arch
Referred- (UMNH 7795) humerus (Carpenter et al., 2005)
?(UUVP 11743) humerus (Carpenter et al., 2005)
?(YPM 1933) tooth (Marsh, 1896)
(YPM 1994) caudal centrum (Ostrom, 1980)
(YPM 1995) caudal vertebra, fragments (Ostrom, 1980)
(YPM 2010; holotype of Coelurus agilis) (subadult) partial dentary, fifth
cervical vertebra (~55 mm), seventh cervical vertebra (~50 mm), eighth cervical
vertebra (~49 mm), first dorsal vertebra (~45 mm), second or third dorsal neural
arch, fifth dorsal vertebra (~34 mm), sixth dorsal neural arch, seventh dorsal
neural arch, eighth dorsal vertebra (~39 mm), ninth dorsal neural arch, tenth
dorsal vertebra (~47 mm), eleventh dorsal vertebra (~44 mm), incomplete twelfth
dorsal vertebra (~47 mm), thirteenth dorsal neural arch, two indeterminate neural
arches, proximal caudal vertebra, proximal scapula, humerus (119 mm), radii
(one proximal) (~81 mm), ulnae (91, 96 mm), distal carpal I, proximal metacarpal
I, phalanx I-1 (~39 mm), metacarpals II (~56 mm), phalanges II-1 (~47 mm; one
incomplete), phalanges II-2 (~55 mm; one incomplete), phalanx III-1 (~16 mm),
phalanx III-2 (~18 mm), distal phalanx III-3, ilial fragment, pubes, femora
(one proximal) (~210 mm), distal tibia, proximal fibula, astragalus (32 mm wide),
distal metatarsal III, metatarsal IV (196 mm), fragments (Ostrom, 1980)
?(YPM 9162) partial sacral vertebra (Marsh, 1884)
?(YPM 9163; not 1252, contra Welles and Long, 1974) astragalus (74 mm wide)
(Welles and Long, 1974)
Diagnosis- (modified from Carpenter et al., 2005) very gracile dentary;
paired pleurocoels on some cervical centra; triangular cervical transverse processes
angled sharply ventrolaterally; pubic foot very acuate ventrally, projected
posterodorsally; interpubic fenestra located at midlength of pubic symphysis;
metatarsus subequal to femur in length.
Comments- YPM 1994, 1995, 2010 and possibly 9162 belong to the syntype
individual (Ostrom, 1980), as they are comparable in size (contra Marsh, 1884),
do not contain duplicated elements and are from the same part of the same quarry.
Thus, Coelurus agilis is an objective junior synonym of Coelurus fragilis.
YPM 9163 was described by Welles and Long, and matches the astragalus of YPM
2010 except for its size. Carpenter et al. (2005) state they may be different
specimens, or Welles and Long could have misreported YMP 9163's size. However,
Ostrom (1980) reports YPM 9163 is from Quarry 9, which would prove it's a different
specimen. Based on stratigraphy, it may belong to the unnamed (?)enigmosaur
of Makovicky (1997) instead.
YPM 1933 is from Quarry 12, and its referral to Coelurus fragilis by
Marsh (1896) is unfounded. It may belong to Ornitholestes, Tanycolagreus
or another small theropod.
Makovicky (1995) and Carpenter et al. (2005) both list two complete dorsal vertebrae,
five centra and six arches. Yet Carpenter et al. illustrate eleven total arches,
which even considering one is attached to the centrum and two are the indeterminate
arches also listed, leaves two extra illustrated arches. Similarly, Carpenter
et al. illustrate a total of six dorsal centra, which means one is unillustrated.
In addition, only four cervical vertebrae are listed among the Coelurus
specimens, yet five are illustrated because Carpenter et al. illustrate dorsal
1 as a cervical. Ostrom (1980) mentions a second cervical vertebrae in YPM 1993,
which he believes was combined with the other cervical to create a composite
Marsh (1881) illustrated. Carpenter et al. (2005) concluded Marsh combined a
Coelurus
cervical with either YPM 1996 or 1997 (belonging to Makovicky's 1997
possible enigmosaur) to create the composite, but this conflicts with
Ostrom's statement. The vertebra illustrated by Marsh (1881) as a
dorsal is a proximal caudal (YPM 1991). Ostrom (1980) lists a
metacarpal III and metacarpal IV fragment as being present, but
Carpenter et al. illustrate a proximal metacarpal I, and a fragment of
a much thinner element. The latter resembles a distal phalanx III-3
most. Carrano (1998) lists femoral and tibial measurements for
"YPM 1991?" as 220.0 and 255.0 respectively, near certainly meaning YPM
2010, but the tibia is missing its proximal end.
Several vertebrae referred to Coelurus fragilis by Gilmore (1920) were
provisionally referred to the unnamed coelurosaur described by Makovicky (1997)
by Carpenter et al. (2005). A partial skeleton was referred to Coelurus
by Miles et al. (1998), also prompting them to refer a manus (AMNH 587) previously
referred to Ornitholestes to Coelurus. However, the skeleton was
later made the holotype of Tanycolagreus topwilsoni by Carpenter et al.
(2005) and the manus was referred to that species instead. A pubis referred
to Coelurus by Gilmore (1920) was also referred to Tanycolagreus
by Carpenter et al. (2005).
Gilmore (1920) doubted the accuracy of the three characters used by Osborn (1903)
to distinguish Coelurus from Ornitholestes, which led to many
synonymizing them until Ostrom (1980) properly differentiated the genera. His
preliminary analysis was confirmed once both Coelurus and Ornitholestes
were redescribed in detail by Carpenter et al. (2005).
References- Marsh, 1879. Notice of new Jurassic reptiles. American Journal
Science. 18, 501-505.
Marsh, 1881. A new order of extinct Jurassic reptiles (Coeluria). American Journal
Science. 21, 339-341.
Marsh, 1884. Principle characters of American Jurassic dinosaurs. Part 8: the
Order Theropoda. American Journal Science. 27, 29-40.
Marsh, 1896. The dinosaurs of North America. Sixteenth Annual Report of the
U.S. Geological Survey. p. 133-230.
Osborn, 1903. Ornitholestes hermanni, a new compsognathid dinosaur from
the Upper Jurassic. American Museum of Natural History Bulletin. 19, 459-464.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. Bulletin of the United States National Museum. 110, 1-154.
Welles and Long, 1974. The tarsus of theropod dinosaurs: Annals of the South
African Museum. 44, 117-155.
Ostrom, 1980. Coelurus and Ornitholestes: Are they the same? In
Jacobs (ed.). Aspects of Vertebrate History. Museum of Northern
Arizona Press. 245-256.
Russell, Beland and McIntosh, 1980. Paleoecology of the dinosaurs of Tendaguru
(Tanzania). M�moires de la Soci�t� g�ologique de
France. 139, 169-175.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). M.S. thesis, University of Copenhagen. 311pp.
Makovicky, 1997. A new small theropod from the Morrison Formation of Como Bluff,
Wyoming. Journal of Vertebrate Paleontology. 17(4), 755-757.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD thesis, The University of Chicago. 424
pp.
Miles, Carpenter and Cloward, 1998. A new skeleton of Coelurus fragilis
from the Morrison Formation of Wyoming. Journal of Vertebrate Paleontology. 18(3) 64A.
Carpenter, Miles, Ostrom and Cloward, 2005. Redescription of the small maniraptoran
theropods Ornitholestes and Coelurus from the Upper Jurassic Morrison
Formation of Wyoming. In Carpenter (ed.). The Carnivorous Dinosaurs. Indiana
University Press. 49-71.
Tanycolageus Carpenter,
Miles and Cloward, 2005
= "Tanycolagreus" Carpenter and Miles vide Anonymous, 2001
T. topwilsoni Carpenter, Miles and Cloward, 2005
?= Elaphrosaurus "philtippettensis" Pickering, 1995b
?= Elaphrosaurus "philtippettorum" Pickering, 1995a
= "Tanycolagreus topwilsoni" Carpenter and Miles vide Anonymous, 2001
Middle Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Wyoming, US
Holotype- (TPII 2000-09-29) (subadult; ~3.3 m) partial skull (premaxilla,
partial nasal, lacrimal, postorbital, squamosal fragment, quadratojugal, quadrate),
premaxillary tooth, two lateral teeth, splenial, articular, two anterior dorsal
centra, four posterior dorsal vertebrae (42, 44, 43, 51 mm), fourteen ribs,
gastralia fragments, first sacral centrum (40 mm), two proximal caudal centra,
two mid caudal centra, three distal caudal vertebrae, seven chevrons, scapulae
(281 mm), coracoid (scapulocoracoid 287 mm), humeri (198 mm), radii (143 mm),
ulnae (152 mm), radiale, semilunate carpal, metacarpal I (37 mm), phalanx I-1
(68 mm), manual ungual I (90 mm straight), metacarpal II (81 mm), phalanx II-1
(65 mm), phalanx II-2 (75 mm), manual ungual II (~55 mm), metacarpal III (55
mm), phalanx III-3 (40 mm), manual ungual III (39 mm straight), distal pubes,
femora (356 mm), tibiae (387 mm), fibulae (one proximal) (370 mm), astragalus
(47 mm wide), calcaneum, metatarsal I (50 mm), phalanx I-1 (40 mm), pedal ungual
I (~32 mm), metatarsal II (196 mm), phalanx II-1 (64 mm), phalanx II-2 (55 mm),
pedal ungual II (~45 mm), metatarsal III (216 mm), phalanx III-1 (73 mm), phalanx
III-2 (55 mm), phalanx III-3 (41 mm), pedal ungual III (57 mm), metatarsal IV
(202 mm), phalanx IV-1 (47 mm), phalanx IV-2 (41 mm), phalanx IV-3 (34 mm),
phalanx IV-4 (28 mm), pedal ungual IV (~45 mm), metatarsal V (50 mm)
Paratype- (AMNH 587) (~2.3 m) metacarpal II (58 mm), phalanx II-1 (40
mm), phalanx II-2 (48 mm), manual ungual II, metacarpal III (44 mm), phalanx
III-1 (14 mm), phalanx III-2 (17 mm), phalanx III-3 (31 mm), manual ungual III
(33 mm), metacarpal IV (9 mm) (Osborn, 1916)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US
Referred- ?(USNM 5737; intended holotype of Elaphrosaurus "philtippettensis"
and "philtippettorum") distal pubes (Gilmore, 1920)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Utah, US
Paratype- (UUVP 2999) (~6.3 m) premaxilla (32 mm) (Madsen, 1974)
Diagnosis- (after Carpenter et al., 2005) short, deep-bodied premaxilla
that is pierced by narial foramen at the base of the nasal process; orbital
process on the postorbital; T-shaped quadratojugal; centrodiapophyseal lamina
on dorsals.
Comments- The first remains of this species were originally referred
to Ornitholestes (Osborn, 1916), Coelurus (Gilmore, 1920) and
Stokesosaurus (Madsen, 1974). The holotype was collected in 1995 and
initially thought to be Coelurus
(Miles et al.,1998). Its name was first published in a guide to the
North American Museum of Ancient Life, credited to Carpenter and Miles.
Assigned to Coeluridae without supporting synapomorphies by Carpenter
et al. (2005), it does indeed fall out as a coelurid in almost all
published analyses such as Brusatte et al. (2014). The genus
could be synonymous with Stokesosaurus clevelandi,
whose holotype and referred ilia, and referred braincase cannot be compared
to Tanycolagreus.
The distal pubes USNM 5737 were discovered in 1884 and provisionally referred
to Coelurus agilis by Gilmore in 1920 based on their size. Pickering
(1995a) listed the name Elaphrosaurus philtippettorum in an unpublished
bibliographic manuscript. In that same year, Pickering printed a packet with
a description of the taxon as ?Elaphrosaurus philtippettensis,
indicating USNM 5737 is the intended type. Both variants on the name are nomina
nuda however, as he didn't follow ICZN Article 8.1.3- it must have been produced
in an edition containing simultaneously obtainable copies by a method that assures
numerous identical and durable copies. Pickering also referred USNM 8414 (two
metatarsals) and 8415 (a humerus) without justification. However, there are
no characters in the diagnosis except that it shares a straight humerus with
Elaphrosaurus and abelisaurids (which does not involve the intended type),
and the only characters listed in the description are those which distinguish
USNM 8415 from Dryosaurus. It is therefore also a nomen nudum in that
it lacks "a description or definition that states in words characters that
are purported to differentiate the taxon." Pickering will also describe
the species in his in progress work Mutanda Dinosaurologica. Carpenter et al.
(2005) referred USNM 5737 to their new taxon Tanycolagreus because of
its straight ventral edge and dorsally placed interpubic fenestra, unlike Coelurus.
Additionally, Ornitholestes lacks an interpubic fenestra altogether.
Why Pickering referred USNM 5737 to Elaphrosaurus is unknown, as he does
not discuss the specimen (except to note it is "elongate, ... lacking a
crest on its craniodorsal surface. In lateral view, the distal foot is ventrally
convex.") and E. bambergi does not preserve the distal pubis. Furthermore,
other ceratosaurs like Ceratosaurus, Kryptops and Carnotaurus
have a very distally placed interpubic fenestra, so USNM 5737 is probably not
a ceratosaur. Carpenter et al.'s assignment is here retained, though it should
be noted Juratyrant also has a distally flat pubic boot and proximally
placed interpubic fenestra.
References- Osborn, 1916. Skeletal adaptations of Ornitholestes,
Struthiomimus and Tyrannosaurus. Bulletin of the American Museum
of Natural History. 35, 733-771.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum, with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. Bulletin of the United States National Museum. 110,
1-154.
Madsen, 1974. A new theropod dinosaur from the Upper Jurassic of Utah. Journal
of Paleontology. 48, 27-31.
Pickering, 1995a. Jurassic Park: Unauthorized Jewish Fractals in Philopatry.
A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola,
California. 478 pp.
Pickering, 1995b. An extract from: Archosauromorpha: Cladistics and osteologies.
A Fractal Scaling in Dinosaurology Project. 2 pp.
Miles, Carpenter and Cloward, 1998. A new skeleton of Coelurus fragilis
from the Morrison Formation of Wyoming. Journal of Vertebrate Paleontology.
18(3), 64A.
Anonymous, 2001. North American Museum of Ancient Life guidebook.
Carpenter, Miles and Cloward, 2005. New small theropod from the Upper Jurassic
Morrison Formation of Wyoming. In Carpenter (ed.). The Carnivorous Dinosaurs.
Indiana University Press. 23-48.
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.
Maniraptoromorpha Cau, 2018
Definition- (Vultur gryphus <- Tyrannosaurus rex) (Cau, 2018)
References- Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25.
Sinocalliopteryx Ji, Ji,
Lu and Yuan, 2007
S. gigas Ji, Ji, Lu and Yuan, 2007
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Beds of the Yixian Formation, Liaoning, China
Holotype-
(JMP-V-05-8-01) (2.37 m) incomplete skull (290 mm), incomplete
mandibles, eleven cervical vertebrae, cervical ribs, twelve dorsal
vertebrae, dorsal ribs, twelve rows of gastralia, forty-nine caudal
vertebrae, chevrons, scapulae, coracoids, humeri, radii (100.7 mm),
ulnae, scapholunare, distal carpal I, distal carpal II, distal carpal
III, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx
II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1,
phalanx III-2, phalanx III-3, manual ungual III, ilium, pubes, ischia,
femora (~236 mm), tibiae, fibulae, astragalus, calcaneum, distal tarsal
III, distal tarsal IV, pes (mtIII 147.3 mm), feathers, four gastroliths
(15-20 mm)
Referred- (CAGS-IG-T1) (~3.1 m) partial skull, fragmentary dentary, six
dorsal vertebrae, eight dorsal ribs, gastralia, eighteen caudal vertebrae, thirteen
chevrons, radius (118.6 mm), phalanx I-1, metacarpal II, phalanx II-1, phalanx
II-2, manual ungual II, metacarpal III, partial manual ungual, manual claw sheath,
ischia (one distal), metatarsal I, phalanx I-1, pedal ungual I, metatarsals
II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsals III (206.3 mm),
phalanges III-1, phalanges III-2, phalanges III-3, pedal ungual III, metatarsals
IV, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, feathers (Xing
et al., 2012)
Comments- Ji et al. (2007) described this as a
compsognathid, and it does emerge as one in most TWiG analyses such as
Brusatte et al. (201) and Senter et al. (2012). However, Cau
(2018) recovers it as the sister to Tyrannoraptora, while Hartman et
al. (2019) find it to be a maniraptoromorph just stemward of
compsognathids. While it can move to Compsognathidae in the
latter matrix with only 4 more steps, most suggested character data was
utilized. Interestingly, only one step moves it to be more
closely related to basal tyrannosauroids like Dilong and proceratosaurids.
References- Ji, Ji, Lu and Yuan, 2007. A new giant compsognathid dinosaur
with long filamentous integuments from Lower Cretaceous of Northeastern China.
Acta Geologica Sinica. 81(1), 8-15.
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.
Xing, Bell, Persons, Ji, Miyashita, Burns, Ji and Currie, 2012. Abdominal contents
from two large Early Cretaceous compsognathids (Dinosauria: Theropoda) demonstrate
feeding on confuciusornithids and dromaeosaurids. PLoS ONE. 7(8), e44012.
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.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25.
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
Huaxiagnathus Hwang, Norell,
Ji and Gao, 2004
= "Huaxiasaurus" Anonymous, 2000
H. orientalis Hwang, Norell, Ji and Gao, 2004
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Beds of Yixian Formation, Liaoning, China
Holotype-
(CAGS-IG02-301) (~1.6 m; subadult) skull (165 mm), mandible, hyoid,
nine cervical vertebrae, cervical ribs, thirteen dorsal vertebrae,
dorsal ribs, gastralia, sacrum, first caudal vertebra (20.44 mm),
second caudal vertebra (21.66 mm), third caudal vertebra (20.46 mm),
fourth caudal vertebra (20.16 mm), fifth caudal vertebra (19.74 mm),
sixth caudal vertebra (19.67 mm), seventh caudal vertebra (21.32 mm),
eighth caudal vertebra (19.60 mm), ninth caudal vertebra (20.71 mm),
tenth caudal vertebra (20.12 mm), eleventh caudal vertebra (20.66 mm),
twelfth caudal vertebra (20.92 mm), thirteenth caudal vertebra (20.73
mm), fourteenth caudal vertebra (20.88 mm), fifteenth caudal vertebra
(22 mm), sixteenth caudal vertebra (21.75 mm), seventeenth caudal
vertebra (22.72 mm), eighteenth caudal vertebra (21.75 mm), nineteenth
caudal vertebra (21.43 mm), twentieth caudal vertebra (22.52 mm),
twenty-first caudal vertebra (23.15 mm), twenty-second caudal vertebra
(22.38 mm), twenty-third caudal vertebra (20.93 mm), twenty-fourth
caudal vertebrae (22 mm), twenty-fifth caudal vertebra (23.29 mm),
twenty-two chevrons, scapulae, coracoids, partial furcula, humeri (90
mm), radii (51 mm), ulnae (55 mm), scapholunares, distal carpals I,
distal carpals II, distal carpals III, metacarpals I (19 mm), phalanges
I-1 (38 mm), metacarpals II (40 mm), phalanges II-1 (26 mm), phalanges
II-2 (35 mm), manual unguals II, metacarpals III (26 mm), phalanges
III-1, phalanges III-2, phalanges III-3, manual unguals III, ilium
(~139 mm), pubes, ischia, femur (163 mm), tibiae (183 mm), fibula,
astragali, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I,
metatarsals II (91 mm), phalanges II-1, phalanges II-2, pedal unguals
II, metatarsals III (102 mm), phalanges III-1, phalanges III-2,
phalanges III-3, pedal unguals III, metatarsals IV (91 mm), phalanx
IV-1, phalanx IV-2, phalanx IV-3, phalanges IV-4, pedal unguals IV,
metatarsal V, stomach contents
Referred- (NGMC 98-5-003; "Huaxiasaurus") (~1.8 m) partial
skeleton including fragmentary skull, dorsal vertebrae, caudal vertebrae, humerus,
radius, ulna, distal carpals, metacarpals, manual unguals, fragmentary pelvis,
femur (~167 mm), tibiae, distal tarsals, metatarsal I, metatarsal II, metatarsal
III, metatarsal IV, metatarsal V (Anonymous, 2000)
Comments- "Huaxiasaurus" was first announced in 2000 in news
articles as a genus of bird. The specimen was later mentioned by Hwang et al.
(2001) in an abstract, and described briefly by Hwang et al. (2004). It is poorly
reconstructed and prepared, with many elements placed in the wrong position.
Hwang et al. (2004) tentatively referred it to their new genus Huaxiagnathus,
as the morphology is identical with the holotype except for a shorter skull
(34% of femoral length instead of 45%). It may be an older individual.
Hwang et al.'s (2004) phylogenetic analysis recovered Huaxiagnathus
as a basal compsognathid, which has also occured in future versions of
the TWiG analysis, such as Senter et al. (2012) and Brusatte et al.
(2014). Hartman et al. (2019) finds it just stemward of
compsognathids, in a clade with Juravenator, but both can be placed in that family with only a single added step.
References- Anonymous, 2000. Feathered dinosaurs on show in Hong Kong.
Xinhua News Agency, May 1.
Anonymous, 2000. New discovery to help solve riddle of bird origin. July 23.
Hwang, Norell, Gao and Ji, 2001. New information on Jehol theropods. Journal
of Vertebrate Paleontology. 21(3), 64A.
Hwang, Norell, Ji and Gao, 2004. A large compsognathid from the Early Cretaceous
Yixian Formation of China. Journal of Systematic Palaentology. 2(1), 13-30.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids
(Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the
evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body
plan culminated in rapid rates of evolution across the dinosaur-bird
transition. Current Biology. 24(20), 2386-2392.
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
Juravenator Gohlich and Chiappe,
2006
J. starki Gohlich and Chiappe, 2006
Late Kimmeridgian, Late Jurassic
Painten Formation, Germany
Holotype- (JME Sch 200; Borsti) (~75-80 cm; juvenile) skull (82 mm), sclerotic
ring, mandible (~77 mm), seven cervical vertebrae, cervical ribs, thirteen dorsal
vertebrae, dorsal ribs, gastralia, three sacral centra, forty-four caudal vertebrae,
chevrons, scapulae (42 mm), coracoids, clavicles, humeri (27, 27.5 mm), radii
(~19.3 mm), ulnae (20.5, 20.5 mm), metacarpals I (4.5 mm), phalanges I-1 (10.5
mm), manual unguals I (~10 mm), metacarpals II (11.5 mm), phalanx II-1 (8 mm),
phalanges II-2 (10, 10 mm), manual unguals II (9, 10 mm), metacarpals III (9
mm), phalanges III-1 (4 mm), phalanges III-2 (4.5, 4.5 mm), phalanges III-3
(5.5 mm), manual unguals III (5.5, 7 mm), manual claw sheaths, ilia (40 mm),
partial pubes, partial ischium?, femora (52 mm), tibiae (58.1, 58.1 mm), fibulae
(55.3, 56 mm), astragali, calcaneum, metatarsals I (4.6, 4.5 mm), phalanges
I-1 (5.8, 6 mm), pedal unguals I (6, 3.5 mm), metatarsals II (26.5 mm), phalanges
II-1 (10.4 mm), phalanges II-2 (9, 8 mm), pedal unguals II (10.7, 11.5 mm),
metatarsals III (34 mm), phalanges III-1 (11.9, 11.5 mm), phalanges III-2 (8.1,
8 mm), phalanges III-3 (7.4 mm), pedal unguals III (7.4, 6.6 mm), metatarsals
IV (29.6, 29.8 mm), phalanges IV-1 (7.4, 7 mm), phalanges IV-2 (5.5, 6.5 mm),
phalanges IV-3 (4.5 mm), phalanges IV-4 (4.2, 4 mm), pedal unguals IV (7.2,
5.8 mm), metatarsals V (8, 6.8 mm), scale impressions, feathers, caudal musculature
impressions
Diagnosis- (modified from Gohlich and Chiappe, 2006) large skull (1.5
times femoral length); eight maxillary teeth; no premaxillary–maxillary
diastema; posterior serrations on premaxillary teeth; concave rostral margin
of the jugal process of the postorbital; relatively long scapula with narrowest
portion at neck; proportionally short feet; antorbital fenestra subequal in
length to the orbit (ontogenetic?); abbreviated deltopectoral crest of the humerus
(ontogenetic?); proximally high manual claws that taper abruptly at midpoint;
bow-like zygapophyses in mid-caudal vertebrae.
Comments- Discovered in 1998, the holotype was given the
informal name of Borsti in 2001 news reports. It was described briefly
by Gohlich and Chiappe (2006), then in more detail by Gohlich et al.
(2006) and Chiappe and Gohlich (2011). Though usually recovered as a
basal coelurosaur (e.g. in Compsognathidae by Brusatte et al., 2014),
Rauhut and Foth (2014) noted it has characters which could suggest a
more basal position- two pairs of cervical pleurocoels; brevis shelf
continuous with supraacetabular crest; well-developed
antitrochanter. Hartman et al. (2019) finds it just stemward of
compsognathids, in a clade with Huaxiagnathus, but both can be placed in that family with only a single added step.
References- Viohl, 1999. Discovery of a new small theropod. Archaeopteryx.
17, 15-19.
Gohlich and Chiappe, 2006. A new carnivorous dinosaur from the Late Jurassic
Solnhofen archipelago. Nature. 440, 329-332.
Gohlich, Tischlinger and Chiappe, 2006. Juravenator starki (Reptilia,
Theropoda) ein neuer Raubdinosaurier aus dem Oberjura der Sudlichen Frankenalb
(Suddeutschland): Skelettanatomie und Weichteilbefunde. Archaeopteryx. 24, 1-26.
Chiappe and Gohlich, 2011. Anatomy of Juravenator starki (Theropoda:
Coelurosauria) from the Late Jurassic of Germany. Neues Jahrbuch f�r Geologie
und Pal�ontologie - Abhandlungen. 258(3), 257-296.
Rauhut and Foth, 2014. New information on the theropod dinosaurs from the Late
Jurassic lithographic limestones of southern Germany. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 212.
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
Neocoelurosauria Hendrickx, Mateus, Ara�jo and Choiniere, 2019
= "Neocoelurosauria" Hendrickx, 2015
Definition- (Compsognathus longipes + Passer domesticus) (modified after Hendrickx,
Mateus, Ara�jo and Choiniere, 2019)
Comments- The earliest use of
this name is online in the Dinosaur Mailing List archives (Kinman, DML
2001), although Gardner (DML 2001) first gives it context in a
cladogram where Maniraptoriformes should be. The term never
gained popularity and went unused for over a decade. Hendrickx
(2015) uses 'Neocoelurosauria' in apostrophes in cladograms in his
thesis for a clade of compsognathids plus maniraptoriforms, while Paul
(2016) uses it informally for "coelurosaurs more derived than
tyrannosauroids". This makes Paul's version another term for
Cau's 2018 Maniraptoromorpha. Alosnso et al. (2017) and Young et
al. (2019) both use neocoelurosaur informally, referring to Hendrickx's
thesis. Hendrickx et al. (2019) officially proposed
Neocoelurosauria, defining it as "the clade Compsognathidae +
Maniraptoriformes", which can be more or less inclusive than
Maniraptoromorpha depending on the topology.
References- Gardner, DML 2001. https://web.archive.org/web/20201110061215/http://dml.cmnh.org/2001Aug/msg00967.html
Kinman, DML 2001. https://web.archive.org/web/20160806135318/http://dml.cmnh.org/2001May/msg01044.html
Hendrickx, 2015. Evolution of teeth and quadrate in non-avian Theropoda (Dinosauria: Saurischia), with the description of Torvosaurus remains from Portugal. PhD thesis, Universidade Nova de Lisboa. 646 pp.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press.
360 pp.
Alonso,
Canudo, Torcida Fernandez-Baldor and Huerta, 2017. Isolated theropod
teeth associated with sauropod remains from El Oterillo II (Early
Cretaceous) site of Salas de los Infantes (Burgos, Spain). Journal of
Iberian Geology. 43(2), 193-215.
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.
Young, Hendrickx, Challands, Foffa, Ross, Butler and Brusatte, 2019.
New theropod dinosaur teeth from the Middle Jurassic of the Isle of
Skye, Scotland. Scottish Journal of Geology.
"Beelemodon" Bakker, 1997
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Wyoming, US
Material- (TATE 546) (~1.5-4 m) tooth (7.1 mm long, FABL 5.4 mm)
(TATE coll.) tooth (~9 mm)
Diagnosis- Currently indeterminate pending more detailed comparison to
several theropod taxa.
Description- This taxon is still a nomen nudum, as it is not yet
diagnosed, nor does it have a species name. Bakker describes it as an "omnivorouscarnivorous
dinosaur of uncertain relations" and an "enigmatic dinosaur".
It is supposedly "coyote-to-wolf size". Although using tooth size
to determine total length is extremely risky, comparison to various theropods
indicates a length of 1.5-4 meters is probable, depending on body form. It is
unclear whether postcranial remains can be referred to the taxon, as only teeth
are described and illustrated. A single tooth is illustrated in side view and
cross section. Another tooth is plotted in the "denticle-width vs. crown
height" graph, indicating a slightly larger specimen is known as well.
The illustrated tooth is slightly recurved, laterally compressed (50% as wide
as anteroposteriorly long) and missing its distal tip. Fluting is present on
the illustrated side. The root is constricted, the mesial carina lacks serrations
and the distal carina has serrations extending to the base. The serrations are
small (4.3 per mm, ~35 on the whole crown), pointed and project slightly distally.
The cross section indicates it was fairly symmetrical labiolingually, narrowing
anteriorly and exhibiting a slight mesial expansion labially(?) and a slight
distal expansion lingually(?).
Comments- At first glance, these specimens look very similar to ornithischian
premaxillary teeth. The posterior two premaxillary teeth of Lesothosaurus
have mesial serrations, but lack them distally except at the tip. This is the
reverse of the case in "Beelemodon". The serrations are comparatively
larger (~15 per tooth if they extended as basally as in "Beelemodon")
and do not extend to the base of the crown. Drinker has a very similar
tooth morphology, with serrations present only on the distal carina. These serrations
are slightly larger (25-30 per tooth) and have longer interdenticle slits. The
tooth itself is not recurved, but is otherwise similar in shape. Galtonia
also has similarily shaped teeth, but with larger serrations and mesial serrations
present apically. "Beelemodon" is obviously based on theropod maxillary
or dentary teeth however, as the premaxillary teeth of most theropods have serrations
displaced so that the distance between them is much longer labially than lingually.
Troodontids, tyrannosaurids and ornithischians have premaxillary teeth that
not only have the latter character, but are also much wider labiolingually than
"Beelemodon". The cross section of "Beelemodon" is very
similar to theropod maxillary and dentary teeth.
While "Beelemodon" is theropod, placing it within that clade is a
more difficult task. Among Jurassic and Cretaceous taxa the constricted root is known in Tanycolagreus, Proceratosaurus, Compsognathus
and most maniraptorans except dromaeosaurids and unenlagiines.
Unserrated mesial carinae and serrated distal carinae are present in
megaraptorans, Guanlong, basal maniraptoromorphs, some troodontids, Caihong and some dromaeosaurids. Thus it is most likely either compsognathid or troodontid. Compsognathus
has some teeth that have unserrated mesial carinae and serrated distal carinae.
These have larger serrations relative to crown height (20-25 per tooth). They
are shaped similarily and have similar serration morphology. Given the amount of variation
in serration number in a single theropod genus (Allosaurus- 20-35; Saurornitholestes-
15-35), there is no way to separate "Beelemodon" from Compsognathus
at this point. Because of this, it must remain indeterminate. The fluting or
serration morphology may eventually prove diagnostic, but this cannot be determined
from the available literature. I recommend classifying "Beelemodon"
as a provisionally indeterminate maniraptoromorph nomen nudum until further research
is done.
Reference- Bakker, 1997. Raptor family values: Allosaur parents brought
great carcasses into their lair to feed their young. In Wolberg, Sump and Rosenberg
(eds.). Dinofest International, Proceedings of a Symposium, Academy of Natural
Sciences. 51-63.
Microvenator? "chagyabi"
Zhao, 1986
= Microvenator "chagyaensis" Zhang and Li, 1997
Early Cretaceous
Lura Formation, Tibet, China
Material- (IVPP coll.) specimen including teeth
Comments- Discovered in 1976 (An et al., 2021), this specimen was first reported by Zhao (1983) who while discussing
the evolution of dinosaurs in China noted "the teeth of coelurosaurids
(Microvenator Ostrom) gradually disappeared and became few in number"
in the Early Cretaceous. He earlier noted the "tooth is thinnest and no
serration occurs" in Early Cretaceous coelurosaurs. As Ostrom did not refer
any teeth to Microvenator, it might be concluded Zhao was referring to
the teeth of an undescribed Chinese specimen of Microvenator. This idea
is strengthened by the later mention of a new Microvenator species from
the same deposits as other Early Cretaceous taxa Zhao mentions (Monkonosaurus,
?Asiatosaurus, ?Prodeinodon). As with other new Tibetan taxa listed
by Zhao (1983), it was probably supposed to be described by Zhao in the published
version of his doctoral dissertation "The Mesozoic vertebrate remains of
Xizang (Tibet), China", in the second Palaeontology of Xizang volume. Yet
this volume is only referenced by Zhao (1983; which was submitted in September
1981) and seems never to have been printed, though the previous volume was published
by the IVPP in 1980 and the third by the NIGP in 1981. Olshevsky (DML, 1999)
notes the IVPP rejected the paper as unpublishable. Zhao (1986) reported Microvenator
chagyabi from the Loe-ein Formation. It was later mentioned by Zhang and
Li (1997) as Microvenator chagyaensis from the Laoran Formation of Qamdun,
Zhag'yab County, Tibet. It is near certainly the same specimen listed as ?Coelurosauria
indet. by Weishampel et al. (2004) from the Lura Formation of Xizang Zizhiqu.
As the specimen has never been described or illustrated, it is a nomen nudum.
If it indeed has serrationless teeth, it is probably a coelurosaur at least as crownward as compsognathids. It may even
be a basal oviraptorosaur like Microvenator celer, as the Early Cretaceous
Chinese taxa Incisivosaurus, Protarchaeopteryx and Caudipteryx
have serrationless teeth as well. However, there is still no published evidence
for this or its generic referral.
Chure and McIntosh (1989) accidentally use the combination Microvenator dayensis,
presumably caused by confusion with the sauropod "Microdontosaurus dayensis".
References- Zhao, "1983" [unpublished]. The Mesozoic vertebrate
remains of Xizang (Tibet), China. The Series of the Scientific Expeditions to
the Qinghai-Xizang Plateau. Palaeontology of Xizang. 2, 1-200.
Zhao, 1983. Phylogeny and evolutionary stages of Dinosauria. Acta Palaeontologica
Polonica. 28(1-2), 295-306.
Zhao, 1986. The Cretaceous biota of China: Reptilia. in Hao, Su, Yu, Li, Li,
Wang, Qi, Guan, Hu, Liu, Yang, Ye, Shou, Zhang, et al. (eds.). The Cretaceous
System of China. Stratigraphy of China. 12, 67-73, plates XI, XII.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the
Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Zhang and Li, 1997. Mesozoic dinosaur localities in China and their stratigraphy.
In Wolberg, Sump and Rosenberg (eds.). Dinofest International, Proceedings of
a Symposium sponsered by Arizona State University. A Publication of The Academy
of Natural Sciences. 265-273.
Olshevsky, DML 1999. https://web.archive.org/web/20200720012936/http://dml.cmnh.org/1999Nov/msg00507.html
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004.
Dinosaur Distribution. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria:
Second Edition. University of California Press. 517-606.
An, Wang, Li, Wang and Wang, 2021. New discovery of Jurassic dinosaur
fossils in Chaya area, Qamdu district, Tibet. Geological Bulletin of
China. 40(1), 189-193.
Aniksosaurus Martinez and Novas,
2006
= "Aniksosaurus" Martinez et al. vide Anonymous, 1997
A. darwini Martinez and Novas, 2006
= "Aniksosaurus darwini" Tronfi, online 2000
Cenomanian, Late Cretaceous
Lower Bajo Barreal Formation, Chubut, Argentina
Holotype- (MTD-PV 1/48) (~2 m) femur, tibia, incomplete fibula, partial
metatarsal I, phalanx I-1 (16 mm), pedal ungual I (15 mm), metatarsal II (98
mm), phalanx II-1 (31 mm), phalanx II-2 (21 mm), metatarsal III (124 mm), phalanx
III-1 (33 mm), phalanx III-2 (30 mm), metatarsal IV (105 mm), phalanx IV-1 (22
mm), phalanx IV-2 (16 mm), phalanx IV-3 (15 mm)
Paratypes- (MTD-PV 1/1) partial tibia
(MTD-PV 1/2) incomplete tibia
(MTD-PV 1/3) femur (247 mm)
(MTD-PV 1/4) metatarsal
(MTD-PV 1/5) partial ilium
(MTD-PV 1/6) fragmentary dorsal vertebra
(MTD-PV 1/7) vertebra
(MTD-PV 1/8) vertebra
(MTD-PV 1/9) vertebra
(MTD-PV 1/10) partial tibia
(MTD-PV 1/11) fragment
(MTD-PV 1/12) fragment
(MTD-PV 1/13) mid caudal vertebra (40 mm)
(MTD-PV 1/14) partial posterior cervical vertebra
(MTD-PV 1/15) vertebra
(MTD-PV 1/16) incomplete humerus (~130 mm)
(MTD-PV 1/17) incomplete ulna (~104 mm)
(MTD-PV 1/18) fragmentary dorsal vertebra
(MTD-PV 1/19) fragment
(MTD-PV 1/20) fragment
(MTD-PV 1/21) neural arch
(MTD-PV 1/22) incomplete tibia
(MTD-PV 1/23) incomplete femur
(MTD-PV 1/24) partial ilium
(MTD-PV 1/25) fragment
(MTD-PV 1/26) incomplete femur
(MTD-PV 1/27) incomplete femur
(MTD-PV 1/28) partial tibia
(MTD-PV 1/29) partial humerus
(MTD-PV 1/30) neural arch
(MTD-PV 1/31) fragment
(MTD-PV 1/32) proximal caudal vertebra (34 mm)
(MTD-PV 1/33) partial ilium
(MTD-PV 1/34) tibia (250 mm)
(MTD-PV 1/35) partial ilium
(MTD-PV 1/36) partial humerus
(MTD-PV 1/37) partial humerus
(MTD-PV 1/38) fragment
(MTD-PV 1/39) fragment
(MTD-PV 1/40) incomplete manual ungual I (44 mm)
(MTD-PV 1/41) partial ischium (~160 mm)
(MTD-PV 1/42) partial humerus
(MTD-PV 1/43) phalanx
(MTD-PV 1/44) partial tibia
(MTD-PV 1/45) metatarsal
(MTD-PV 1/46) neural arch
(MTD-PV 1/47) vertebra
(MTD-PV 1/52) vertebra
(MTD-PV coll.) fragmentary ribs
Diagnosis- (modified after Martinez and Novas, 2006) cervical vertebrae
with the neural arch pedicels unusually deep (2.5 times the height of the centrum);
wide neural canal on cervical vertebrae (also in Avimimus); transversely
broad manual ungual I (also in Alvarezsauridae); caudolateral surface of proximal
femur with strong depression and rugosities presumably for the attachment for
M. ischiotrochantericus; metatarsal IV and its correspondent digit transversely
narrow (also in Nqwebasaurus).
Comments- The holotype and paratypes represent at least five individuals,
based on the number of right tibiae.
Discovered in 1995 and originally mentioned as a nomen nudum in an Argentinian
newpaper article, with the discovery attributed to Martinez et al.. In 1997,
the taxon was briefly described (but still not named) in an abstract by Martinez
and Novas. Tronfi (online 2000) wrote "To the list of discoveries was added, in 1995, aniksosaurus darwini 130 kilometers north of Sarmiento." Its formal description was finally published
in 2006.
Phylogenetic relationships- In its description, Martinez and Novas (2006) proposed Aniksosaurus
was a coelurosaur outside Maniraptoriformes, the latter containing
tyrannosaurids in their phylogeny. While not included in a
phylogenetic analysis, they did list three characters supporting this
placement. "Distal tibia with astragalar surface proportionally
low" is also true in basal tyrannosauroids (Coelurus, Guanlong) and maniraptoromorphs (Aorun, Compsognathus, alvarezsauroids including Nqwebasaurus).
"Insertion of the M. caudifemoralis longus extensive and deep" is
plesiomorphic for maniraptoromorphs, also being found in Ornitholestes, Juravenator
and all ornithomimosaurs. Robust limb elements were stated to
resemble carnosaurs and megalosauroids more than most coelurosaurs,
specifically the humerus, ulna, femur, tibia and pes. The humerus
is also robust in compsognathids and alvarezsaurids, while the
metatarsus is less robust than Ornitholestes, therizinosaurs or Deinocheirus.
However, any measure of basic robusticity such as femoral circumference
should also account for size correlation as larger taxa are typically
less gracile than smaller taxa.
Although Dal Sasso and Maganuco (2011) show Aniksosaurus as sister to Nedcolbertia
outside Tyrannoraptora in their Figure 113, this is based on a majority
consensus and the taxon can actually fall out as a tyrannosauroid or
maniraptoromorph instead. Similarly, in Choiniere et al. (2010)
it is in a polytomy with tyrannosauroids, compsognathid-grade taxa,
ornithomimosaurs and maniraptorans. Novas et al. (2012) recovered
it as the sister taxon to Maniraptoriformes. Brusatte (2013:395)
stated "it is unclear if it possesses any clear coelurosaurian
characters" so did not include it in his TWiG analysis. However,
as outlined above every analysis including Aniksosaurus
has recovered it within Coelurosauria, and only unquantified
appendicular robusticity has even been suggested to validly support a
more rootward placement than Tyrannoraptora. Most recently,
Hartman et al. (2019) recovered it in a polytomy with Scipionyx,
compsognathids and maniraptoriforms. While there are some
alvarezsauroid-like characters (e.g. large presacral neural canals*,
ventrally keeled proximal caudal centra, transversely broad manual
ungual I*, laterally expanded brevis shelf*, distally projecting
lateral femoral condyle), enforcing this result in the Hartman et al.
matrix adds 9 steps. However, the asterisked characters were not
included, so it could potentially only be 6 steps longer.
References- Anonymous, 1997. [title] Pagina/12. [pp]
Martinez and Novas, 1997. A new tetanuran (Dinosauria: Theropoda) from
the Bajo Barreal Formation (Upper Cretaceous), Patagonia. XIII Jornadas
Argentinas de Paleontologia de Vertebrados, resumenes. Ameghiniana.
34(4), 538.
Tronfi, online 2000. https://web.archive.org/web/20010216191134/http://tierraaustral.com/informacion/nota_paleo.htm
Mart�nez and Novas, 2006. Aniksosaurus darwini gen. et sp. nov.,
a new coelurosaurian theropod from the early Late Cretaceous of central Patagonia,
Argentina. Revista del Museo Argentino de Ciencias Naturales. 8(2), 243-259.
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal
alvarezsauroid theropod from the early Late Jurassic of Xinjiang,
China. Science. 327, 571-574.
Dal Sasso and Maganuco, 2011. Scipionyx samniticus (Theropoda: Compsognathidae)
from the Lower Cretaceous of Italy: Osteology, ontogenetic assessment, phylogeny,
soft tissue anatomy, taphonomy, and palaeobiology. Memorie della Societ�
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
281 pp.
Novas, Ezcurra, Agnolin, Pol and Ortiz, 2012. New Patagonian Cretaceous theropod
sheds light about the early radiation of Coelurosauria. Revista del Museo Argentino
de Ciencias Naturales. 14(1), 57-81.
Brusatte, 2013. The phylogeny of basal coelurosaurian theropods
(Archosauria: Dinosauria) and patterns of morphological evolution
during the dinosaur-bird transition. PhD thesis, Columbia University.
944 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
Archaeornithoididae Elzanowski and Wellnhofer, 1992
Archaeornithoides Elzanowski
and Wellnhofer, 1992
A. deinosauriscus Elzanowski and Wellnhofer, 1992
Late Campanian, Late Cretaceous
Djadokhta Formation, Mongolia
Holotype- (ZPAL MgD-II/29) (juvenile) (skull ~50 mm) maxillae, maxillary
teeth, anterior jugal, possible vomer fragment, palatine, possible ectopterygoid
fragment, possible parasphenoid, dentaries, dentary tooth
Comments- Discovered in 1965, this specimen was originally mentioned
by Elzanowski (1983), then by Paul (1988) as a possible aublysodontine tyrannosaurid.
Elzanowski and Wellnhofer (1992, 1993) originally suggested Archaeornithoides
was most closely related to spinosaurids, troodontids and Lisboasaurus,
and that these were all closer to birds than other known theropods. Spinosaurids are now known to be basal tetanurines
while Lisboasaurus is a crocodiliform. However, in their 1993 article,
they suggest Archaeornithoides is more closely related to birds than
Lisboasaurus, confusing matters slightly. They rejected a troodontid
relationship based on the broad maxillary palatal shelf and unserrated teeth,
but Currie (2000) noted this is invalid as Troodon has the former, while
Clark et al. (2002) noted it's invalid because Byronosaurus has both
features. Indeed, both characters are now recognized as primitive for troodontids,
and maniraptoriforms in general. Currie further suggested that Archaeornithoides
may be a juvenile Saurornithoides mongoliensis, while Averianov and Sues
(2007) suggested it could be a juvenile Byronosaurus. Besides the latter,
there are other Djadokhta troodontids with serrationless teeth- Almas, Gobivenator and
IGM 100/1128, but Archaeornithoides doesn't show a strong
resemblence to these or other paravians. Although Averianov and Sues stated
that Chiappe et al. (1996) proposed Archaeornithoides was a juvenile
dromaeosaurid, the latter authors actually only state they believe the lack
of serrations to be a juvenile character of birdlike theropods, as they mistakenly
assigned IGM 100/972 and 100/974 to Dromaeosauridae at the time. Archaeornithoides emerges in a polytomy with Scipionyx,
compsognathids and maniraptoriforms in Hartman et al.'s (2019)
analysis, the only published one its been included in. However,
only a single step moves it to Paraves where it emerges as a troodontid
near Mei and Xiaotingia, leaving its position still highly uncertain.
References- Elzanowski, 1983. Birds in Cretaceous Ecosystems. Acta Palaeontologia
Polonica. 28(1-2), 75-92.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster Co..
464 pp.
Elzanowski and Wellnhofer, 1992. A new link between theropods and birds from
the Cretaceous of Mongolia. Nature. 359, 821-823.
Elzanowski and Wellnhofer, 1993. Skull of Archaeornithoides from the
Upper Cretaceous of Mongolia. American Journal of Science. 293-A, 235-252.
Chiappe, Norell and Clark, 1996. Phylogenetic position of Mononykus (Aves:
Alvarezsauridae) from the Late Cretaceous of the Gobi Desert. Memoirs of the
Queensland Museum. 39(3), 557-582.
Currie, 2000. Theropod dinosaurs from the Cretaceous of Mongolia. In Benton,
Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia.
434-455.
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.
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.
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
Scipionyx Dal Sasso and Signore,
1998a
= "Dromaeodaimon" Signore, 1995
S. samniticus Dal Sasso and Signore, 1998a
= "Dromaeodaimon irene" Signore, 1995
Early Albian, Early Cretaceous
Pietraroja Formation, Italy
Holotype- (SBA-SA 163760) (~461 mm; <3 week old juvenile) skull (51.7
mm), sclerotic ring, mandibles (47.3 mm), hyoids (18.7 mm), atlantal neurapophysis,
atlantal intercentrum, axis (4.8 mm), third cervical vertebra (4.3 mm), fourth
cervical vertebra (4.5 mm), fifth cervical vertebra (4.7 mm), sixth cervical
vertebra (~4.7 mm), seventh cervical vertebra (4.9 mm), eighth cervical vertebra
(5.4 mm), ninth cervical vertebra (~5 mm), tenth cervical vertebra (4.9 mm),
nine pairs of cervical ribs (some partial; 9.1-15.9 mm), (dorsal series 69 mm)
first dorsal vertebra (5.1 mm), second dorsal vertebra (4.9 mm), third dorsal
vertebra, fourth dorsal vertebra, fifth dorsal vertebra, sixth dorsal vertebra,
seventh dorsal vertebra, eighth dorsal vertebra (~5.9 mm), ninth dorsal vertebra
(6 mm), tenth dorsal vertebra (6.5 mm), eleventh dorsal vertebra, twelfth dorsal
vertebra (6.3 mm), thirteenth dorsal vertebra (6.3 mm), twelve pairs of dorsal
ribs (some partial; 15.3-30.6 mm), thirteenth dorsal rib frament, eighteen rows
of gastralia, (sacrum ~23 mm) first sacral vertebra (7.4 mm), second sacral
centrum, fourth sacral neural arch, fourth sacral rib, fifth sacral vertebra
(5.1 mm), fifth sacral rib, first caudal vertebra (5.4 mm), second caudal vertebra
(5.4 mm), third caudal vertebra (5.6 mm), fourth caudal vertebra (5.7 mm), fifth
caudal vertebra (7.4 mm), sixth caudal vertebra (7.6 mm), seventh caudal vertebra
(6.8 mm), incomplete eighth caudal vertebra, ninth caudal neural arch fragment,
fourth chevron (8.1 mm), fifth chevron (6.7 mm), sixth chevron fragment, scapulae
(23.8 mm), coracoids (6.8, 6.8 mm), furcula (~12 mm), humeri (26.3 mm), radii
(17.5 mm), ulnae (19.3 mm), scapholunares (1.7 mm), semilunate carpals (2.4, 3 mm),
metacarpals I (4, 4 mm), phalanges I-1 (9.1, ~9.5 mm), manual unguals I (8.2
mm), metacarpals II (10.6, 10.6 mm), phalanges II-1 (7.3, ~6.7 mm), phalanges
II-2 (10.4, 10.2 mm), manual unguals II (one proximal; 8.1 mm), metacarpals
III (8.6, 8.7 mm), phalanges III-1 (3.1, 2.9 mm), phalanges III-2 (3.1 mm),
phalanges III-3 (one partial; 7.4, 6.7), manual unguals III (6.1 mm), horny
manual claws, incomplete ilia (~26.7 mm), pubes (27.3 mm), ischia (~20.4 mm),
femora (37.3 mm), proximal tibiae, proximal fibulae, cartilage, ligaments, tracheal
fragment, liver, esophagal fragment, stomach, intestine, rectum, cervical muscles,
dorsal epaxial muscles, puboischiofemoral muscle, caudifemoralis longus muscle,
lateral caudal musculature, fecal pellets, fish vertebrae, teleost scales, lepidosaur
pedal elements, lepidosaur scales
Diagnosis- (modified from Dal Sasso and Signore, 1998a) accessory transverse
postorbital ridge at fronto-parietal contact; compressed scapholinare and semilunate
carpal.
(after Dal Sasso and Maganuco, 2011) five premaxillary teeth; ventral squamosal
process squared; only two carpals; distal carpals I and II fused; manual digit
III much longer (123%) than digit I; preacetacetabular anterior concavity slightly
developed and facing anteriorly; obturator process quadrangular.
Comments- This specimen was first mentioned by Leonardi and Teruzzi (1993)
and described in depth in Signore's (1995) unpublished thesis. It was preliminarily
described and named by Dal Sasso and Signore in 1998 and monographed by Dal
Sasso and Maganuco (2011)
The sternal plate identified by Dal Sasso and Signore (1998a) is actually the left proximal
humerus (Dal Sasso and Maganuco, 2011).
Initially assigned to Maniraptoriformes by Dal Sasso and Signore
(1998a), Dal Sasso and Maganuco (2011) used a TwiG matrix to recover it
as a basal compsognathid. Most recently, it emerges in a polytomy
with Aniksosaurus, compsognathids and maniraptoriforms in Hartman et al.'s (2019) analysis.
References- Leonardi and Teruzzi, 1993. Prima segnalazione di uno scheletro
fossile di dinosauro (Theropoda, Coelurosauria) in Italia (Cretacico di Pietraroia,
Benevento). Paleocronache. 1993, 7-14.
Signore, 1995. Il teropode del Plattenkalk della Civita di Pietraroia (Cretaceo
inferiore, Benevento). Unpublished thesis. University of Napoli "Federico II". [pp]
Dal Sasso and Signore, 1998a. Exceptional soft tissue preservation in a theropod
dinosaur from Italy. Nature. 392, 383-387.
Dal Sasso and Signore, 1998b. Scipionyx samniticus (Saurischia, Theropoda):
The first Italian dinosaur. Third European Workshop on Vertebrate Paleontology,
Abstract: 23.
Dal Sasso and Signore, 1998c. Scipionyx samniticus (Theropoda: Coelurosauria)
and its exceptionally well preseved internal organs. Journal of Vertebrate Paleontology.
18(3), 37A.
Ruben, Dal Sasso, Geist, Hillenius, Jones and Signore, 1998. Pulmonary function
and metabolic physiology of theropod dinosaurs. Science. 283, 514-516.
Galliano and Signore, 1999. Parental care in theropod dinosaurs: possible evidences
from Scipionyx samniticus. Journal of Vertebrate Paleontology. 19(3),
46A.
Signore, 2001. Scipionyx samniticus (Theropoda, Maniraptoriformes) and the palaebiology of some maniraptoran theropods. PhD thesis, University of Bristol. 159 pp.
Dal Sasso and Maganuco, 2009. Osteology, ontogenetic assessment, phylogeny,
paleobiology, and soft-tissue anatomy of Scipionyx samniticus. Journal
of Vertebrate Paleontology. 29(3), 84A.
Dal Sasso and Maganuco, 2011. Scipionyx samniticus (Theropoda: Compsognathidae)
from the Lower Cretaceous of Italy: Osteology, ontogenetic assessment, phylogeny,
soft tissue anatomy, taphonomy, and palaeobiology. Memorie della Societ�
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
281 pp.
Klingler, 2015. Tracheal and esophageal displacement in the remarkably preserved
compsognathid Scipionyx samniticus. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 156.
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
Compsognathidae Cope, 1871
Definition- (Compsognathus longipes <- Passer domesticus)
(Holtz, Molnar and Currie, 2004)
Other definitions- (metacarpal I very short (MII/MI < 35%, with discrete
extensor tubercle directed proximo-radially (ratio 1.8–1.4 proximally to
radially), and barely asymmetrical distal condyles [< 5� offset ulnar
to radial condyles] as in Compsognathus longipes) (Gishlick and Gauthier,
2007)
= Compsognatha Huxley, 1870
= Compsognathinae Cope, 1871 vide Nopcsa, 1923
= Compsognathia Paul, 1988
= Sinosauropterygiformes Ji and Ji, 1996
= Sinosauropterygidae Ji and Ji, 1996
= Aptilonia Ji and Ji, 2001
= Eoptilonia Ji and Ji, 2001
Comments- Both Aptilonia and Eoptilonia were named by Ji and Ji (2001)
in a cladogram, the former including Compsognathus and the latter including
Sinosauropteryx. Though not defined, their etymology suggests reference
to Sinosauropteryx's preserved primitive feathers and Compsognathus'
lack of well preserved feathers. The latter is probably preservational, neither
state is apomorphic, and both names are best seen as junior synonyms of Compsognatha
and Sinosauropterygiformes respectively.
Placement of Compsognathidae relative to the base of Tyrannoraptora has
been contentious. Most recently, Hartman et al. (2019) recovered
them as non-maniraptoriform maniraptoromorphs. Eight more steps
are required to move them to Maniraptora as in Gauthier (1986), 13 more
to move them to Tyrannosauroidea as in Olshevsky (1991) and 15 more to
move them outside Tyrannoraptora as in Paul (1988).
References- Huxley, 1870. On
the classification of the Dinosauria, with observations on the
Dinosauria of the Trias. Quarterly Review of the Geological Society of
London, 26, 32-51.
Cope, 1871. On the homologies of some of the cranial bones of the
Reptilia, and on the systematic arrangement of the class. Proceedings
of the American Association for the Advancement of Science. 19, 194-247.
Nopcsa, 1923. Die Familien der Reptilien. Forschritte der Geologie und
Palaeontologie. Verlag von Gebr�der Borntraeger, Berlin. 2, 1-210.
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 and Schuster.
464 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope,
1869, excluding the advanced Crocodylia. Mesozoic Meanderings. 2, 196
pp.
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, 2001. How can we define a feathered dinosaur as a bird? In
Gauthier and Gall (eds.). New Perspectives on the Origin and Early
Evolution of Birds: Proceedings of the International Symposium in Honor
of John H. Ostrom. 43-46.
Holtz, Molnar and Currie, 2004. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 71-110.
Gishlick and Gauthier, 2007. On the manual morphology of Compsognathus longipes
and its bearing on the diagnosis of Compsognathidae. Zoological Journal of the
Linnean Society. 149, 569-581.
Tugulusaurus Dong, 1973
T. faciles Dong, 1973
Early Cretaceous
Lianmugin Formation of Tugulu Group, Xinjiang, China
Holotype- (IVPP V4025) dorsal rib, four incomplete mid caudal vertebrae
(23, 25, 34 mm), metacarpal I (26 mm), manual phalanx I-1 (54 mm), manual ungual
I (70 mm), femora (one proximal) (215 mm), tibia (~240 mm), astragalus (32 mm
wide), astragalar fragment, calcaneum, distal metatarsal III, distal metatarsal
IV, pedal phalanx IV-? (27 mm), pedal ungual III
Diagnosis- (after Rauhut and Xu, 2005) proximal mid-caudal vertebrae
with neural arch placed only on anterior two thirds of centrum and centrum considerably
broader than high (ratio width/height ca. 1.5); caudal centra rapidly increasing
in length distally; minimal length of metacarpal I less than width of this bone;
tibia with pronounced, semicircular lateral expansion of lateral malleolus.
Comments- The tibia is referred to as a radius in the translation of
Dong 1973, which explains the rather odd statement that the radius exceeds femoral
length in Glut (1997). Although Dong states Tugulusaurus differs from
other ornithomimids in that the proximal third metatarsal does not constrict,
the proximal end is unpreserved.
Phylogenetic relationships- Dong (1973) refers this genus to the Coelurosauria based
on hollow long bones and tibia longer than femur, and to the Ornithomimidae
based on the outline and characteristics of the metatarsus and phalanges.
Tugulusaurus was redescribed
by Rauhut and Xu (2005) and recovered as the first branching
coelurosaur in their analysis. Instead, the Hartman et al. (2019)
places it in Compsognathidae. Rauhut and Xu's position basal to Coelurus and compsognathids (which consisted of Compsognathus, Sinosauropteryx, Aristosuchus and the then-unnamed Mirischia)
was based on three characters. "Ascending process of astragalus:
arising out of lateral part of astragalar body" (127:0) is equivalent
to Hartman et al. character 209:0, which is quantified as "halfway up,
<58% width of astragalocalcaneum." This is also true in Coelurus, scored unknown by Rauhut and Xu. Among compsognathids, Sinosauropteryx and Compsognathus
only expose their ascending process in section or obliquely so cannot
be measured precisely, but are not obviously broad as Rauhut and Xu
score the family. Nqwebasaurus was misscored 1 but has state 0. Additionally, the compsognathids Huaxiagnathus and Aorun have narrow processes, as does the basal tyrannosauroid Guanlong and the basalmost scorable therizinosaur Erliansaurus.
"Ascending process of astragalus: ... [less] than twice height of
astragalar body" (128:0/1) is similar to Hartman et al. character
208:1, although that compares process height to astragalocalcanear
width instead of astragalar body height. While Rauhut and Xu
score Tugulusaurus with an
uncertainty polymorphism regarding whether the process is lower or
higher than the body, their figures 4B and C combine to clearly show it
is higher so should be rescored 1. Coelurus again has a low process but was scored unknown. Of their included compsognathids, Sinosauropteryx does have state 2 but Compsognathus has a process higher than its body but by less than twice, so the family should be scored 1+2. Nqwebasaurus was misscored 2 but has state 1. Among taxa they did not include Guanlong has state 1, and Aorun
has state 0. "Anterior side of distal tibia: with distinct
"step", running obliquely from medio-distal to latero-proximal" (122:0)
is not in the Hartman et al. character list. Contra Rauhut and
Xu's scoring, it is present in Coelurus. Among compsognathids (scored unknown by Rauhut and Xu), Compsognathus appears to have a high angled ridge on its right tibia and Aorun possesses it as well. Harpymimus has the ridge too, as does Kinnareemimus, so ornithomimosaurs should be polymorphic. Therizinosaurs were scored unknown by Rauhut and Xu, but not only do Falcarius and Neimongosaurus have a ridge, Segnosaurus itself seems to as well (photo of IGM 100/81 courtesy of Zanno). Given its absence in Nothronychus, therizinosaurs should be rescored polymorphic. Basal maniraptoromorph Aniksosaurus exhibits a ridge but was not included by Rauhut and Xu. Basal tyrannosauroids not used by Rauhut with a ridge include Guanlong, Dilong and Juratyrant.
When the asterisked ten changes are made to Rauhut and Xu's matrix, trees result where Tugulusaurus can fall out in Compsognathidae.
Smith et al. (2007) also recovered Tugulusaurus outside tyrannosauroids plus maniraptoromorphs (including compsognathids and Coelurus) based on the same three characters (their 310, 320 and 321).
Rauhut et al. (2010) recovered the genus as the first branching coelurosaur (before compsognathids, Coelurus, Sinosauropteryx, Shaochilong and tyrannoraptorans) based on the narrow ascending process (their 198).
Although not indicated by their published figure 18, Choiniere et al. (2010) recovered Tugulusaurus
as sister to Tyrannoraptora. Within Tyrannoraptora,
tyrannosauroids, ornithomimosaurs and maniraptorans (including
compsognathids and Ornitholestes
plus Pennaraptora) form a trichotomy. This exclusion from
Tyrannoraptora is based on four characters. Three of these are
the same as in Rauhut and Xu (437, 449, 450), and the fourth is a
result of misscoring numerous taxa- "Lateral accessory cnemial crest
... present" (427:1) was misscored among non-pennaraptoran coelurosaurs
as unknown in Tanycolagreus (absent), Guanlong (present), Compsognathus (present), 'Ornithomimus edmontonensis' (= Dromiceiomimus) (present) and Falcarius (present), and misscored as absent in Tyrannosaurus (present) and Garudimimus (present). Thus a lateral crest is typical in basal maniraptoromorphs.
Novas et al. (2012) recovered Tugulusaurus basal to Bicentenaria and Tyrannoraptora, based on Rauhut and Xu's three characters.
Godefroit et al. (2013) used Cau's large theropod matrix to recover Tugulusaurus outside Tyrannoraptora (including Coelurus, Tanycolagreus and Guanlong as tyrannosauroids, and Sinocalliopteryx, compsognathids, Ornitholestes and Juravenator
as maniraptoromorphs) based on two characters. One is "Femur,
head and neck, proximodistal axis in proximal view: ... slightly
anteromedially directed (describes an angle of 40�-20� with the
mediolateral axis of the distal condyles)" (431:1), which is present in
Tugulusaurus whereas most
tyrannoraptorans have state 2, a strictly medially projected
head. These states are generally not determinable from figures or
slab specimens, and were not recognized as distinct by workers before
the mid-2000's so that prior to that both were considered simply
medially directed (e.g. Rauhut's 2003 character 195). Further,
Godefroit et al.'s given angle ranges aren't typical, as Benson
(2009:11) describes state 1 as "around 20 degrees", compared to state
0's "around 40-45 degrees." Indeed Zuolong, scored state 1 by Godefroit et al., has an angle of ~15 degrees. Given these caveats, Godefroit's scoring of 2 for Guanlong
is incorrect as "the head appears to be angled slightly anteromedially
... but deformation of the specimen precludes confirmation of the true
angle" (Choiniere, 2010:176-177), and no other basal tyrannosauroid
condition has been reported until taxa as close to Tyrannosaurus as Dryptosaurus
(Brusatte et al., 2011:26- "the head projects medially and not at all
anteriorly"). Among maniraptoromorphs most basal taxa preserving
proximal femora are slab specimens, and note Mirischia's
condition has not been reported in a modern context, where earlier
reports of a medially directed head (e.g. Rauhut, 2003:112) indicate
only state 1/2. However, basal therizinosaur Falcarius
has "a slight cranial deflection from the transverse plane relative to
the distal condyles" (Zanno, 2010:217; misscored by Godefroit et al.),
basal ornithomimosaur Deinocheirus' head is "twisted 15 degrees anteromedially to the femoral shaft" (Lee et al., 2014:259) and basal maniraptoromorph Aniksosaurus
has a "craniomedially projected" head (Martinez and Novas,
2006:250). Thus, like astragalar morphology, a strictly medial
femoral head angle may have evolved convergently in several
tyrannoraptoran clades. The other character supporting this is
"Tibia, fibular condyle, posteriormost extent in proximal view: ... at
the same level of ... the posterior margin of the medial condyle"
(449:1), whereas most tyrannoraptorans are scored as having a fibular
condyle whose posterior margin is anteriorly offset. While not
quantified by Godefroit et al., if we limit state 1 to those taxa with
medial condyles extending 11% or less posteriorly past the lateral
condyle (using the distance from the cnemial crest tip to the
intercondylar groove as 100%; Tugulusaurus' measures 7%), Garudimimus (6%), Gallimimus (11%) and Falcarius
(10%) were misscored by Godefroit et al.. Indeed,
ornithomimosaurs and therizinosaurs exhibit the primitive condition
except for Deinocheirus and Alxasaurus. Other basal maniraptoromorphs without posteriorly restricted lateral condyles are Scipionyx (lateral condyle extends posterior to medial), Kinnareemimus (4%) and Nedcolbertia (4%), none of which Godefroit et al. included.
In more recent analyses, Tugulusaurus can fall out more crownward. Choiniere et al. (2013) resolved Tugulusaurus
as a coelurid maniraptoran when ontogeny was accounted for.
Brusatte et al. (2014) recovered it in a trichotomy with
tyrannosauroids and maniraptoromorphs. Xu et al. (2018) resolved Tugulusaurus as an alvarezsaur sister to their new taxon Xiyunykus, closer to alvarezsaurids than Haplocheirus or their uniquely alvarezsaurian Aorun.
This position in Alvarezsauroidea was based on four characters not
included by Hartman et al. ("strong medial tab on metacarpal [I]";
"dorsolaterally and dorsomedially facing lateral and medial surfaces of
phalanx [I]-1 that are shallowly concave"; "axial furrow along the
flexor surface of phalanx [I]-1"; "partially enclosed ‘flexor notches’
on the medial and lateral surfaces of the proximal end of the ventral
surface of manual ungual [I]-2"), where it takes 5 more steps to force
as an alvarezsauroid. Thus the true difference between a
compsognathid and alvarezsauroid placement could be as low as a single
step. Thus it is only placed tentatively in Compsognathidae
here. In any case, Tugulusaurus'
basal position in analyses a decade after its redescription is based on
misscorings and a few characters which more parsimoniously converge in
tyrannosaurids, ornithomimosaurs and pennaraptorans.
References- Dong, 1973. Dinosaurs from Wuerho. In Reports of paleontological expedition to Sinkiang
(II), pterosaurian fauna from Wuerho, Sinkiang. Memoirs of the Institute of
Vertebrate Paleontology and Paleoanthropology Academia Sinica. 11, 45-52.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press. 1076 pp.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 213 pp.
Rauhut and Xu, 2005. The small theropod dinosaurs Tugulusaurus and Phaedrolosaurus
from the Early Cretaceous of Xinjiang, China. Journal of Vertebrate Paleontology.
25(1), 107-118.
Mart�nez and Novas, 2006. Aniksosaurus darwini
gen. et sp. nov., a new coelurosaurian theropod from the early Late
Cretaceous of central Patagonia, Argentina. Revista del Museo Argentino
de Ciencias Naturales. 8(2), 243-259.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti
(Dinosauria: Theropoda) from the Early Jurassic of Antarctica and
implications for early theropod evolution. Zoological Journal of the
Linnean Society. 151, 377-421.
Benson, 2009. An assessment of variability in theropod dinosaur remains
from the Bathonian (Middle Jurassic) of Stonesfield and New Park
Quarry, UK and taxonomic implications for Megalosaurus bucklandii and Iliosuchus incognitus. Palaeontology. 52(4), 857-877.
Choiniere, 2010. Anatomy and systematics of coelurosaurian theropods
from the Late Jurassic of Xinjiang, China, with comments on forelimb
evolution in Theropoda. PhD thesis. George Washington University. 994
pp.
Choiniere, Clark, Forster and Xu, 2010. A basal coelurosaur
(Dinosauria: Theropoda) from the Late Jurassic (Oxfordian) of the
Shishugou Formation in Wucaiwan, People's Republic of China. Journal of
Vertebrate Paleontology. 30(6), 1773-1796.
Rauhut, Milner and Moore-Fay, 2010. Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi
(Woodward, 1910) from the Middle Jurassic of England. Zoological
Journal of the Linnean Society. 158(1), 155-195.
Zanno, 2010. Osteology of Falcarius utahensis (Dinosauria: Theropoda):
Characterizing the anatomy of basal therizinosaurs. Zoological Journal
of the Linnaean Society. 158(1), 196-230.
Brusatte, Benson and Norell, 2011. The anatomy of Dryptosaurus aquilunguis
(Dinosauria: Theropoda) and a review of its tyrannosauroid affinities.
American Museum Novitates. 3717, 53 pp.
Novas, Ezcurra, Agnolin, Pol and Ortiz, 2012. New Patagonian Cretaceous
theropod sheds light about the early radiation of Coelurosauria.
Revista del Museo Argentino de Ciencias Naturales. 14(1), 57-81.
Choiniere, Clark, Forster, Norell, Eberth, Erickson, Chu and Xu, 2014 (online 2013).
A juvenile specimen of a new coelurosaur (Dinosauria: Theropoda) from
the Middle-Late Jurassic Shishugou Formation of Xinjiang, People's
Republic of China. Journal of Systematic Palaeontology. 12(2), 177-215.
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, Barsbold, Currie, Kobayashi, Lee, Godefroit, Escuillie and
Tsogtbaatar, 2014. Resolving the long-standing enigmas of a giant
ornithomimosaur Deinocheirus mirificus. Nature. 515, 257-260.
Xu, Choiniere, Tan, Benson, Clark, Sullivan, Zhao, Han, Ma, He, Wang,
Xing and Tan, 2018. Two Early Cretaceous fossils document transitional
stages in alvarezsaurian dinosaur evolution. Current Biology. 28, 1-8.
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
Shishugounykus Qin, Clark, Choiniere and Xu, 2019
S. inexpectus Qin, Clark, Choiniere and Xu, 2019
Early Oxfordian, Late Jurassic
Wucaiwan, Middle Shishugou Formation, Xinjiang, China
Holotype- (IVPP V23567) (6.8
kg; 9 year old young adult) fragmentary frontals, incomplete parietal,
partial laterosphenoid, partial angular, articular, partial anterior
dorsal vertebra (21 mm), partial mid dorsal vertebra, partial last
dorsal vertebra (21.5 mm), few rib fragments, fused partial first and
second sacral vertebrae (17.3, 17.4 mm), ?last two sacral centra (16.9,
15.7 mm), proximal caudal vertebra (17.4 mm), incomplete mid caudal
vertebra (20.3 mm), incomplete mid caudal centum, distal caudal
vertebra (26.9 mm), mid chevron, incomplete scapula, proximal humerus,
proximal radius, proximal ulna, metacarpal I (21.7 mm), incomplete
phalanx I-1 (45.6 mm), incomplete manual ungual I, metacarpal II (~42
mm), phalanx II-1 (31.6 mm), phalanx II-2 (41.7 mm), incomplete manual
ungual II (36.5 mm), incomplete phalanx III-1 (13.7 mm), phalanx III-2
(13.5 mm), manual ungual III (16.9 mm), partial ilium, pubic shaft
fragment, incomplete ischium, femora (one incomplete; 175.8 mm), tibiae
(one incomplete, one partial; 222.7 mm), proximal fibulae, distal
tarsal, distal metatarsal II, distal metatarsal III, phalanx III-1
(31.5 mm), phalanx III-2 (23.8 mm), phalanx IV-1 (22.1 mm), phalanx
IV-2 (19.4 mm), phalanx IV-4 (13.1 mm), fragments
Diagnosis- (after Qin et al.,
2019) supratemporal fossa occupying large portion of frontal; frontal
with indistinct anterior border; scapula with hollow acromial process
but without lateral concavities; humeral internal tuberosity
mediolaterally constricted distally, giving it a 'pinched' appearance;
metacarpal II straight in dorsal view; manual ungual II subequal in
size to ungual I; iliac medial surface with step-wise transition from
ischial peduncle to pubic peduncle; distal end of metatarsal II
asymmetrically ginglymoid.
Comments- Note the skeletal
reconstruction in Qin et al. (2019) does not include the anterior
dorsal, rib fragments or chevron, and incorrectly shows distal
metatarsal IV preserved instead of II. The distal caudal vertebra
illustrated is not mentioned in the text but is in the measurements
table.
Qin et al. announced this in an abstract (2018) and later described it
preliminarily (2019). They used Choiniere's coelurosaur matrix to
recover it as an alvarezsauroid more basal than Haplocheirus, Xiyunykus and Bannykus, in a trichtomy with Aorun and more derived taxa. As with Haplocheirus and Aorun,
Hartman et al.'s maniraptoromorph analysis instead recovers it in
Compsognathidae. Only four steps move it to Alvarezsauroidea
however (between Nqwebasaurus and Pelecanimimus),
suggesting further studies are needed. Agnolin et al. (2022) were
also skeptical of an alvarezsauroid relationship, although they
suggested it was similar to Haplocheirus which they considered could be closer to ornithomimosaurs.
References- Qin, Clark and Xu,
2018. A new Jurassic alvarezsaurian theropod from the Shishugou
Formation of western China demonstrates an early diversification of the
group. Journal of Vertebrate Paleontology. Program and Abstracts, 200.
Qin, Clark, Choiniere and Xu, 2019. A new alvarezsaurian theropod from
the Upper Jurassic Shishugou Formation of western China. Scientific
Reports. 9:11727.
Agnolin, Lu, Kundrat and Xu, 2022 (online 2021). Alvarezsaurid
osteology: New data on cranial anatomy. Historical Biology. 34(3),
443-452.
Sinosauropteryx Ji and Ji,
1996
S. prima Ji and Ji, 1996
= Compsognathus prima (Ji and Ji, 1996) Morell, 1997
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Beds of Yixian Formation, Liaoning, China
Holotype- (GMV 2123, NIGP 127586) (680 mm; subadult) skull (62.5 mm),
sclerotic plates, mandible, hyoids, eight cervical vertebrae, nine cervical
ribs (third 13 mm, sixth 10 mm, eighth 6 mm), eleven dorsal centra, twenty dorsal
ribs, gastralia, fifty-nine caudal vertebrae, thirty-four chevrons, scapulae,
coracoids, humeri (20.3 mm), radii (12.4 mm), ulnae, distal carpal I (2.9 mm),
metacarpals I (4.2 mm), phalanges I-1, manual unguals I, metacarpals II (10.2
mm), phalanges II-1, phalanges II-2, manual unguals II, metacarpals III, phalanx
III-1, phalanx III-2, phalanx III-3, manual ungual III, ilium (39 mm), pubes
(41.3 mm), ischia, femora (53.2 mm), tibiae (61 mm), fibulae, astragali, calcanea,
distal tarsals III, distal tarsals IV, metatarsals II, metatarsals III (39.9
mm), phalanx III-1, phalanx III-2, metatarsals IV (36.8 mm), phalanx IV-1, phalanx
IV-2, phalanx IV-3, phalanx IV-4, six pedal phalanges, metatarsal V (8.1 mm),
feathers, viscera
Referred- (NIGP 127587) (1.07 m; young adult) incomplete skull (97.2
mm), sclerotic plates, incomplete mandibles, hyoids, ten cervical vertebrae
(third cervical vertebra 9.6 mm), twelve dorsal vertebrae, sixteen dorsal ribs,
dorsal rib fragments, gastralia, partial sacrum, twenty-three caudal vertebrae,
twenty-three chevrons, scapulae, coracoids, humeri (35.5 mm), radii (21 mm),
ulnae, scapholunare (3 mm), distal carpal I (5.6 mm), distal carpal II (1.8 mm),
metacarpals I (8.6 mm), phalanges I-1 (19.4 mm), manual ungual I (~25 mm), metacarpals
II (17.1 mm), phalanges II-1 (9.5 mm), phalanx II-2 (11.8 mm), manual ungual
II (~14 mm), metacarpals III (12.7, 13.7 mm), phalanges III-1 (5, 4.6 mm), phalanges
III-2 (3.7, 3.8 mm), phalanges III-3 (5.6 mm), manual unguals III (9.9 mm),
ilia (67.5 mm), pubis (74 mm), ischia, femora (86.4 mm), tibiae (97 mm), astragalus,
calcaneum, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsals
II (~58 mm), phalanx II-1, phalanges II-2, pedal unguals II, metatarsals III
(~65 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III,
metatarsals IV (~60 mm), phalanges IV-1, phalanges IV-2, phalanx IV-3, phalanx
IV-4, pedal ungual IV, metatarsal V fragment, feathers, intestine (Chen et al., 1998)
Early Aptian, Early Cretaceous
Dawangzhangzi Beds of Yixian Formation, Liaoning, China
(D 2141) skull (86.6 mm), mandible, hyoid (23.3 mm), cervical vertebrae, cervical
ribs, dorsal vertebrae, fifteen rows of gastralia, sacrum, twenty caudal vertebrae,
chevrons, partial scapulae, coracoids, humerus (24.7 mm), radius, ulna (21.3
mm), metacarpal I, phalanx I-1, metacarpal II, phalanx II-1, phalanx II-2, manual
ungual II, manual digit III, ilia, ischia, femora, tibiae (72.8 mm), fibulae,
metatarsals II, metatarsals III 52.3 mm), metatarsals IV, pedal phalanges, pedal
unguals, feathers (Ji et al., 2007)
(IVPP V12415) specimen including skull, mandible, hyoid, cervical series, cervical
ribs, dorsal series, dorsal ribs, gastralia, caudal series, chevrons, scapula,
coracoids, humeri, ulna, manual elements, manual ungual I, ilium, pubis, ischia,
tibiae, phalanx I-1, pedal ungual I, pedal digits, feathers (Lingham-Soliar
et al., 2007)
(IVPP V14202) specimen including sacrum, eleven proximal caudal vertebrae, ten
chevrons, ilium and feathers (Zhang et al., 2010)
Diagnosis- (after Currie and Chen, 2001) first manual digit is longer
than the humerus or the radius; powerful proximomedial flange on first metacarpal.
Comments-
A lizard skeleton is preserved in the gut region of NIGP 127587.
Wang et al. (2022) stated "Ovoid structures in the abdominal cavity of
a specimen of Sinosauropteryx [NIGP 127587], and argued to be eggs, closely match in shape and position the duodenal portion of Scipionyx intestine and the ventral part of the bluish layer in Daurlong, and are here re-interpreted as intestinal remnants."
A specimen described by Ji and Ji (1997), NGMC 2124, seems to be a different
taxon. This was first suggested by Longrich (DML, 2000), who later wrote an
abstract on it in 2002. This is agreed on by Ji et al. (2007) and Gishlick and
Gauthier (2007), who label it Sinosauropteryx? sp..
References- 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. Advances in the study of the avian Sinosauropteryx prima.
Chinese Geology. 242, 30-32.
Morell, 1997. The origin of birds: the dinosaur debate. Audubon Magazine, April,
36-45.
Chen, Dong and Zhen, 1998. An exceptionally well-preserved theropod dinosaur
from the Yixian Formation of China. Nature. 391, 147-152.
Longrich, DML 2000. https://web.archive.org/web/20201115172810/http://dml.cmnh.org/2000Apr/msg00300.html
Currie and Chen, 2001. Anatomy of Sinosauropteryx prima from Liaoning,
northeastern China. Canadian Journal of Earth Sciences. 38(12), 1705-1727.
Longrich, 2002. Systematics of Sinosauropteryx. Journal of Vertebrate
Paleontology. 22(3), 80A.
Gishlick and Gauthier, 2007. On the manual morphology of Compsognathus longipes
and its bearing on the diagnosis of Compsognathidae. Zoological Journal of the
Linnean Society. 149, 569–581.
Ji, Gao, Liu, Meng and Ji, 2007. New material of Sinosauropteryx (Theropoda:
Compsognathidae) from western Liaoning, China. Acta Geologica Sinica. 81(2),
177-182.
Lingham-Soliar, Feduccia and Wang, 2007. A new Chinese specimen indicates that
'protofeathers' in the Early Cretaceous theropod dinosaur Sinosauropteryx
are degraded collagen fibres. Proceedings of the Royal Society B. 274, 1823-1829.
Zhang, Kearns, Orr, Benton, Zhou, Johnson, Xu and Wang, 2010. Fossilized melanosomes
and the colour of Cretaceous dinosaurs and birds. Nature. 463, 1075-1078.
Smithwick, Nicholls, Cuthill and Vinther, 2017. Countershading and stripes in the theropod dinosaur Sinosauropteryx reveal heterogeneous habitats in the Early Cretaceous Jehol Biota. Current Biology. 27(21), 3337-3343.
Smithwick, Mayr, Saitta, Benton and Vinther, 2017. On the purported
presence of fossilized collagen fibres in an ichthyosaur and a theropod
dinosaur. Palaeontology. 60(3), 409-422.
Saitta, Gelernter and Vinther, 2018 (online 2017). Additional
information on the primitive contour and wing feathering of paravian
dinosaurs. Paleontology. 61(2), 273-288.
Wang, Cau, Guo, Ma, Qing and Liu, 2022. Intestinal preservation in a
birdlike dinosaur supports conservatism in digestive canal evolution
among theropods. Scientific Reports. 12:19965.
Santanaraptor Kellner, 1999
S. placidus Kellner, 1999
Albian, Early Cretaceous
Romualdo Formation of Santana Group, Brazil
Holotype- (MN 4802-V) (~1.25 m; juvenile) three distal caudal vertebrae,
distal chevrons, ischia (91 mm), femora (~167 mm), partial tibiae, partial fibulae,
metatarsal I, metatarsals II, phalanges, II-1, phalanges II-2, pedal unguals
II, metatarsals III (~136 mm), 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 unguals IV, skin, musculature
Diagnosis- (modified from Kellner, 1999) foramen at medial base of anterior
trochanter; well developed sulcus on posterior femoral head; fibular trochlea
of triangular shape and constricted at base.
Comments- This theropod was known first announced by Kellner in 1996
and emphasis was placed on the soft tissue preserved with the specimen. At that
time, it was identified as a probable maniraptoran. It was later preliminarily
described and named in 1999, though a more detailed description is planned.
Initially described briefly as a maniraptoriform (Kellner, 1999) based
on Sereno's (1999) character "obturator notch U-shaped with slightly
divergent sides" (which I find highly variable), this has been
recovered in an uncertain position within Tyrannoraptora in the two
TWiG analyses which added it. Dal Sasso and Maganuco (2011) found
it emerged as a tyrannoraptoran excluded from Coeluridae plus other
tyrannosauroids, Ornithomimosauria and Therizinosauria plus Metornithes
(note its maniraptoran position in their Fig. 113 is based on a
majority rules tree), while Novas et al. (2012) recovered it as a
tyrannoraptoran excluded from Tyrannosauridae, Ornithomimosauria and
Therizinosauria plus Pennaraptora. Most recently, Hartman et al.
(2019) recover it as a compsognathid, but Santanaraptor has usually been posited to be a tyrannosauroid in current literature.
Holtz (2004) questionably referred it to Tyrannosauroidea,
though without supporting evidence. Similarly, Agnolin et al. (2004) refer it
to Noasauridae without reason, though this seems less plausible.
Novas et al. (2013) and derivatives recovered it as a tyrannosauroid
sister to Megaraptora plus Tyrannosauridae based on six
characters. "Caudal vertebrae, ventral surface: flat" (their
113:0) must refer to distal caudals, as three of these are the only
preserved vertebrae in Santanaraptor. While not verified by text or illustration in Santanaraptor, of the taxa Hartman et al. recover in Compsognathidae, Tugulusaurus also lacks a median groove in the middle of its four preserved caudals. This is similar to tyrannosauroids as close to Tyrannosaurus as Juratyrant
but most compsognathids are not scorable due to lateral exposure of the
caudals. "Ischium, ischial tubercle ventral to iliac peduncle:
... present as a convex bulge on the posterior surface of the ischium"
(their 167:1) is equivalent to Hartman et al.'s 180:1. Although Sinosauropteryx was not interpreted by Novas et al. as having a proximodorsal process, it does show a convexity comparable to Santanaraptor.
"Ischium distal expansion: absent" (their 169:0) is equivalent to
Hartman et al.'s 189:1, but contrary to Novas et al., Hartman et al.
interpret the distal ischium as expanded in Santanaraptor (~1.44 times minimum shaft depth). This doesn't matter in respect to Novas et al.'s topology though, as Appalachiosaurus was also misscored and has a distal expansion and megaraptorans are now known to exhibit this state too (e.g. Murusraptor).
Thus an unexpanded ischium is convergent in tyrannosaurids and
pennaraptorans. "Femur, fossa on the medial surface of the head,
lateral to the trochanteric fossa, form: ... deep" (their 175:1) was
"modified from Brusatte et al., 2010a: 286", but as Brusatte and Carr
(2016:S33) note, the fossa is on the posterior surface and IS the
trochanteric fossa. It cannot be verified in Santanaraptor from available images and description. Contrary to Novas et al.'s scoring it seems to be present in Sinosauropteryx-
"the femoral head ... is separated from the low, ridge-like greater
trochanter by a shallow depression" (Currie and Chen, 2001:1718),
notable when the holotype's femur is exposed in posterior view.
They similarly misscored Compsognathus
as state 0 when the only specimen preserving a proximal femur has it
exposed in lateral view and crushed, so cannot be coded. It's
also developed in the unincluded Sinocalliopteryx,
but is generally difficult to discern in photos unless bound distally
by a ridge as in most tyrannosaurines. "Tibia, form of medial
malleolus: ... oriented almost medially, 'shoulder' absent" (their
187:2) and "Tibia, medial expansion of distal medial malleolus: ...
expanded 40% or more than tibial mid-shaft width" (their 190:1).
These are not determinable in Santanaraptor
as the distal tibia is shattered with the only preserved surface facing
into the sediment (photo of MN 4802-V courtesy of Bruno Campos).
Delcourt and Nelson Grillo (2018) recently analyzed Santanaraptor
in the matrices of Choiniere et al. (2012), Carr et al. (2017) and
Porfiri et al. (2018), recovering it as a tyrannosauroid in each based
on several characters. "A deep fossa on the medial surface of the
femoral head, lateral to the trochanteric fossa (175:1 in Porfiri et
al., 2018 and 362:1 in Carr et al., 2017)" was discussed above.
"An ischial medial apron positioned along the anterior margin of its
shaft in medial view (357:1 in Carr et al., 2017)" as scored for
tyrannosaurines is not present in Santanaraptor,
which has the posterior shaft most projected medially. "The
proximal margin of the femur is concave in posterior view due to a
greater trochanter that is elevated substantially relative to the
lateral portion of the proximal surface of the head (361:1 in Carr et
al., 2017)" is equivalent to Hartman et al.'s 490:1 and is not present
in Santanaraptor, which has a
slightly convex proximal margin and unelevated greater
trochanter. "The lesser trochanter and the greater trochanter
extending to approximately the same level proximally (360:1 in Carr et
al., 2017)" is similar to Hartman et al.'s 319:2 and is not present in Santanaraptor.
"A shallow femoral extensor groove on the anterior surface of the
distal end that is expressed as a broad concave anterior margin in
distal view but present as an extensive depression on the anterior
surface of the femur (366:1 in Carr et al., 2017)" is equivalent to
Hartman et al.'s 655:1 and would be similar to Juratyrant and more derived tyrannosauroids if verified in Santanaraptor.
"The absence of an accessory ridge on the lateral surface of the
cnemial crest (510:0 in Choiniere et al., 2012)" was discussed above
(see Tugulusaurus entry) and its distribution in Choiniere's matrices is based on misscoring several taxa. In actuality only Tanycolagreus has state 0 among tyrannosauroids, with additional tyrannosauroids exhibiting a ridge not mentioned in the Tugulusaurus discussion including- Yutyrannus, Juratyrant, Dryptosaurus, Alectrosaurus, Appalachiosaurus and Albertosaurus. Thus, if Santanaraptor
does have state 0, it would not support a tyrannosauroid
identification. "The absence of a horizontal groove across the
astragalar condyles anteriorly (539:0 in Choiniere et al., 2012)" is
700:0 in Hartman et al.'s analysis. While true of tyrannosauroids
at least as close to Tyrannosauridae as Dryptosaurus, it is also seen in almost every maniraptoromorph including Sinosauropteryx.
Uncertainty in the case of Santanaraptor
is due partly to its brief initial description, as a lack of a median
ventral groove in distal caudals and deep femoral extensor groove may
be tyrannosauroid-like, but have not been verified by text or
illustration. Other suggested characters are based on misscorings
or also present in maniraptoromorphs. As it only requires 5 steps
to place in Tyrannosauroidea in the Hartman et al. matrix, it is
tentatively assigned to Compsognathidae here pending better description
or photos.
References- Kellner, 1996. Fossilized theropod soft tissue. Nature. 379,
32.
Kellner and Campos, 1998. Archosaur soft tissue from the Cretaceous of the Araripe
Basin, northeastern Brazil. Boletim do Museu Nacional, Geologia. 42, 1-22.
Kellner, 1999. Short note on a new dinosaur (Theropoda, Coelurosauria) from
the Santana Formation (Romualdo Member, Albian), northeastern Brazil. Boletim
do Museu Nacional. 49, 1-8.
Currie and Chen, 2001. Anatomy of Sinosauropteryx prima from Liaoning,
northeastern China. Canadian Journal of Earth Science. 38(12), 1705-1727.
Agnolin, Apesteguia, and Chiarelli, 2004. The end of a myth: The mysterious
ungual claw of Noasaurus leali. Journal of Vertebrate Paleontology. 24(3),
301A.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria
Second Edition. University of California Press. 111-136.
Dal Sasso and Maganuco, 2011. Scipionyx samniticus (Theropoda: Compsognathidae)
from the Lower Cretaceous of Italy: Osteology, ontogenetic assessment, phylogeny,
soft tissue anatomy, taphonomy, and palaeobiology. Memorie della Societ�
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
281 pp.
Choiniere, Forster and de Klerk, 2012. New information on Nqwebasaurus thwazi,
a coelurosaurian theropod from the Early Cretaceous (Hauteriverian?)
Kirkwood Formation in South Africa. Journal of African Earth Sciences.
71-72, 1-17.
Novas, Ezcurra, Agnolin, Pol and Ortiz, 2012. New Patagonian Cretaceous
theropod sheds light about the early radiation of Coelurosauria.
Revista del Museo Argentino de Ciencias Naturales. 14(1), 57-81.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Brusatte and Carr, 2016. The phylogeny and evolutionary history of
tyrannosauroid dinosaurs. Scientific Reports. 6, 20252.
Carr, Varricchio, Sedlmayr, Roberts and Moore, 2017. A new tyrannosaur
with evidence for anagenesis and crocodile-like facial sensory system.
Scientific Reports. 7:44942.
Delcourt
and Nelson Grillo, 2018. Tyrannosauroids from the southern hemisphere:
Implications for biogeography, evolution, and taxonomy.
Palaeogeography, Palaeoclimatology, Palaeoecology. 511, 379-387.
Porfiri, Valieri, Santos and Lamanna, 2018. A new megaraptoran theropod
dinosaur from the Upper Cretaceous Bajo de la Carpa Formation of
northwestern Patagonia. Cretaceous Research. 89, 302-319.
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
Haplocheirus Choiniere, Xu,
Clark, Forster, Guo and Han, 2010
H. sollers Choiniere, Xu, Clark, Forster, Guo and Han, 2010
Early Oxfordian, Late Jurassic
Wucaiwan, Middle Shishugou Formation, Xinjiang, China
Holotype- (IVPP V15988) (~1.9-2.3 m; 19 kg; subadult) skull (212.8 mm), sclerotic
rings, mandibles, hyoid fragment, partial axis, incomplete third cervical vertebra
(26.5 mm), incomplete fourth cervical vertebra (28.3 mm), incomplete fifth cervical
vertebra, incomplete sixth cervical vertebra (32.5 mm), seventh cervical vertebra
(32.1 mm), eighth cervical vertebra (28 mm), ninth cervical vertebra (28.8 mm),
incomplete tenth cervical vertebra (25.4 mm), twelve cervical ribs, incomplete
first dorsal vertebra (25.2 mm), incomplete second dorsal vertebra (25.7 mm),
incomplete third dorsal vertebra (19.9 mm), incomplete fourth dorsal vertebra
(24.1 mm), incomplete fifth dorsal vertebra (21.4 mm), incomplete sixth dorsal
vertebra (24.4 mm), incomplete seventh dorsal vertebra (28.4 mm), incomplete
eighth dorsal vertebra (25.1 mm), ninth dorsal vertebra (25.6 mm), tenth dorsal
vertebra (28 mm), eleventh dorsal vertebra (26.5 mm), twelfth dorsal vertebra
(29 mm), thirteenth dorsal vertebra (28 mm), twenty-six dorsal ribs, eighteen
incomplete rows of gastralia, partial first sacral vertebra (29.1 mm), second
sacral centrum (25 mm), third sacral centrum (20.1 mm), partial fourth sacral
vertebra (21 mm), partial fifth sacral vertebra (21.5 mm), incomplete first
caudal vertebra (24.7 mm), incomplete second caudal vertebra (28.6 mm), third
caudal vertebra (27.6 mm), incomplete fourth caudal vertebra, incomplete fifth
caudal vertebra (30.1 mm), sixth caudal vertebra (32.8 mm), seventh caudal vertebra
(32 mm), eighth caudal vertebra, ninth caudal vertebra (32.3 mm), tenth caudal
vertebra (35.1 mm), eleventh caudal vertebra (33.8 mm), twelfth caudal vertebra
(34.2 mm), thirteenth caudal vertebra (35.3 mm), fourteenth caudal vertebra
(37.8 mm), eight distal caudal centra, chevrons 1-14, scapulae (~135 mm), coracoids
(32.7 mm tall), humeri (104.3 mm; one incomplete), radii (86, 83 mm), ulnae
(90.1 mm), scapholunare, distal carpal I, distal carpal II, metacarpals I (22.6 mm),
phalanx I-1 (49.9 mm), manual unguals I (55, 48.1 mm), metacarpals II (57 mm),
phalanx II-1 (33.7 mm), phalanges II-2 (50.6, 42 mm), manual unguals II (47.8,
50.4 mm), metacarpals III (26.2 mm), phalanx III-1 (14.7 mm), phalanx III-2
(15.6 mm), phalanx III-3 (30.2 mm), manual unguals III (30.7, 31.2 mm), partial
ilium, incomplete pubes (159.8 mm), ischia (121 mm), femora (214.3 mm), tibiae
(277.1, 269.3 mm), fibulae, astragali, calcanea, distal tarsal IV, metatarsal
II, phalanx II-1 (33.2 mm), phalanx II-2 (26 mm), pedal ungual II (32.3 mm),
metatarsal III (144.6 mm), partial phalanx III-1 (38.6 mm), incomplete phalanx
III-2 (26 mm), phalanx III-3 (26 mm), pedal ungual III (26.7 mm), metatarsal
IV (134.5 mm), phalanx IV-1 (27.6 mm), phalanx IV-2 (22 mm), phalanx IV-3 (15.8
mm), partial phalanx IV-4 (~13.5 mm), pedal ungual IV (~20.3 mm), metatarsal
V (44.6 mm)
Diagnosis- (after Choiniere et al., 2010) metacarpal III one-half the length of metacarpal II.
(after Choiniere et al., 2014) dorsally expanded distal end of posterior maxillary
process; ventral edge of distal paroccipital process twisted posteriorly.
Other diagnoses- Choiniere et al. (2010) suggested a
supposed second external mandibular fenestra was diagnostic, but
Choiniere (2010) later showed this was caused by breakage and
disarticulation. Choiniere et al. also suggested distally serrated
lateral teeth were diagnostic among alvarezsaurs, but these are
probably plesiomorphic, certainly if it is a compsognathid.
Similarly, "alveolar margin of anterior end of dentary is dorsally
convex" was listed by Choiniere et al. (2010) but is is found in other
compsognathids (Sinosauropteryx, Compsognathus, Aorun, Sciurumimus). They also list "heterodont dentary tooth row
with enlarged dentary tooth 4", but the first few dentary teeth being larger than others is also present in Compsognathus, Aorun and Sciurumimus.
Comments- While generally recovered as a basal
alvarezsauroid, Hartman et al. (2019) found it to be a compsognathid
instead, a placement originally proposed non-quantitatively by Alifanov
and Saveliev (2011:184). Similarly, Choiniere et al. (2011) found
the braincase anatomy of parvicursorines was more similar to avialans
than to Haplocheirus, suggesting homoplasy or alvarezsauroid
polyphyly. Among published phylogenetic analyses, Lee and Worthy (2011) recovered Haplocheirusas
the basalmost ornithomimosaur in their Bayesian reanalysis of a TWiG
matrix, not separated from compsognathids by a majority bootstrap
value. Enforcing a placement in Alvarezsauroidea requires 9 more
steps, but eight characters used by Choiniere et al. (2010) to place it
in the clade were not included. This suggests neither a
compsognathid nor an alvarezsauroid identification is well supported
and more study is needed. Agnolin et al. (2022) have since
contested Choiniere et al.'s characters, and concluded "there are some
features reminiscent to ornithomimosaurs and different from those of
alvarezsaurs."
References- Choiniere, Clark, Xu and Han, 2009. A new basal alvarezsaur
from the Shishugou Formation. Journal of Vertebrate Paleontology. 29(3), 77A.
Choiniere, 2010. Anatomy and systematics of coelurosaurian theropods from the
Late Jurassic of Xinjiang, China, with comments on forelimb evolution in Theropoda.
PhD thesis. George Washington University. 994 pp.
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal alvarezsauroid theropod
from the Early Late Jurassic of Xinjiang, China. Science. 327, 571-574.
Alifanov and Saveliev, 2011. Brain structure and neurobiology of alvarezsaurians
(Dinosauria), exemplified by Ceratonykus oculatus (Parvicursoridae) from
the Late Cretaceous of Mongolia. Paleontological Journal. 45(2), 183-190.
Choiniere, Norell and Dyke, 2011. The anatomy of the parvicursorine braincase
and its implications for alvarezsauroid systematics and evolution. Journal of
Vertebrate Paleontology. Program and Abstracts 2011, 88.
Lee and Worthy, 2011. Likelihood reinstates Archaeopteryx as a primitive bird. Biology Letters. 8(2), 299-303.
Choiniere, Clark, Norell and Xu, 2014. Cranial osteology of Haplocheirus
sollers Choiniere et al., 2010 (Theropoda: Alvarezsauroidea). American Museum
Novitates. 3816, 44 pp.
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.
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
Agnolin, Lu, Kundrat and Xu, 2022 (online 2021). Alvarezsaurid
osteology: New data on cranial anatomy. Historical Biology. 34(3),
443-452.
Choiniere, Clark and Xu, in prep.. The anatomy of Haplocheirus sollers.
"Ubirajara" Smyth, Martill, Frey, Rivera-Sylva and Lenz, 2020 online
"U. jubatus" Smyth, Martill, Frey, Rivera-Sylva and Lenz, 2020 online
Aptian, Early Cretaceous
Nova Olinda Member, Crato Formation, Brazil
Material- (SMNK PAL 29241) axis
(~12.8 mm), third cervical vertebra, fourth cervical vertebra, fifth
cervical vertebra, sixth cervical vertebra, seventh cervical vertebra,
eighth cervical vertebra, ninth cervical vertebra, tenth cervical
vertebra, partial cervical rib, first dorsal vertebra, second dorsal
vertebra, third dorsal vertebra, fourth dorsal neural arch, fifth
dorsal vertebra, sixth dorsal vertebra, partial seventh dorsal
vertebra, eighth dorsal neural arch, ninth dorsal neural arch, tenth
dorsal neural arch, partial eleventh dorsal vertebra, twelfth dorsal
vertebra, thirteenth dorsal vertebra, partial dorsal ribs, gastralia,
fused first and second sacral vertebrae, scapulae (68 mm), coracoids,
humerus (84 mm), radius (53 mm), ulna (58 mm), metacarpal I, phalanx
I-1, incomplete metacarpal II, incomplete phalanx II-1, phalanx II-2
(28 mm), manual ungual II (18 mm), incomplete metacarpal III, phalanx
III-1 (8 mm), phalanx III-2 (O10 mm), manual ungual III (14 mm), manual
claw sheaths, follicles, body feathers
Diagnosis- (after Smyth et al.,
2020) dorsal margins of the sacral neural spine tips are between 15 and
27% longer than their base; humerus (h) ~25% longer than the scapula
(s), with the ratio s/h being 0.81.
Comments- The specimen was
exported from Brazil in 1995 with an Article In Press published online
on December 13 2020. Due to legal issues regarding this export,
the pdf status was changed to "TEMPORARY REMOVAL" on December 21, with
the comment "The publisher regrets that this article has been
temporarily removed. A replacement will appear as soon as possible
in which the reason for the removal of the article will be specified,
or the article will be reinstated." On September 25 2021 this was
changed to "WITHDRAWN" with the comment "This article has been
withdrawn at the request of the editor. The Publisher apologizes for
any inconvenience this may cause." Given the facts it was never
printed in the paper version of the journal (which would have not been
until 2021 in any case) and the included lsid
zoobank.org:act:9467530F-3807-4B95-BCE4-28776E811182 does not generate
a search result on ZooBank as of 2-25-2021 (ICZN Article
8.5.3. states names published electronically must "be registered in the
Official Register of Zoological Nomenclature (ZooBank) (see Article
78.2.4) and contain evidence in the work itself that such registration
has occurred"), the name is here considered a nomen nudum.
Smyth et al. (2020 online) identify two subparallel structures as broad
monofilamentous feathers equivalent to the 'elongated broad filamentous
feathers (EBFFs)' Xu et al. 2009 reported in Beipiaosaurus.
Yet the latter are ~5% of the length of the axial centrum compared to
~35% in "Ubirajara" and lack the midline ridge found in the latter
taxon's structures. Instead, they resemble the rachis-dominated
retrices of basal pygostylians in length (~150 mm), having a rachidial
ridge and being paired. While probably not associated with the
tail of "Ubirajara", these could belong to a fortuitously placed basal
pygostylian whose body (if it remained) would be placed in the large
mass of pseudomorphosed adipocere at the center of the specimen.
Or the retrices may have been left behind (in the adipocere?) as
happened many times in Myanmar amber where pairs are preserved (Xing et
al., 2018). Basal pygostylians were common in the Aptian and
present in the Nova Olinda Member, where the juvenile Cratoavis
holotype has paired retrices 80 mm long. Or they may be
separately evolved in "Ubirajara" after all despite the absence of
other examples outside Paraves.
Smyth et al. use Brusatte's TWiG matrix to recover "Ubirajara" as a compsognathid sister to Compsognathus plus Sinosauropteryx. Entering it into Hartman et al.'s maniraptoromorph matrix results in a a sister group relationship with Haplocheirus that shares absent cervical pleurocoels and a flexor fossa on manual phalanx I-1, with both genera being compsognathids.
References-
Xing, Cockx, McKellar and O'Connor, 2018. Ornamental feathers in
Cretaceous Burmese amber: Resolving the enigma of rachis-dominated
feather structure. Journal of Palaeogeography. 7, 13.
Smyth, Martill,
Frey, Rivera-Sylva and Lenz, 2020 online. A maned theropod dinosaur
from Gondwana with elaborate integumentary structures. Cretaceous
Research. Withdrawn Article in Press. DOI: 10.1016/j.cretres.2020.104686
Xunmenglong Xing, Miyashita, Wang, Niu and Currie, 2020
= "Xunmenglong" Xing, Miyashita, Wang, Niu and Currie, online 2019
X. yinliangis Xing, Miyashita, Wang, Niu and Currie, 2020
= "Xunmenglong yinliangis" Xing, Miyashita, Wang, Niu and Currie, online 2019
Late Hauterivian, Early Cretaceous
Sichakou Sedimentary Member of the Huajiying Formation, Hebei, China
Holotype- (YLSNHM-00005)
fragmentary thirteenth dorsal vertebra?, incomplete sacrum (~18.6 mm),
first to eleventh caudal vertebrae, proximal chevrons, ilia (one
partial, one fragmentary), ischial fragment, femora (one proximal, one
distal), tibiae (one incomplete; 50.74 mm), incomplete fibulae (50.03
mm), astragali (5.5, 6.89 mm wide), calcanea, distal tarsals III,
distal tarsals IV, metatarsal I (7.29 mm), phalanges I-1 (6.7 mm),
proximal pedal ungual I, metatarsals II (one incomplete; 25, 28.88 mm),
phalanges II-1 (one partial; 8.42, 7.15 mm), phalanges II-2 (7.58, 6.91
mm), pedal unguals II (7.3, 6.82 mm), metatarsals III (26.78 mm),
phalanges III-1 (7.82, 8.84 mm), phalanges III-2 (7.74, 8.68 mm),
phalanges III-3 (6.22, 7.28 mm), pedal unguals III (8.73, 10 mm),
metatarsals IV (24.44, 26.15 mm), phalanges IV-1 (5.74, 4.54 mm),
phalanges IV-2 (3.74, 3.79 mm), phalanges IV-3 (~3.08 mm), phalanges
IV-4 (~3.08 mm), pedal unguals IV (6.41, 6.66 mm), metatarsals V (94.9
mm), pedal claw sheaths
Diagnosis- (after Xing et al.,
2020) vertical neural spine of each proximal caudal vertebrae
dorsoventrally taller than (or as tall as) associated anteroposterior
centrum length [only valid for c1-5]; pedal digit III (excluding
ungual) subequal in length to metatarsal III [93%].
Other diagnoses- Xing et al.
(2020) listed sacral and proximal caudal centra dorsoventrally taller
than (or as tall as) anteroposteriorly long as being diagnostic, but
the sacral vertebrae are too poorly preserved to determine centrum
height while the caudal centra are all longer than tall (e.g. c3 15%
longer than tall). They also listed preacetabular process as long as
postacetabular process, but this is not evident from the specimen where
the pubic peduncle and anterior and ventral extent of the preacetabular
process is unpreserved. Similarly, when they list "dorsal outline
straight and anterior margin round in lateral view" as a diagnostic
character of the ilium, the dorsal margin is convex and the anterior
and ventral edges of the preacetabular process are unpreserved so that
its shape is unknown. A further supposed character "tibia markedly
longer than predicted by an allometric trend among theropods
(tibia/femur length ratio is 1.53)", is seemingly due to incorrectly
estimating the femoral length, as the proximal end of the left femur
extends under the ilium and probably the sacrum as well. The
ossification of distal tarsal II is another listed diagnostic feature,
but all saurischians lack this and given the preservation it is
probably the medial portion of distal tarsal III in both feet. Xing et
al. also list pedal phalanx IV-4 markedly longer than IV-3 as being
diagnostic, but neither illustration of either pes shows this, which
also seems to be the case for the photos. Adjusted measurements for
IV-3 and IV-4 (excluding proximal lips) of the left pes are given above
based on the photo, whereas the authors list 2.96 and 3.88 mm
respectively. The right pes is in dorsal view so cannot be measured as
precisely (Xing et al.'s measurements are 2.89 and 3.62 mm).
Comments- The specimen was discovered in 2013, and described and named by Xing et al. (2019). Unfortunately this was
in a journal pre-proof posted October 25 2019, but was electronic
and had no mention of ZooBank, so was a nomen nudum (ICZN Article
8.5.3. states names published electronically must "be registered in the
Official Register of Zoological Nomenclature (ZooBank) (see Article
78.2.4) and contain evidence in the work itself that such registration
has occurred") until officially published in March 2020.
As Xing et al. (2020) noted, "YLSNHM-00005 was used by a private
collector as a component to assemble a chimaeric small theropod. The
slabs of the chimaera represent at least three different animals, but
were disassembled at YLSNHM under the supervision of the authors."
"The right femur of unknown origin was fitted in the original position
of the pubes by the collector. This retrofitted femur is nearly
identical in size to the left femur of YLSNHM-00005, and may belong to
the same animal. However, the planes of breakage did not match
perfectly between the retrofitted femur and the proximal portion of the
right femur that is preserved on the main slab. To be conservative,
this femur is treated separately from the holotype until there is
compelling evidence for its identity."
Although the authors identify their first illustrated largely preserved
vertebra as the first sacral, no actual structure is visiable in this
area, including the anterior and posterior rims, neurocentral suture
and fossa or rib articulation in their line drawing. Based on
comparison with Compsognathus
MNHN CNJ 79, a sacral series of five would actually begin with the
following vertebrae, which is the measurement and interpretation used
here. Its possible that one or more dorsals were incorporated
however given the uncertain anterior extent of the preacetabular
process. In general, Xing et al. describe and score much more
information than
the poor preservation allows, which in addition to features noted above
include the tall sacral neural spines
(seemingly low and backswept in the last two sacrals), damage to the
sides of the sacrals and proximal caudals being viewed as "sacral ribs
[that] appear not to have been fully ossified", any absence of
accessory caudal neural spines, and brevis fossa development.
Xing et al. added Xunmenglong to Choiniere's coelurosaur analysis and
found it to be in Compsognathidae in both the primary dataset and one
where young taxa were scored conservatively for ontogenetic
characters. When added to Hartman et al.'s maniraptoromorph analysis, it emerges as a compsognathid closest to Haplocheirus.
Reference- Xing, Miyashita,
Wang, Niu and Currie, 2020 (online 2019). A new compsognathid theropod dinosaur from
the oldest assemblage of the Jehol Biota in the Lower Cretaceous
Huajiying Formation, northeastern China. Cretaceous Research. 107, 104285.
Compsognathus Wagner, 1859
C. longipes Wagner, 1859
= Compsognathus corallestris Bidar, Demay and Thomel, 1972
Early Tithonian, Late Jurassic
Solnhofen Formation, Germany
Holotype- (BSP AS I 536) (~.86 m, .58 kg) incomplete skull (75 mm), mandibles,
hyoids, atlas, axis (8.7 mm), third cervical vertebra (9.5 mm), fourth cervical
vertebra (11 mm), fifth cervical vertebra (12.3 mm), sixth cervical vertebra
(12.7 mm), seventh cervical vertebra (12.7 mm), eighth cervical vertebra (11.3
mm), ninth cervical vertebra (10.9 mm), tenth cervical vertebra (10.9 mm), fourteen
cervical ribs, first dorsal vertebra (9.9 mm), second dorsal vertebra (9.4 mm),
third dorsal vertebra (~9.8 mm), fourth dorsal vertebra (~9.1 mm), fifth dorsal
vertebra (~9.7 mm), sixth dorsal vertebra (9.9 mm), seventh dorsal vertebra
(10.5 mm), eighth dorsal vertebra (10.2 mm), ninth dorsal vertebra (12.2 mm?),
tenth dorsal vertebra (10.75 mm), eleventh dorsal vertebra (11.4 mm), twelfth
dorsal vertebra (~11.5 mm), thirteenth dorsal vertebra (~12 mm), twenty-two
partial dorsal ribs, gastralia, third sacral vertebra , fourth sacral vertebra
(8.6 mm), fifth sacral vertebra, first caudal vertebra (10.9 mm), second caudal
vertebra (11.2 mm), third caudal vertebra (11.5 mm), fourth caudal vertebra
(11.8 mm), fifth caudal vertebra (12.1 mm), sixth caudal vertebra (12.6 mm),
seventh caudal vertebra (12.9 mm), eighth caudal vertebra (13.2 mm), ninth caudal
vertebra (13.3 mm), twenth caudal vertebra, eleventh caudal vertebra, twelfth
caudal vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra, fifteenth
caudal vertebra, ten chevrons, partial scapula (~38 mm), partial coracoids,
humeri (~38-40 mm), radii (24.7 mm), ulnae (28.5 mm), two carpals, metacarpal
I (5.85 mm), phalanx I-1 (17.6 mm), manual ungual I (10.4, 10.4 mm), metacarpal
II (13.95 mm), phalanx II-1 (7.7, 7.8 mm), phalanx II-2 (14.5, 14.45 mm), manual
ungual II (9.6, 9.7 mm), metacarpal III (13.1 mm), phalanx III-1, partial ilia,
incomplete pubes (~60 mm), ischia (~40 mm), femora (~67 mm), tibiae (87.7, 87.6mm),
fibulae (82.1 mm), astragalus?, distal tarsal IV, metatarsal I (12 mm), phalanx
I-1 (9 mm), pedal ungual I (4.5 mm), metatarsal II (50.4 mm), phalanx II-1 (15
mm), phalanx II-2 (15 mm), pedal ungual II (13 mm), metatarsal III (56 mm),
phalanx III-1 (18 mm), phalanx III-2 (15 mm), phalanx III-3 (13 mm), pedal ungual
III (13 mm), metatarsal IV (51.8 mm), phalanx IV-1 (12 mm), phalanx IV-2 (10
mm), phalanx IV-3 (10 mm), phalanx IV-4 (10 mm), pedal ungual IV (10 mm), metatarsal
V (17 mm), eichstaettisaurid skeleton
Early Tithonian, Late Jurassic
Lithographic Portlandian Limestone, France
Referred- ?(MNHN CNJ 79; holotype of Compsognathus corallestris)
(~1.4 m, 2.5 kg) incomplete skull (100 mm), incomplete mandibles, hyoids, atlas,
axis, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra,
sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra,
ninth cervical vertebra, tenth cervical vertebra, seven 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, several
dorsal ribs, gastralia, sacrum, thirty-one caudal vertebrae, thirty-one chevrons,
scapulae (51.2 mm), coracoid, furcula, humeri (56.3, 51.9 mm), radii (41 mm),
ulnae (46.4 mm), scapholunare, distal carpal I, distal carpal II, metacarpal I (6.8
mm), phalanx I-1 (21.6 mm), metacarpal II (27.3, 25.4 mm), phalanx II-1 (13.4,
13 mm), metacarpal III (22.9, 24.2 mm), phalanx III-1 (1.4 mm), phalanx III-2
(3.3 mm), ilium (77.8 mm), pubes (103.4 mm), ischia (65.8 mm), femora (108.8
mm), tibiae (131, 131.8 mm), fibulae (one partial; 124.9 mm), astragali, calcanea,
distal tarsal III, distal tarsal IV, metatarsal I (17 mm), phalanx I-1 (13.8
mm), pedal ungual I (6.6 mm), metatarsal II (72.5, 73.1 mm), phalanx II-1 (22.1
mm), phalanx II-2 (19.4, 19.7 mm), pedal ungual II (15 mm), metatarsal III (79.6,
80.9 mm), phalanx III-1 (24.6, 23.6 mm), phalanx III-2 (19.5, 20.8 mm), proximal
phalanx III-3, metatarsal IV (72.5, 73.2 mm), phalanx IV-1 (15.2, 16 mm), phalanx
IV-2 (14.9, 12.3 mm), phalanx IV-3 (10.9 mm), phalanx IV-4 (8.8 mm), pedal ungual
IV, metatarsal V (24.5 mm), skin impressions (Bidar, Demay and Thomel, 1972)
Diagnosis- (after Peyer, 2006) ventral process at the posterior end of
premaxillary body; opisthocoelous cervical vertebrae; metacarpal I less than
one third as long as metacarpal II; no fourth trochanter on femur; hallux ends
at or below the distal end of phalanx 1 of digit II.
Comments- The genus and its type species were first named and briefly
described by Wagner in 1859, then more extensively described by him in 1861,
the date usually given for these taxa.
While MNHN CNJ 79 was originally described as a new species, C. corallestris,
until Ostrom (1978) synonymized it with C. longipes. Recently, Rauhut
and Foth (2014) state newly discovered casts of the holotype indicate it was
more complete when found and indicates "it is not the same taxon as the
French Compsognathus."
Dames (1884) described three metapodials and a proximal phalanx (HMN coll.)
from the Solnhofen Formation, which was questionably referred to Compsognathus
by Huene (1925). However, Ostrom (1978) showed that the shortest metapodial
is too short to be a Compsognathus metatarsal II (which is the shortest
of its main three metatarsals) and that the phalanx associated with it is too
long to be II-1. These may not be theropod, and may not even be metatarsals.
Gauthier and Gishlick (2000) reinterpreted the manus of Compsognathus.
"Metacarpal I" is really phalanx I-1. The mystery element above the
skull is a very short metacarpal I. There is a collateral ligament pit on metacarpal
III, but no preserved phalanges. Thus, there may have been a third digit or
not.
References- Wagner, 1859. �ber einige im lithographischen Schiefer
neu aufgefundene Schildkr�ten und Saurier: Gelehrte Anzeigen der Bayerischen
Akademie der Wissenschaften. 49, 553.
Wagner, 1861. Neue Beitrige zur Kenntis der urweltlichen Fauna des lithographischen
Schiefers. V. Compsognathus longipes Wagner. Abhandlungen der Bayerischen Akademie der Wissenschaften. 9,
30-38.
Dames, 1884. Uber Metatarsen eines Compsognathus - ahnlichen Reptils
von Solnhofen. Sitzungsberichte der Gesellschaft Naturforschender
Freunde zu Berlin. 1884, 179-180.
Huene, 1925. Eine neue Rekonstrucktion von Compsognathus longipes. Zentralblatt f�r Mineralogie, Geologie und Pal�ontologie. Jahrgang 1925, Abteilung B(5), 157-160.
Bidar, Demay and Thomel, 1972. Compsognathus corallestris, nouvelle espece
de dinosaurien theropode du Portlandien de Canjuers (Sud-Est de la France).
Annales du Mus�um d’Histoire Naturelle de Nice. 1, 9-40.
Ostrom, 1978. The osteology of Compsognathus longipes. Zitteliana. 4, 73-118.
Michard, 1991. Description du Compsognathus (Saurischia, Theropoda) de
Canjuers (Jurassique sup�rieur du Sud-est de la France), position phylog�n�tique,
relation avec Archaeopteryx et implications sur l’origine th�ropodienne
des oiseaux. PhD thesis. Mus�um National d’Histoire Naturelle. 328 pp.
Gauthier and Gishlick, 2000. Re-examination of the manus of Compsognathus
and its relevance to the original morphology of the coelurosaur manus. Journal
of Vertebrate Paleontology. 20(3), 43A.
Peyer, 2003. A complete redescription of the French Compsognathus with
special consideration of the anatomy of the hand. Journal of Vertebrate Paleontology.
23(3), 87A.
Peyer, 2004. The phylogenetic relationship of the French Compsognathus
within the Compsognathidae and coelurosaurs. Journal of Vertebrate Paleontology.
24(3), 144A-145A.
Peyer, 2006. A reconsideration of Compsognathus from the Upper Tithonian
of Canjuers, southeastern France. Journal of Vertebrate Paleontology. 26(4),
879-896.
Gishlick and Gauthier, 2007. On the manual morphology of Compsognathus longipes
and its bearing on the diagnosis of Compsognathidae. Zoological Journal of the
Linnean Society. 149, 569-581.
Conrad, 2014. The lizard (Squamata) in Compsognathus (Theropoda) is a
new species, not Bavarisaurus. Journal of Vertebrate Paleontology, Program
and Abstracts. 111.
Rauhut and Foth, 2014. New information on the theropod dinosaurs from the Late
Jurassic lithographic limestones of Southern Germany. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 212.
C. sp. (Zinke, 1998)
Kimmeridgian, Late Jurassic
Guimarota Formation, Portugal
Material- (IPFUB GUI D 28-65, 98, 103, 105-110, 112, 113) 49 teeth (~1.71
mm)
Diagnosis- differs from C. longipes in that posterior teeth have
serrations on mesial carinae.
Reference- Zinke, 1998. Small theropod teeth from the Upper Jurassic
coal mine of Guimarota (Portugal). Palaontologische Zeitschrift. 72, 179-189.
Aorun Choiniere, Clark, Forster, Norell,
Eberth, Erickson, Chu and Xu, 2013
= "Farragochela" Choiniere, 2010
A. zhaoi Choiniere, Clark, Forster, Norell, Eberth, Erickson,
Chu and Xu, 2013
= "Farragochela zhaoi" Choiniere, 2010
Early Oxfordian, Late Jurassic
Wucaiwan, Middle Shishugou Formation, Xinjiang, China
Holotype- (IVPP V15709) (4 kg; <1 year old juvenile) incomplete skull (~49
mm), incomplete mandibles, partial sclerotic rings, hyoids, anterior cervical
vertebra (14 mm), posterior(?8-10) dorsal vertebra (10 mm), proximal caudal
vertebra, proximal caudal vertebra, partial proximal caudal vertebra, incomplete
ulna, scapholunare, distal carpal I, metacarpal I, phalanx I-1 (21.5 mm), manual
ungual I (19.2 mm), metacarpal II (21.9 mm), phalanx II-1 (13.4 mm), phalanx
II-2 (22 mm), manual ungual II (~15 mm), incomplete metacarpal III (~19.9 mm),
phalanx III-3 (15 mm), partial manual ungual III, distal pubes, tibiae (one
incomplete, one proximal; 123.3 mm), partial fibula, astragalus, distal tarsal
III, distal tarsal IV, metatarsals I (6.8, 8.7 mm), phalanges I-1 (3, 3.8 mm),
pedal ungual I (6.4 mm), metatarsals II (one incomplete, one distal), phalanges
II-1 (17.1, 16 mm), phalanges II-2 (15.5, 14.7 mm), pedals ungual II (15.5 mm),
metatarsals III (one incomplete, one distal), phalanges III-1 (17.8, 19 mm),
phalanges III-2 (14.4, 14.2 mm), phalanx III-3 (13.4 mm), partial pedal unguals
III, metatarsals IV (one incomplete, one distal), phalanges IV-1 (12.7 mm),
phalanges IV-2 (10.6, 10.7 mm), phalanges IV-3 (9.4 mm), phalanx IV-4 (6.9 mm),
partial pedal ungual IV
Diagnosis- (after Choiniere et al., 2013) large maxillary fenestra occupying
most of antorbital fossa; maxillary teeth with very small, apically directed
serrations restricted to distal carinae; weakly opisthocoelous cervical centra;
heterogeneous manual ungual morphology with large, recurved ungual I and smaller
unguals II and III that have straight ventral surfaces; tibia with mediolaterally
narrow, proximodistally tall articular groove accepting the astragalar ascending
process that is only developed on anterolateral margin; ascending process of
astragalus low and restricted to lateral side of tibia.
Comments- The holotype was discovered in 2006 and initially described
by Choiniere (2010) in his thesis as "Farragochela", then published online on May 3 2013 with a different
name. As the authors included a Zoobank registration number, the name was valid
under new ICZN rules despite the physical version being unpublished until 2014.
Choiniere et al. recover Aorun as the most basal maniraptoromorph
without ontogenetic consideration, and as a coelurid in basal Maniraptora if
ontogeny is taken into account, using a version of Choiniere's matrix. Xu et al. (2018) recently proposed Aorun
is the basalmost alvarezsauroid based on four unambiguous
synapomorphies- dorsoventrally flattened internarial bar; collateral
ligament fossae on metacarpal I absent; manual digit I bearing large
ungual and all other unguals distinctly smaller; proximodistally
oriented step-like ridge on anterior surface of tibia that braces
astragalar ascending process. Most recently, Hartman et al.
(2019) recovered Aorun as a
compsognathid, and found it takes 17 more steps to move to
Alvarezsauroidea. This is similar to Cau's (2018) results, where Aorun and Compsognathus
are both non-tyrannoraptoran coelurosaurs and can form a
Compsognathidae with no extra steps, but it takes 9 extra steps to
force an alvarezsauroid Aorun. Notably, even Xu et al.'s matrix only needs 4 steps to move Aorun
to Compsognathidae. Of Xu et al.'s proposed alvarezsauroid
characters, the flattened internarial bar was used by Hartman et al.
and is shared with the controversially compsognathid Haplocheirus. Aorun
actually seems to have a weak medial ligament pit on metacarpal I
(Choiniere et al., 2013: Fig. 15A). The ratio between manual
ungual I and II lengths (1.28) in Aorun is not more than Sciurumimus (~1.29) or Sinosauropteryx
(~1.79). Finally, the ridge bracing the astragalar ascending
process is primitive for maniraptoriforms and also present in Compsognathus and Tugulusaurus. Thus a compsognathid identity is the best supported given proposed characters.
References- Choiniere, 2010. Anatomy and systematics of coelurosaurian
theropods from the Late Jurassic of Xinjiang, China, with comments on forelimb
evolution in Theropoda. PhD Thesis. George Washington University. 994 pp.
Choiniere, Clark, Forster, Norell, Eberth, Erickson, Chu and Xu, 2014
(online 2013). A juvenile specimen of a new coelurosaur (Dinosauria:
Theropoda) from the Middle-Late Jurassic Shishugou Formation of
Xinjiang, People's Republic of China. Journal of Systematic
Palaeontology. 12(2), 177-215.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25.
Xu, Choiniere, Tan, Benson, Clark, Sullivan, Zhao, Han, Ma, He, Wang,
Xing and Tan, 2018. Two Early Cretaceous fossils document transitional
stages in alvarezsaurian dinosaur evolution. Current Biology. 28, 1-8.
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
Sciurumimus Rauhut, Foth,
Tischlinger and Norell, 2012
S. albersdoerferi Rauhut, Foth, Tischlinger and Norell, 2012
Late Kimmeridgian, Late Jurassic
Painten Formation, Bavaria, Germany
Holotype- (BMMS BK 11) (719 mm; juvenile) skull (79 mm), sclerotic ring,
mandibles (73.2 mm), hyoids, (cervical series 69 mm) atlas, axis, third cervical
vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth cervical
vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical
vertebra, tenth cervical vertebra, cervical ribs, (dorsal series 102 mm) 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, (sacrum 37.3 mm) first sacral centrum, fourth sacral ventrum,
fifth sacral centrum, about sixty caudal vertebrae (432 mm), chevrons, scapula,
coracoids, clavicles, humeri (26.8 mm), radii (17 mm), ulnae, distal carpals
I, metacarpals I, phalanges I-1, manual unguals I, metacarpals II (11 mm), phalanges
II-1, phalanges II-2, manual unguals II, metacarpals III, phalanges III-1, phalanges
III-2, phalanges III-3, manual unguals III, ilium, pubes, ischia, femora (50.6
mm), tibiae (54.2 mm), fibulae, calcaneum, distal tarsal IV, metatarsal I, phalanx
I-1, pedal ungual I, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals
II, metatarsals III (32.1 mm), phalanges III-1, phalanges III-2, phalanges III-3,
pedal unguals III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges
IV-3, phalanges IV-4, pedal unguals IV, pedal claw sheaths, metatarsals V, hindlimb
muscle, skin impressions, feathers
Diagnosis- (after Rauhut et al., 2012) axial neural spine symmetrically
hatchet-shaped in lateral view; posterior dorsal neural spines with rectangular
edge anteriorly and lobe-shaped dorsal expansion posteriorly; anterior margin
of ilium with semioval anterior process in its dorsal half.
Comments- In 2011 a complete juvenile theropod skeleton was announced
at the Munich Show Mineralientage M�nchen. Rauhut and Foth (2011) had given
a presentation on this specimen the previous month, and the next year Rauhut
et al. (2012) described it as Sciurumimus albersdoerferi. Rauhut et al. scored it for three analyses- Smith et al.'s (2008) found it as an orionidan
more closely related to Monolophosaurus and Avetheropoda; Choiniere et
al.'s (2010) found it as an orionidan in a polytomy with Afrovenator,
spinosauroids and Monolophosaurus+Avetheropoda; Benson et al. (2010)
found it as a basal megalosaurid. This is interesting as the specimen is extremely
similar to the supposed basal coelurosaur Juravenator, though both are
juveniles. When Juravenator is added to the former two matrices with
Sciurumimus (it was not added to Benson et al.'s inexplicably), it ends
up sister to Sciurumimus in the same positions described above outside
Coelurosauria. Yet when Juravenator is added and Sciurumimus is
not, Juravenator is a coelurosaur as usual. Cau (online, 2012) included Sciurumimus in his much larger
analysis and found it to be a non-tyrannoraptoran coelurosaur by Tugulusaurus and Zuolong, with Juravenator a maniraptoromorph. This was eventually published in Godefroit et al. (2013). Hartman et al. (2019) examined the Sciurumimus
issue in depth and found numerous inaccurate scorings in Smith et
al.'s, Choiniere et al.'s and Benson et al.'s matrices were to blame,
along with a few misscorings of Sciurumimus. When these are corrected in each matrix, Sciurumimus and Juravenator can be compsognathids. This coincides with Hartman et al.'s recovery of Sciurumimus in Compsognathidae, requiring 7 steps to move to Megalosauroidea. Sciurumimus
does however possess several characters which are similar to some basal
tetanurines and are thus autapomorphies if it is a compsognathid- axial
pleurocoels absent, no posteroventral coracoid process, cuppedicus
fossa absent, obturator process not proximally defined. Hartman
et al. included all but the second character in their analysis.
References- Smith, Makovicky, Agnolin, Ezcurra, Pais and Salisbury, 2008. A Megaraptor-like
theropod (Dinosauria: Tetanurae) in Australia: Support for faunal
exchange across eastern and western Gondwana in the Mid-Cretaceous.
Proceedings of the Royal Society B. 275(1647), 2085-2093.
Benson, Carrano and Brusatte, 2010 (online 2009). A new clade of archaic large-bodied
predatory dinosaurs (Theropoda: Allosauroidea) that survived to the
latest Mesozoic. Naturwissenschaften. 97(1), 71-78.
Choiniere, Clark, Forster and Xu, 2010. A basal coelurosaur
(Dinosauria: Theropoda) from the Late Jurassic (Oxfordian) of the
Shishugou Formation in Wucaiwan, People's Republic of China. Journal of
Vertebrate Paleontology. 30(6), 1773-1796.
Rauhut and Foth, 2011. New information on Late Jurassic theropod
dinosaurs from southern Germany. IV Congresso Latinoamericano Paleontologia
de Vertebrados.
Cau, online 2012. http://theropoda.blogspot.com/2012/07/sciurumimus-albersdoerferi-rauhut-et-al.html
Rauhut, Foth, Tischlinger and Norell, 2012. Exceptionally preserved juvenile
megalosauroid theropod dinosaur with filamentous integument from the Late Jurassic
of Germany. Proceedings of the National Academy of Sciences. 109(29), 11746-11751.
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.
Foth, Haug, Haug, Tischlinger and Rauhut, 2014. New details on the integumental
structures in the juvenile megalosauroid Sciurumimus albersdoerferi from
the Late Jurassic of Germany using different auto-fluorescence imaging technique.
Journal of Vertebrate Paleontology. Program and Abstracts 2014, 131-132.
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 compsognathid (Ji and JI, 1997)
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Beds of Yixian Formation, Liaoning, China
Material- (NGMC 2124) (1.06 m) incomplete skull (113 mm), mandibles,
hyoids, cervical vertebrae, dorsal vertebrae, dorsal ribs, sacrum, thirty-eight
caudal vertebrae, chevrons, incomplete scapula, incomplete coracoid, forelimb
elements, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx
II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx
III-2, phalanx III-3, manual ungual III, ilia (91 mm), pubes (96 mm), ischia
(56 mm), femora (108 mm), tibiae (151 mm), fibulae, calcanea, metatarsals I,
phalanges I-1, pedal unguals I, metatarsals II, phalanges II-1, phalanges II-2,
pedal unguals II, metatarsals III (90 mm), phalanges III-1, phalanges III-2,
proximal phalanx III-3, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges
IV-3, phalanges IV-4 (one partial), pedal unguals IV (one partial), metatarsals
V, feathers, Sinobaatar mandible, Zhangheotherium mandibles (Ji and Ji, 1997)
Comments- This was described as a new specimen of Sinosauropteryx
prima by Ji and Ji (1997). Longrich (DML, 2000) noted it differed from Sinosauropteryx
in several characters. He published an abstract in 2002 detailing his reasoning,
proposing NGMC 2124 was a compsognathid/coelurid-grade coelurosaur, while Sinosauropteryx
was a very basal coelurosaur or even a basal carnosaur. Gishlick and Gauthier
(2007) refer to the specimen as Sinosauropteryx? sp. and figure the manus.
Ji et al. (2007) also agree it does not belong in Sinosauropteryx.
Hartman et al. (2019) were the first authors to include NGMC 2124 in a
phylogenetic analysis and found it resolves as a compsognathid closest
to Aorun then Sciurumimus, widely separated from Sinosauropteryx. Only three steps are needed to make NGMC 2124 and Sinosauropteryx sister taxa, however, suggesting this is still quite possible.
Hurum et al. (2006) determined "in the abdomen of a specimen (CAGS GMV 2124) of the feathered dinosaur Sinosauropteryx prima,
three lower jaws of mammals have been preserved, rather than two as
previously mentioned by Ackerman (1998). Two of them belong to Zhangheotherium, the third to the multituberculate Sinobaatar..."
References- Ji and Ji, 1997. Advances in the study of the avian Sinosauropteryx
prima. Chinese Geology. 242, 30-32.
Ackerman, 1998. Dinosaurs take wing. National Geographic. 194(1), 189-192.
Longrich, DML 2000. https://web.archive.org/web/20201115172810/http://dml.cmnh.org/2000Apr/msg00300.html
Currie and Chen, 2001. Anatomy of Sinosauropteryx prima from Liaoning,
northeastern China. Canadian Journal of Earth Science. 38(12), 1705-1727.
Longrich, 2002. Systematics of Sinosauropteryx. Journal of Vertebrate
Paleontology. 22(3), 80A.
Hurum, Luo and Kielan-Jaworowska, 2006. Were mammals originally venomous? Acta Palaeontologica Polonica. 51(1), 1-11.
Gishlick and Gauthier, 2007. On the manual morphology of Compsognathus longipes
and its bearing on the diagnosis of Compsognathidae. Zoological Journal of the
Linnean Society. 149, 569–581.
Ji, Gao, Liu, Meng and Ji, 2007. New material of Sinosauropteryx (Theropoda:
Compsognathidae) from Western Liaoning, China. Acta Geologica Sinica. 81(2),
177-182.
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
Maniraptora sensu Choiniere, Xu, Clark, Forster, Guo and Han, 2010
Definition- (Ornitholestes hermanni + Archaeopteryx lithographica)
(modified)
"Fukuivenator" Azuma, Xu, Shibata,
Kawabe, Miyata and Imai, 2016
"F. paradoxus" Azuma, Xu, Shibata, Kawabe, Miyata and Imai, 2016
Middle-Late Aptian, Early Cretaceous
Kitadani Dinosaur Quarry, Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Material-
(FPDM-V8461) (~2.5 m, ~25 kg subadult) maxillae (one partial),
lacrimals (one partial), jugal, frontals, parietals (26.2, 28.8 mm),
quadrate (40.3 mm), incomplete braincase, palatine, ectopterygoids,
dentary fragment, three teeth, atlantal neural arches,
axis, third cervical vertebra, fifth cervical vertebra, sixth cervical
vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth
cervical vertebra, eleventh cervical vertebra, nine partial to complete
cervical ribs (cr9 41.2, 41.7 mm), incomplete first dorsal vertebra,
second dorsal centrum (23.7 mm), third dorsal vertebra, fourth dorsal
vertebra, fifth dorsal vertebra, incomplete sixth dorsal vertebra,
seventh dorsal vertebra, eighth dorsal vertebra, ninth dorsal vertebra,
tenth dorsal vertebra, eleventh dorsal vertebra, fifteen fragmentary to
incomplete dorsal ribs, several gastralial fragments, incomplete fused
first-fourth sacral vertebrae (101.0 mm), fifth sacral vertebra, first
caudal vertebra, second caudal vertebra, incomplete third caudal
vertebra, fourth caudal neural arch fragment, fifth caudal vertebra,
sixth caudal vertebra, seventh 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, eigthteenth caudal vertebra, nineteenth caudal
vertebra, twentieth caudal vertebra, twenty-first caudal vertebra,
partial twenty-second caudal vertebra, twenty-third caudal vertebra,
twenty-fourth caudal vertebra, twenty-fifth caudal vertebra,
twenty-sixth caudal vertebra, twenty-seventh caudal vertebra,
twenty-eighth caudal vertebra, twenty-ninth caudal vertebra, thirtieth
caudal vertebra, pygostyle, incomplete ninth chevron, incomplete
eleventh chevron, fourteenth chevron (19.3 mm), partial fifteenth
chevron, sixteenth chevron (15.0 mm), partial seventeenth chevron,
scapulae (one incomplete; 132.9 mm), coracoids (one incomplete),
incomplete humeri (one incomplete, one distal), incomplete radiii (~107
mm), ulnae (one incomplete, one proximal), metacarpals I (31.1, 31.1
mm), phalanx I-1 (47.0 mm), manual ungual I (35.4 mm), metacarpal II
(62.8 mm), phalanx II-1 (42.3 mm), phalanges II-2 (46.9, 46.5 mm),
manual ungual II (44.5 mm), metacarpal III (~50.4 mm), phalanges III-1
(18.8, 20.3 mm), phalanx III-2 (16.7 mm), phalanx III-3 (36.1 mm),
manual ungual III (37.0 mm), incomplete pubis, incomplete femora (~190
mm), tibiae (one incomplete, one partial), partial fibula, astragali
(31.5, 32.6 mm), calcaneum, metatarsals I (28.8 mm), phalanx I-1, pedal
unguals I (16.9 mm), distal tarsal III, metatarsals II (one incomplete;
180.6 mm), phalanges II-1 (33.2, 32.1 mm), phalanx II-2 (29.3 mm),
partial pedal ungual II, metatarsals III (one incomplete; 118.3 mm),
phalanges III-1 (32.3, 31.9 mm), phalanx III-2 (24.7 mm), phalanges
III-3 (24.2, 24.3 mm), pedal ungual III (19.9 mm), metatarsal IV (105.1
mm), phalanges IV-1 (23.7, 23.5 mm), phalanx IV-2 (20.2 mm), phalanges
IV-3 (18.1, 17.8 mm), phalanges IV-4 (17.7, 17.1 mm), pedal unguals IV
(17.1, 16.8 mm), metatarsal V (34.9 mm)
Diagnosis- (after Azuma et al., 2016) large oval
lacrimal pneumatic recess; elongate
tubercle on posterior surface of basal tuber; highly heterodont dentition featuring
robust unserrated teeth including small spatulate anterior teeth, large and
posteriorly curved middle teeth, and small and nearly symmetrical posterior
teeth; cervical vertebrae with complex lamina system surrounding neural canal
resulting in deep and wide grooves for interspinous ligaments and additional
deep sockets; anterior cervical vertebrae with interprezygapophyseal, postzygadiapophyseal,
prezygadiapophyseal, and interpostzygapophyseal laminae connecting to each other
to form extensive platform; eighth and ninth cervical vertebrae with transversely
bifid neural spines; dorsal, sacral and proximal caudal vertebrae with strongly
laterally curved hyposphene and centropostzygapophyseal laminae that, together
with postzygapophyseal facet, form socket-like structure for receiving the prezygapophyses;
caudal zygapophyseal facets expanded to be substantially wider than zygapophyseal
processes; mid caudal vertebrae with transversely and distally bifid prezygapophyses
(also in eudromaeosaurs).
(after Hattori et al., 2021) large maxillary fenestra expanded well
dorsally above suprantral strut; jugal anterior process with thick and
rounded dorsal margin continuous with lateral surface; bifid posterior
end of ectopterygoid for contact with pterygoid.
Other diagnoses- Azuma et al. (2016) listed "unusually
large external naris (slightly smaller than antorbital fenestra in
dorsoventral height)" as diagnostic, but this was based on the
misidentified right maxilla's antorbital fenestra being mistaken for an
external naris. Another proposed character "large premaxillary
[sic] fenestra subequal in size to maxillary fenestra" is untrue as the
photo indicates the promaxillary fenestra is ~43% the length of the
maxillary fenestra, though both have some broken margins. The character
"lacrimal with a distinct groove on lateral surface of anterior process
and a ridge on lateral surface of descending process" is not valid
given the reidentification of the anterior process as the ventral
process and vice versa. They also list "postorbital frontal
process with T-shaped cross section and laterally-flanged squamosal
process", but does the first part indicate the postorbital process of
the frontal or the frontal (i.e. anterior) process of the postorbital?
Similarly, the squamosal process of the postorbital has no obvious
flange in the figure. They list "dorsoventrally bifurcated sacral ribs"
as being diagnostic, but this is true in e.g. the middle four sacrals
of ornithomimosaurs (e.g. Deinocheirus,
Gallimimus) and Suzhousaurus and the fourth sacral of Zuolong.
Comments-
Discovered in Summer 2007, and initially announced as a dromaeosaurid
(Anonymous, 2009; Shibata and Azuma, 2010), this was later described by
Azuma et al. on February 23 2016 as a new taxon of basal
coelurosaur. However, this paper has no mention of ZooBank and as
of March 3 2022
"Fukuivenator" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Fukuivenator paradoxus" Azuma
et al., 2016 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
The initial description of "Fukuivenator" included several problematic
interpretations and assertions (Mortimer, 2016 online), many cleared up
in the detailed redescription by Hattori et al. (2021). Contra
Azuma et al., it cannot be determined if the premaxillary subnarial
process extends posterior to the external naris like in dromaeosaurids,
as the anterior maxillary surface shows no differentiation of
premaxillary vs. nasal sutures and the tip of the process itself (and
indeed the entire premaxilla) is missing. The text claims the lacrimal
is T-shaped, but the figure shows the posterodorsal process is
unpreserved, and the character is coded scored in their matrix. Hattori
et al. specify this supposed left lacrimal is actually the right
lacrimal, with the apparent ventral process actually being an anterior
process plus tip of the dorsal maxillary process. The
posterodorsal process of the reinterpreted lacrimal is still very short
regardless. Although the authors claim the frontals have
dromaeosaurid-like anterolateral notches, Hattori et al.'s figures show
they do not. The fenestra labeled "IX?" in figure 4 is the otic
fenestra, the basipterygoid processes are labeled as laterosphenoids
twice in that figure, and the supposed medial eustachian foramina are
the paired foramina of the basisphenoid recess, all corrected by
Hattori et al.. Azuma et al.'s description states "ten cervical
vertebrae are preserved, missing at least the atlas" but the materials
list only says "eight cervical vertebrae" are present, and Hattori et
al. reveal nine cervicals are preserved "missing 4th and 7th" with the
atlantal neurapophyses having been initially identified as axial ribs.
Azuma et al. say "likely pleurocoels are present in all dorsal
vertebrae in the form of longitudinal fossae on the lateral surfaces of
centra", but only foramina are considered pleurocoels by most authors,
and Hattori et al. find that not even the anteriormost dorsals have
pleurocoels. Thus this character was incorrectly scored as
dromaeosaurid-like. Azuma et al. also state "the parapophyses of the
dorsal vertebrae including the posterior ones are stalk-like as in
derived alvarezsauroids and dromaeosaurids, though they are not as
prominent as in the latter groups", but they're actually short. Those
authors also say "the most unusual feature is that the prezygapophyses
of the middle caudal vertebrae are distally bifid (Fig. 6i), which has
not been reported in any dinosaurs", but this is a standard
dromaeosaurid character reported in e.g. Deinonychus and Velociraptor. Contra the text
and scoring, the coracoid is proximodistally shallow, unlike
pennaraptorans. Figure 7's caption is incorrect and the humeri shown
are a right in anterior view and left in lateral view, not the left "in
lateral (left) and posterior (right) view." Also, the femur in figure
7f is in medial and posterior views, not lateral and posterior. In
figure 7h, the pedal phalanges are placed incorrectly, with II-1 and
II-2 switched with IV-1 and IV-2. The measurement table is partly
inconsistent as it has IV-2 subequal to IV-3 and IV-4 in length, unlike
the figure. Hattori et al. agree with this based on their figures and
description, but their materials list incorrectly says supposed IV-2 is
actually III-2 and supposed III-1 is actually II-2, which besides just
being wrong anatomically and in their own figures would leave us with
an extra III-2 and no III-1. Contra the text and coding, I don't think
the second pedal digit looks particularly deinonychosaurian- Tanycolagreus has the same
dorsally prominent distal articular surface on II-1, and the ungual in Ornitholestes
is comparatively larger. Hattori et al. also list the folowing
"Re-identified elements: right maxilla, originally identified as right
premaxilla; ... partial neural arch of fifth dorsal vertebra,
originally identified as left squamosal; right manual phalanx III-1,
originally identified as right pedal phalanx I-1; right pubis,
originally identified as left pubis; left metatarsals II and V,
originally identified as right metatarsals IV and V,
respectively." They list the following "Additionally identified
elements: right quadrate; both parietals; right ectopterygoid; centrum
of 2nd dorsal vertebra; neural spine of 9th dorsal vertebra; partial
neural arch of 4th caudal vertebra; prezygapophyses of 18th caudal
vertebra; 29th caudal vertebra; several rib elements; distal end of
right humerus; left manual phalanx I-2; left femoral head; distal part
of right fibula; right distal tarsal III; left metatarsal III; right
metatarsal IV; left pedal phalanx I-2". Finally, they list three
"Withdrawn elements:
left pterygoid; posterior caudal vertebra; left ischium" without
comment, but Hattori (pers. comm. 3-4-2022) indicates the pterygoid and
ischium cannot be identified in the existing materials and that the
caudal is theropodan but not part of the holotypic block.
Azuma et al. (2016) add "Fukuivenator" to Turner et al.'s TWiG matrix and recover
it as a coelurosaur in a polytomy with compsognathids, Ornitholestes,
ornithomimosaurs and maniraptorans, contra their statements that they
found the taxon to be a basal maniraptoran. Experimentation shows the
polytomy exists regardless of "Fukuivenator"'s
presence in the tree. Azuma et al. find that constraining the genus to
be a paravian, deinonychosaur or dromaeosaurid only takes three more
steps. As indicated above, many of the supposedly dromaeosaurid-like
characters are incorrectlyscored. Cau (2018) recovered the taxon
as either the most basal
paravian or a basal dromaeosaurid. More recently, Hartman et al.
(2019) found "Fukuivenator"
to be the basalmost alvarezsauroid, though it moves to a basal
therizinosaurian position in only two steps. A more stemward
position seems more likely than a relationship with dromaeosaurids, as
it can be a coelurid with only 4 more steps, but takes 7 steps to be
sister to Pennaraptora and 11 steps to be paravian. Forcing it to
be a dromaeosaurid is 27 steps longer, so is extremely unlikely.
Hattori et al. (2021) in their redescription added it to a later TWiG
matrix and recovered it as the basalmost therizinosaurian, while adding
it to Hartman et al.'s analysis using the new data results in it being
sister to Ornitholestes with this pair sister to Maniraptoriformes.
References- Anonymous, 2009. [3rd new species? Small-sized meat diet
dinosaur to restoration Fukui] msn.com 3/18/2009
Shibata and Azuma, 2010. New dinosaurs from the Lower Cretaceous Kitadani Formation
of the Tetori Group, Fukui, Central Japan. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 163A-164A.
Azuma, Xu, Shibata, Kawabe, Miyata and Imai, 2016. A bizarre theropod from the
Early Cretaceous of Japan highlighting mosaic evolution among coelurosaurians.
Scientific Reports. 6, 20478.
Mortimer, 2016 online. https://theropoddatabase.blogspot.com/2016/02/fukuivenator-thoughts.html
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25.
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.
Hattori, Kawabe, Imai, Shibata, Miyata, Xu and Azuma, 2021. Osteology of Fukuivenator paradoxus:
A bizarre maniraptoran theropod from the Early Cretaceous of Fukui,
Japan. Memoir of the Fukui Prefectural Dinosaur Museum. 20, 1-82.
Maniraptoriformes Holtz, 1995
Definition- (Ornithomimus velox + Passer domesticus) (Maryanska, Osmolska and Wolsan, 2002; modified from Holtz, 1996)
Other definitions- (Ornithomimus velox + Dromaeosaurus albertensis + Passer domesticus) (modified from Holtz and Padian, 1995)
(Ornithomimus edmontonicus + Passer domesticus)
(Turner, Makovicky and Norell, 2012)
(Ornithomimus velox + Vultur gryphus) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
= Ornithomimoidea Marsh, 1890 sensu Zhao, 1983
Definition- (Ornithomimus velox + Shuvuuia deserti) (modified
from Sereno, 1999)
Other definitions- (Ornithomimus edmontonicus <- Tyrannosaurus rex, Nqwebasaurus thwazi, Shuvuuia
deserti, Therizinosaurus cheloniformis, Oviraptor philoceratops, Troodon formosus,
Passer domesticus) (Sereno, 2017)
(Deinocheirus mirificus + Ornithomimus velox) (Hendrickx,
Mateus, Ara�jo and Choiniere, 2019)
= Protoavia Paul, 1988
= "Eumaniraptora" Holtz, 1992
= "Pneumatocrania" Holtz, 1992
= Bullatosauria Holtz, 1994
Definition- (Ornithomimus velox + Troodon formosus) (modified
from Holtz, 1996)
= Maniraptoriformes sensu Holtz and Padian, 1995
Definition- (Ornithomimus velox + Dromaeosaurus albertensis + Passer domesticus) (modified)
= Ornithomimosauria sensu Padian, Hutchinson and Holtz, 1999
Definition- (Pelecanimimus polyodon + Ornithomimus edmontonicus)
(modified)
= Ornithomimosauria sensu Senter, 2011
Definition- (Pelecanimimus polyodon + Harpymimus okladnikovi +
Shenzhousaurus orientalis + Ornithomimus velox) (modified)
= Maniraptoriformes sensu Turner, Makovicky and Norell, 2012
Definition- (Ornithomimus edmontonicus + Passer domesticus)
= Maniraptoriformes sensu Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Ornithomimus velox + Vultur gryphus)
Comments-
Zhao (1983) named Ornithomimoidea as a new superfamily of toothless
Late Cretaceous coelurosaurs which excluded podokesaurids and
coelurids. It was later used by Sereno (1999) as an arctometatarsalian
clade containing ornithomimosaurs and alvarezsauroids, but not
therizinosaurs. Sereno (2017) redefined it to include
ornithomimosaurs closer to ornithomimids than Nqwebasaurus,
but as the latter is an alvarezsauroid here, his Ornithomimoidea is a
junior synonym of Ornithomimosauria here. Hendrickx et al. (2019)
redefined it to refer "to toothless ornithomimosaurs, which includes
the 'family'-level clades Ornithomimidae and Deinocheiridae", but the
Hartman et al matrix found Deinocheirus to be sister to toothed Hexing, further from toothless ornithomimids than toothed Harpymimus and Shenzhousaurus. Thus all ornithomimosaurs are covered under their definition of (Deinocheirus mirificus + Ornithomimus velox) and Deinocheirus is convergently toothless.
Paul (1988) used Protoavia for a clade conatining what are today recognized
as maniraptoriforms, which would also correspond to the modern definition of
Coelurosauria in his topology. The name has not seen much use since, and is
inadvisable due to the eponymous Protoavis being a chimaera of taxa less
closely related to birds.
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. He also named "Pneumatocrania" there, to contain his Arctometatarsalia
(caenagnathids, Avimimus, tyrannosaurids, troodontids and ornithomimosaurs)
plus oviraptorids, though the name was left out of the 1994 published version.
No subsequent analysis has recovered this group, which seems largely based on
miscodings.
In the late 1980s and 1990s, a sister group relationship between ornithomimosaurs
and troodontids was popular based on the bulbous cultriform process and dental
anatomy of Pelecanimimus. This was formalized by Holtz (1994) as the
clade Bullatosauria, defined by him in 1996. The discovery of basal troodontids
like Sinovenator showed they were ancestrally bird-like, and bullatosaurs
have not been supported by many studies since. While Bullatosauria predates
Maniraptoriformes and was defined in the same publication, its limited concept
has led to the widespread use of Maniraptoriformes for the now far more inclusive
Ornithomimus+Troodon clade.
Maniraptoriformes defined-
Holtz and Padian (1995) define Maniraptoriformes as "the node
connecting Arctometatarsalia with Maniraptora", but since their
Maniraptora was a node-based taxon of Dromaeosaurus plus birds, the modern node-stem triplet wasn't formed yet.
Unlike Turner et al. (2012), Maryanska et
al. (2002) used Ornithomimus velox, the type species, as dictated by
Phylocode. To illustrate why this is a good idea, consider the fact that Makovicky
et al. (2004) synonymized O. edmontonicus with Dromiceiomimus.
They listed the species as O. edmontonicus, but brevitertius has
priority, so the species should be Ornithomimus brevitertius. Ornithomimus
velox, on the other hand, remains valid. Makovicky et al. also considered
the possibility O. brevitertius (as O. edmontonicus) may be a
junior synonym of O. velox, and deCourten and Russell (1985) suggested
it (again as O. edmontonicus) may warrant generic separation from O.
velox if the specimen they describe is properly referred to the latter species.
Then Turner et al.'s redefinitions of taxa eponymous with Ornithomimus
would not be based on Ornithomimus. Sereno also used edmontonicus
online and claims O. edmontonicus is the taxon represented by most analyses,
not O. velox, but only the TWG matrix (from Ji et al., 2003 onward) and
Kobayashi's work (Kobayashi and Lu, 2003; Kobayashi, 2004; Kobayashi and Barsbold,
2005; Kobayashi and Barsbold, 2005) have used Ornithomimus as an OTU,
and the latter uses both species as references. So this is not a valid rationale.
References- Marsh, 1890. Description of new dinosaurian reptiles. The
American Journal of Science, series 3. 39, 81-86.
Zhao, 1983. Phylogeny and evolutionary stages of Dinosauria. Acta Palaeontologica
Polonica. 28(1-2), 295-306.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
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.
Holtz, 1995. A new phylogeny of the Theropoda. Journal of Vertebrate Paleontology. 15(3), 35A.
Holtz and Padian, 1995. Definition and diagnosis of Theropoda and related taxa. Journal of Vertebrate Paleontology. 15(3), 35A.
Holtz, 1996. Phylogenetic taxonomy of the Coelurosauria (Dinosauria: Theropoda).
Journal of Paleontology. 70, 536-538.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions
and nomenclature of the major taxonomic categories of the carnivorous Dinosauria
(Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Makovicky, Kobayashi and Currie, 2004. Ornithomimosauria. In Weishampel, Dodson
and Osmolska (eds.). The Dinosauria Second Edition. University of California
Press. 137-150.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological
continuity within Dinosauria. Journal of Evolutionary Biology. 24(10), 2197-2216.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Zanno and Makovicky, 2011. Body mass evolution in omnivorous/herbivorous coelurosaurian
dinosaurs. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 219.
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.
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.
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.
unnamed maniraptoriform (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-38424) tooth (?x.5x? mm)
Comments- This is slightly labiolingually compressed, strongly recurved,
lacks a basal constriction, and has weak carinae without serrations.
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the
Late Cretaceous of the San Juan Basin, northwestern New Mexico and their implications
for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
unnamed Maniraptoriformes (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Late Campanian, Late Cretaceous
La�o, Sedano Formation, Spain
Material- (MCNA 14523) tooth (2.8x3.5x1.6 mm)
(MCNA 14524) tooth (2.8x2.2x1.6 mm)
(MCNA 14525) tooth (1.5x1.2x.7 mm)
(MCNA 14526) tooth (3.9x4.2x1.4 mm)
(MCNA 14527) tooth (1.5x1.1x.8 mm)
(MCNA 14528) tooth (4.1x2.8x1.2 mm)
(MCNA 14529) tooth (4.8x2.6x1.3 mm)
(MCNA 14530) tooth (4.5x1.8x1.2 mm)
(MCNA 14531) tooth (3.1x2.1x1.1 mm)
(MCNA 14532) tooth (2.4x1.1x.4 mm)
(MCNA 14533) tooth (1.4x.8x.6 mm)
(MCNA 14534) tooth (1.7x1.6x.8 mm)
(MCNA 14535) tooth (1.5x.9x.6 mm)
(MCNA 14536) tooth (2.1x.6x.5 mm)
(MCNA 14537) tooth (1.6x1.4x.7 mm)
(MCNA 14538) tooth (2.8x1.2x.8 mm)
(MCNA 14539) tooth (2x1.3x.8 mm)
(MCNA 14540) tooth (1.8x1.2x.6 mm)
(MCNA 14541) tooth (2.4x1.8x.9 mm)
(MCNA 14542) tooth (1.7x1.3x.6 mm)
(MCNA 14543) tooth (1.4x1.2x.5 mm)
(MCNA 14544) tooth (3.1x1.8x1.1 mm)
(MCNA 14545) tooth (8.1x4.3x2.3 mm)
(MCNA 14546) tooth (5.8x1.8x1.5 mm)
(MCNA 14547) tooth (8.1x3.4x1.9 mm)
(MCNA 14548) tooth (2.6x2.1x1.7 mm)
(MCNA 14549) tooth (2.4x1.5x1 mm)
(MCNA 14550) tooth (5.3x2.5x1.9 mm)
(MCNA 14551) tooth (5.5x1.8x1.2 mm)
(MCNA 14552) tooth (4.3x1.9x1.5 mm)
(MCNA 14553) tooth (4x1.6x1.1 mm)
(MCNA 14554) tooth (2.9x1.9x1.4 mm)
(MCNA 14555) tooth (3.5x1.9x1.2 mm)
(MCNA 14556) tooth (3.8x1.5x1.2 mm)
(MCNA 14557) tooth (3.2x1.4x.9 mm)
(MCNA 14558) tooth (1.8x1.4x1.1 mm)
(MCNA 14559) tooth (2.4x1.2x.8 mm)
(MCNA 14560) tooth (2.6x1.8x1.2 mm)
(MCNA 14561) tooth (1.6xx1.1x.7 mm)
(MCNA 14564) tooth (2.3x.9x.6 mm)
(MCNA 14565) tooth (1.6x.8x.5 mm)
Comments-
These are recurved and unserrated but lack a constricted base.
Isasmendi et al. (2020) referred at least two of the teeth called
Coelurosauria indet. by Torices et al. (2015) to cf. Paronychodon (MCNA 14562 and 14563),
as they have longitudinal grooves. It is likely more La�o teeth listed here belong to Paronychodon
as well. Isasmendi et al. referred the rest of these teeth to
Paraves indet., whuch is possible although some could plausibly be e.g.
alvarezsaurid as well.
Reference- Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod
dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta
Palaeontologica Polonica. 60(3), 611-626.
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.
undescribed maniraptoriform (Company, Torices, Pereda-Suberbiola and
Ruiz-Omenaca, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Sierra Perenchiza Formation, Valencia, Spain
Material- teeth (~6 mm)
Comments- Described as distally recurved, strongly compressed labiolingually,
and lacking serrations.
Reference- Company, Torices, Pereda-Suberbiola and Ruiz-Omenaca, 2009.
Theropod teeth from the Late Cretaceous of Chera (Valencia, Eastern Spain).
Journal of Vertebrate Paleontology. 29(3), 81A.
unnamed Maniraptoriformes (Torices, 2002)
Late Campanian, Late Cretaceous
Vicari 4, Tremp Formation, Spain
Material- (DPM-VIR4-T5) tooth (2.9x1.3x.9 mm) (Torices, 2002)
Late Campanian-Early Maastrichtian, Late Cretaceous
Montrebei, Tremp Formation, Spain
(DPM-MON-T3) tooth (2.8x1.6x.7 mm) (Torices, 2002)
(DPM-MON-T6) tooth (3.3x1.4x1.2 mm) (Torices, 2002)
Late Maastrichtian, Late Cretaceous
Blasi 2B, Tremp Formation, Spain
(MPZ98/79) tooth (1.9x1.9x1 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
(MPZ98/80) tooth (2.6x1.8x.9 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
(MPZ98/81) tooth (1.9x1.4x.7 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
(MPZ98/82) tooth (2.4x1.2x.7 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Comments- These are recurved and unserrated but lack a constricted base.
References- Torices, 2002. Los dinosaurios ter�podos del Cret�cico
Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de
Paleontolog�a. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
undescribed possible maniraptoriform (Nessov, 1995)
Coniacian-Santonian, Late Cretaceous
Geisu River, Armenia
Material- (CCMGE coll.) partial radius or ulna or proximal tibiotarsus
Comments- Initially reported by
Nessov (1995) as "Geisu, river in Northern Armenia. Upper Cretaceous,
Coniacian - Santonian based on discoveries of ammonites (data on the
age and discovery by A. A. Atabekyan). Tubular limb bone of a theropod"
(translated). Averianov and Atabekyan (2005) resolve some ambiguity by
stating "A fragment of a tubular bone of a predatory dinosaur was found
in the Upper Cretaceous (Coniacian-Santonian) marine deposits of the
Geisu River in northern Armenia." Averianov (pers. comm., 5-2023) says
it is stored in the CCMGE, while an unpublished manuscript by Atabekyan
and Nessov reveals it was discovered in 1975 and is a gracile element
with a preserved length of ~175 mm, shaft diameter of 22x17 mm and bone
wall thickness of 2 mm.
References- Atabekyan and Nessov, MS (~1987-1989). Первне находки костей летающего ящера и динозавров в мезозое Армении [The first finds of bones of a flying lizard and dinosaurs in the Mesozoic of Armenia]. 14 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, St. Petersburg. 1-156.
Averianov and Atabekyan, 2005. The first discovery of a flying reptile
(Pterosauria) in Armenia. Paleontological Journal. 39(2), 210-212.
undescribed maniraptoriform (Nessov, 1995)
Late Santonian-Early Campanian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- astragalocalcaneum
Comments-
Nessov (1995) referred this to Troodontidae based on the fused tarsus,
but as pointed out by Averianov and Sues (2007) this is also known in
other maniraptoriforms (e.g. deinocheirids, derived alvarezsauroids,
caenagnathids, some oviraptorids, some unenlagiids, most
dromaeosaurids, most avialans).
References- Nessov, 1995. Dinosaurs of Northern Eurasia: New data about
assemblages, ecology and paleobiogeography. Scientific Research Institute of
the Earth's Crust, St. Petersburg State University.
156 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.
undescribed maniraptoriform (Watabe, Tsogtbaatar, Ichinnorov and Barsbold, 2004)
Cenomanian-Turonian, Late Cretaceous
Bayshin Tsav, Bayanshiree Formation, Mongolia
Material- (uncollected?) manual phalanx, manual ungual, fragments
Comments- Watabe et al. (2004)
photograph "an uncus and phalanges of a theropod discovered in Baynshin
Tsav", showing a strongly curved manual ungual with large proximally
placed flexor tuber, and a slender bowed phalanx perhaps a
therizinosaur, oviraptorosaur or deinonychosaur.
Reference- Watabe, Tsogtbaatar, Ichinnorov and Barsbold, 2004. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 2001. Hayashibara Museum
of Natural Sciences Research Bulletin. 2, 69-96.
Maniraptoriformes indet. (Janensch, 1925)
Late Kimmeridgian, Late Jurassic
Quarry Ig, Middle Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (HMN MB R 1762) manual phalanx I-1 (55 mm)
Comments- Janensch (1925) refers a manual phalanx (HMN M.B.R. 1762) from contemporaneous Quarry Ig to Elaphrosaurus bambergi, but Rauhut and Carrano state "as there is no
overlap with the type specimen, its referral to Elaphrosauruscannot
be tested, and therefore we do not consider it further here."
While Janensch tentatively identified the phalanx as II-2, Rauhut and
Carrano reidentified it as I-1 which seems to be correct based on its
asymmetry. Based on its elongation it is here placed in
Maniraptoriformes indet..
References-
Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten Deutsch-Ostafrikas.
Palaeontographica. (Supp. 7)1, 1-99.
Rauhut and Carrano, 2016. The theropod dinosaur Elaphrosaurus bambergi Janensch, 1920, from the Late Jurassic of Tendaguru, Tanzania. Zoological Journal of the Linnean Society. 178(3), 546-610.
unnamed Maniraptoriformes (Lasseron, 2020)
Early Bathonian, Middle Jurassic
GEA 2, Guelb el Ahmar,
Anoual Formation, Morocco
Material- (MNHN GEA2-57; Theropoda gen. et sp. indet. morphotype VI) lateral tooth (3.06x1.59x1.12 mm)
Early Bathonian, Middle Jurassic
GEA 7, Guelb el Ahmar,
Anoual Formation, Morocco
(MNHN GEA7-15; Theropoda gen. et sp. indet. morphotype VI) lateral tooth (2.40x1.21x.73 mm)
Comments- Discovered in 2015 and/or 2018, these are assigned to Maniraptoriformes here based on the lack of serrations.
Reference- Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.
unnamed Maniraptoriformes (Knoll and Ruiz-Omenaca, 2009)
Beriassian, Early Cretaceous
KM 1983, Ksar Metlili Formation, Morocco
Material- (MNHN SA 2004/2B; lost) anterior tooth (2.60x1.40x.88 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA 2004/4C; lost) tooth (3.68x2.28x1.08 mm) (Knoll and Ruiz-Omenaca, 2009)
(MNHN SA A0; lost) anterior tooth (1.68x.88x.56 mm) (Knoll and Ruiz-Omenaca, 2009)
Beriassian, Early Cretaceous
KM-D2, Ksar Metlili, Ksar Metlili Formation, Morocco
?(FSAC-KM-D2-8; Theropoda gen. et sp. indet. morphotype IV) lateral tooth (3.12x1.22x.78 mm), two teeth (Lasseron, 2020;
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
Comments- The MNHN specimens were collected in 1983, 1986 and/or 1999, while the
FSAC specimen was collected in 2010, 2015 or 2018 (Lasseron, 2020). Lasseron noted the "dinosaur remains (Knoll, 2000; Knoll & Ruiz-Ome�aca, 2009) ... were taken out of the MNHN and lost." The
MNHN teeth are referred to Maniraptoriformes by Knoll and Ruiz-Omenaca
(2005) as they lack serrations. Similarities were noted to
isolated teeth from Cretaceous Europe described as Coelurosauria indet..
FSAC-KM-D2-8 has a more elongate slender curve compared to most
theropod teeth, so may belong to another group (Pterosauria?).
Lasseron et al. list all three FSAC-KM-D2-8 teeth as Maniraptoriformes.l
References- Knoll and Ruiz-Omenaca, 2005. Theropod teeth from the Berriasian
of Anoual (Morocco). Journal of Vertebrate Paleontology. 25(3), 78A.
Knoll and Ruiz-Omenaca, 2009. Theropod teeth from the basalmost Cretaceous of
Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine.
146(4), 602-616.
Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020 (online 2019). New data on the microvertebrate fauna from
the Upper Jurassic or lowest Cretaceous of Ksar Metlili (Anoual
Syncline, eastern Morocco). Geological Magazine. 157, 367‑392.
Arctometatarsalia
Maniraptora Gauthier, 1986
Definition- (Passer domesticus <- Ornithomimus velox)
(Maryanska, Osmolska and Wolsan, 2002; modified from Padian, Hutchinson and Holtz, 1997; modified from Gauthier,
1984)
Other definitions- (bowed ulna, semilunate carpal, slender metacarpal
III) (Holtz, 1994)
(Dromaeosaurus albertensis + Passer domesticus) (modified from
Holtz and Padian, 1995)
(Oviraptor philoceratops + Passer domesticus) (modified from Sereno,
1998)
(Ornitholestes hermanni + Archaeopteryx lithographica) (modified
from Choiniere, Xu, Clark, Forster, Guo and Han, 2010)
(Passer domesticus <- Ornithomimus edmontonicus) (Turner, Makovicky and Norell, 2012)
(Vultur gryphus <- Ornithomimus velox) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
= "Maniraptora" Gauthier, 1984
= Therizinosauridae sensu Sereno, 1998
Definition- (Erlikosaurus andrewsi <- Ornithomimus velox) (modified)
= Alvarezsauridae sensu Sereno, 1999
Definition- (Shuvuuia deserti <- Ornithomimus velox) (modified)
= Enigmosauria Naish, Hutt and Martill, 2001
= Maniraptora sensu
Turner, Makovicky and Norell, 2012
Definition- (Passer domesticus <- Ornithomimus edmontonicus)
= Maniraptora sensu Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Vultur gryphus <- Ornithomimus velox)
Comments-
Maniraptora was first named by Gauthier (1984) in his unpublished
thesis, using a cladistic philosophy but no quantitative
analysis. Proposed as "birds and all coelurosaurs that are closer
to birds than they are to Ornithomimidae" (p297), it included Avialae,
Deinonychosauria (with Saurornitholestes and Hulsanpes as different OTUs), oviraptorosaurs (as Caenagnathidae, Elmisauridae and Microvenator), Compsognathus, Ornitholestes and Coelurus.
Gauthier and Padian (1985) later presented this concept in published
form, albeit without naming the deinonychosaur plus bird clade.
The full analysis with data matrix was published by Gauthier (1986),
although my reanalysis shows Compsognathus, Ornitholestes and Coelurus
only fell out as maniraptorans there due to misscorings (see Evaluating
Phylogenetic Analyses). This concept of grouping oviraptorosaurs,
deinonychosaurs and birds to the exclusion of ornithomimosaurs,
carnosaurs and coelophysoids was rather novel for the time. For
one thing, few explicit hypotheses of theropod relationships included
birds until the 1990s. On the other hand, Paul (1984, 1988) was
noncommital with regard to ornithomimosaur relationships, viewing them
as potentially closer to birds than Archaeopteryx
and dromaeosaurids. Thulborn (1984) would only count Avimimus and birds as maniraptoran, with troodontids being closer to ornithomimids, and Archaeopteryx,
oviraptorosaurs and dromaeosaurids outside what would now be
Maniraptoriformes. Bakker's (1986) dinosaur family tree in the
non-technical book "The Dinosaur Heresies" does show dromaeosaurs
closer to birds than ornithomimids or tyrannosaurs however.
In the 1990s it was common to place troodontids, oviraptorosaurs (at least caenagnathids and/or Avimimus),
therizinosaurs, alvarezsauroids and/or tyrannosauroids in the sister
clade to Maniraptora- Arctometatarsalia, those taxa closer to
ornithomimosaurs than to birds. The discovery of bird-like basal
oviraptorosaurs and troodontids around 2000, like Caudipteryx and Sinovenator,
led to these clades confidently being placed in Maniraptora, while the
discovery of more coelurid-like basal tyrannosauroids lessened that
superfamily's similarity to ornithomimosaurs. Alvarezsauroids and
therizinosaurs are still controversial members however, moving to be
arctometatarsalians with 4 steps in the matrix of Hartman et al.
(2019). While usually ignored due to the homogeniety of TWiG
results, these two clades can plausibly be arctometatarsalians in Cau's
(2018) matrix as well, with only 5 and 6 steps respectively.
While more stemward coelurosaurs like Compsognathus, Ornitholestes and Coelurus
are still recovered as maniraptorans in some analyses, Hartman et al.'s
results suggest these are unlikely to be correct, requiring 8, 13 and
13 more steps respectively.
Therizinosaurs plus oviraptorosaurs?-
For a time, a clade of therizinosaurs plus oviraptorosaurs seemed
likely, beginning with the results of Makovicky's (1995) axial analysis
and being common until Zanno's 2010 osteology of Falcarius
and revision of Therizinosauria. Naish et al. (2001) named
Enigmosauria in a cladogram for this concept. It was not defined or
mentioned in the text, as the
authors had only accidentally left in in the figure after they decided
not to formally name the clade in that publication. Other defined
names that could function for this clade include Caenagnathiformes and
Oviraptoriformes (see comments for Oviraptorosauria). Such a
pairing now seems
unlikely however, considering Hartman et al. (2019) requires 13 steps
to enforce
it.
Maniraptora defined- Holtz (1994) has been the only
author to attempt to explicitly change the content of Maniraptora to
include ornithimosaurs in his topology. Correctly predicting
ornithomimosaurs and tyrannosaurids evolved from ancestors with the
namesake 'snatching hands' of Maniraptora, Holtz changed the definition
to be "the first theropod possessing the derived fore limb structures
described by Gauthier (including a semilunate carpal structure, a thin
narrow metacarpal III, and bowed ulna) and its descendants." This
covered all theropods in his cladogram except Compsognathus.
As is usual for apomorphy-based definitions, this one has issues with
ambiguous character definition and homoplasy, compounded here by using
three independent features. For instance, in the Hartman et al.
(2019) matrix a semilunate carpal (defined by its deep semicircular
shape) is recovered as a pennaraptoran character convergent in
therizinosaurians and derived alvarezsauroids (due to its absence in Pelecanimimus, Nqwebasaurus and Protarchaeopteryx). A bowed ulna is ambiguous though, as the most basal therizinosaur and alvarezsauroid (Falcarius and Fukuivenator)
have one but other members of those clades don't. The degree of
slenderness in metacarpal III is never specified, making it more
difficult to comment on. In any case, Holtz saw this folly and
changed it to a node-based definition the next year.
That and the other two node-based definitions listed here all have the
same thing in common- they function to recover the same known content
as Gauthier's definition, but only in their respective
topologies. So for example, Sereno's (1998) definition of Oviraptor plus Passer
works in his topology where oviraptorosaurs are the first branching
maniraptorans, and alvarezsauroids and therizinosaurs are
arctometatarsalians. But in most topologies such as Hartman et
al.'s, this defines a maniraptoran subgroup subsequently named
Pennaraptora. Similarly, Holtz and Padian's (1995) definition
worked in their topology that had even oviraptorosaurs and troodontids
in Arctometatarsalia, but the dromaeosaurid plus bird clade was named
Eumaniraptora two years later by those same authors. Fianally,
Choiniere et al. (2010) had Ornitholestes as the basalmost maniraptoran, but their definition would encompass all Maniraptoromorpha in Hartman et al.'s tree.
See the comments under Maniraptoriformes for why Turner
et al.'s (2012) definition of Maniraptora using Ornithomimus edmontonicus
is inferior to Maryanska et al.'s (2002) using O. velox.
References-
Gauthier, 1984. A cladistic analysis of the higher systematic
categories of the Diapsida. PhD thesis. University of California. 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 and Padian, 1985. Phylogenetic, functional, and aerodynamic
analyses of the origin of birds and their flight. In Hecht, Ostrom,
Viohl and Wellnhofer (eds.). The Beginnings of Birds: Proceedings of
the International Archaeopteryx Conference, Eichst�tt 1984. Freunde des Jura-Museums Eichst�tt, Eichst�tt. 185-197.
Bakker, 1986. The Dinosaur Heresies. Kensington. 481 pp.
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.
464 pp.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Holtz and Padian, 1995. Definition and diagnosis of Theropoda and related taxa. Journal of Vertebrate Paleontology. 15(3), 35A.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). Masters thesis, Copenhagen University. [pp]
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.
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.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: Theropods. In
Martill and Naish (eds). Dinosaurs of the Isle of Wight. The
Palaeontological Association. 242-309.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Tsuihiji, 2004. The neck of non-avian maniraptorans: How bird-like was the cervical
musculature of the "bird-like" theropods? Journal of Vertebrate Paleontology.
24(3), 21A-22A.
Codd and Manning, 2007. Uncinate processes: A unique synapomorphy for maniraptoran
and avian theropods? Journal of Vertebrate Paleontology. 27(3), 60A.
Dececchi and Larsson, 2008. Critical analysis of arboreality in maniraptoran
theropods. Journal of Vertebrate Paleontology. 28(3), 70A.
Dececchi, Harrison and Larsson, 2009. Up in arms: An analysis of evolutionary
trends within the maniraptoran appendicular skeleton using allometric and Baysian
phylogenetic approaches. Journal of Vertebrate Paleontology. 29(3), 86A.
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal alvarezsauroid theropod
from the Early Late Jurassic of Xinjiang, China. Science. 327, 571-574.
Zanno, 2010. A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria:
Maniraptora). Journal of Systematic Palaeontology. 8(4), 503-543.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Balanoff, 2014. Archaeopteryx and the evolution of the paravian brain.
Journal of Vertebrate Paleontology. Program and Abstracts 2014, 84.
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.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Bradycnemidae Harrison and Walker, 1975
Bradycneme Harrison and Walker,
1975
B. draculae Harrison and Walker, 1975
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Holotype- (NHMUK A1588) distal tibiotarsus (37.8 mm wide)
Comments- The holotype was originally referred to Elopteryx (Lambrecht,
1929, 1933), then considered a pelecaniform. Harrison and Walker (1975) later
separated the material and named Bradycneme as a new taxon of strigiform.
Later authors agreed Bradycneme was a non-avian theropod, beginning with
Brodkorb (1978). Glut (1982) notes Brett-Surman proposed it was non-avian
at the 1978 Society of Vertebrate Paleontology meeting, "an opinion
supported by Dr. Storrs Olson and Dr. R. T. Bakker." Martin (1983) suggested it was ornithomimid. Paul (1988) and
Osmolska and Barsbold (1990) thought it was troodontid. Le Loeuff et al. (1992)
suggested it was synonymous with Elopteryx, which they placed in the
Dromaeosauridae. Csiki and Grigorescu (1998) made Heptasteornis a junior
synonym and proposed it was a non-maniraptoran tetanurine. Naish and Dyke (2004)
noted the craniocaudally compressed rectangular shape in distal view was similar
to maniraptorans, while the astragalar ascending process lacks the alvarezsaurid
notched medial margin seen in Heptasteornis. They thus assigned Bradycneme
to Maniraptora indet.. Most recently, Hartman et al. (2019) scored Bradycneme in their phylogenetic analysis and found it emerges as a maniraptoran excluded from Ceratonykus+Mononykus, Therizinosauria, Oviraptorosauria and Deinonychosauria. This makes an alvarezsauroid or avialan identity most likely
References- Lambrecht, 1929. Mesozoische und tertiare Vogelreste aus
Siebenburgen. In Csiki (ed.). Xe Congres International de Zoologie. 1262-1275.
Lambrecht, 1933. Handbuch der Palaeornithologie. Gebr�der Borntraeger.
1024 pp.
Harrison and Walker, 1975. The Bradycnemidae, a new family of owls from the
Upper Cretaceous of Romania. Palaeontology. 18(3), 563-570.
Brodkorb, 1978. Catalogue of fossil birds. Part 5, Passeriformes. Bulletin of
the Florida State Museum, Biological Sciences. 23, 139-228.
Glut, 1982. The New Dinosaur Dictionary. Citadel Press. 288 pp.
Martin, 1983. The origin and early radiation of birds. In Brush and Clark, (eds.).
Perspectives in Ornithology. 291-338.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Osmolska and Barsbold, 1990. Troodontidae. In Weishampel, Dodson and
Osm�lska (eds.). The Dinosauria. University of California Press. 259-268.
Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992. The first record of dromaeosaurid
dinosaur (Saurichia, Theropoda) in the Maastrichtian of southern Europe: Palaeobiogeographical
implications. Bulletin de la Societe Geologique de France. 163(3), 337-343.
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.
Naish and Dyke, 2004. Heptasteornis was no ornithomimid, troodontid,
dromaeosaurid or owl: the first alvarezsaurid (Dinosauria: Theropoda) from Europe.
Neus Jahrbuch f�r Geologie und Pal�ontologie. 7, 385-401.
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 maniraptoran (Naish, 1998)
Barremian, Early Cretaceous
Wessex Formation, England
Material-
(MIGW 6214) (juvenile) femur (123 mm)
References- Naish, 1998. A small Wealden theropod, represented only by a femur.
Progressive Palaeontology 1998, Programme and Abstracts. 16.
Naish, 2000. A small, unusual theropod (Dinosauria) femur from the Wealden Group
(Lower Cretaceous) of the Isle of Wight, England. Neues Jahrbuch f�r Geologie
und Pal�ontologie Monatshefte. 2000, 217-234.
unnamed maniraptoran (Sanchez-Hernandez and Benton, 2014)
Early Barremian, Early Cretaceous
Camarillas Formation, Aragon, Spain
Material- (MPG-KPC24; holotype of Camarillasaurus cirugedae in part) incomplete anterior dorsal centrum (~30 mm)
Comments- Sanchez-Hernandez and Benton (2014) said this "may belong to the Camarillasaurus
skeleton, having been collected in close proximity to the other
elements", interpreting it as the posterior half of an anterior
cervical centrum. Wang et al. (2016) noted it "most likely
represents an anterior dorsal based on
the position of the parapophyses and presence of a hypapophysis" and
that "the morphology of the hypapophysis in Camarillasaurus more
strongly resembles those found in some coelurosaurs (e.g. Nomingia,
Avimimus, Mononykus) than the structure on dorsal 1 of Elaphrosaurus."
Samathi (2019) found "It is too small to belong to the same individual or even the same taxon as
Camarillasaurus", and Samathi
et al. (2021) excluded the element from
the type material. Contra both Sanchez-Hernandez and Benton and
Samathi et al., the preserved articular surface is the anterior one, as
it is adjacent to the parapophysis. Also, the labeled
parapophyses in figure 3B are the neural arch peduncles and the supposed
pneumatic foramen is merely a fossa. The combination of large
hypapophysis, concave to flat anterior articular surface and no dorsal
pleurocoels suggests a basal alvarezsauroid, Mahakala relative, troodontid or pygostylian, so it is here assigned to Maniraptora.
References- Sanchez-Hernandez and Benton, 2014 (online 2012). Filling the ceratosaur
gap: A new ceratosaurian theropod from the Early Cretaceous of Spain. Acta Palaeontologica
Polonica. 59(3), 581-600.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2016. Extreme ontogenetic
changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
Samathi, 2019. Theropod dinosaurs from Thailand and southeast Asia
Phylogeny, evolution, and paleobiogeography. PhD thesis, Rheinischen
Friedrich-Wilhelms-Universit�t Bonn. 249 pp.
Samathi, Sander and Chanthasit, 2021 online. A spinosaurid from
Thailand (Sao Khua Formation, Early Cretaceous) and a reassessment of
Camarillasaurus cirugedae from the Early Cretaceous of Spain.
Historical Biology. Latest Articles. DOI: 10.1080/08912963.2021.1874372
Metornithes Perle, Norell, Chiappe and Clark,
1993
Definition- (Mononykus olecranus + Passer domesticus) (modified
from Chiappe, 1995)
Comments- Metornithes was named by Perle et al. (1993) for a clade containing
Mononykus and Ornithothoraces, but not Archaeopteryx and non-bird
theropods. Chiappe (1995) was the first author to define the clade, making it
a node containing Mononykus and Neornithes. Under the current topology
it's a maniraptoran clade that may contain therizinosaurs, though if alvarezsauroids
are arctometatarsalians (4 more steps in Hartman et al., 2019 analysis) it will be a senior synonym of Maniraptoriformes.
Paul (2016) used the informal name therizinosauriforms for
"jeholornids, therizinosaurians, and avians and their common ancestor,
operative only if three groups form a clade that excludes all other
dinosaurs except oviraptorosaurs." This would correspond to a
clade 'Therizinosauriformes', defined as "Jeholornis prima, + Therizinosaurus cheloniformis + Passer domesticus, - Oviraptor philoceratops."
Yet such a clade would be highly unparsimonious, and indeed Paul's
listed characters are either meaningless ("Head somewhat elongated",
"Tail from very long to very short", "[semi]lunate carpal from well to
poorly developed", "three to four load-bearing toes"), absent in
therizinosaurs ("teeth ... not serrated", "Gastroliths often
present"), not demonstrated in jeholornithids ("teeth ... blunt,
leaf shaped"), or vague and/or also present in e.g. Archaeopteryx and/or Sapeornis ("blunt upper beak, extra joint in lower jaw absent, teeth small", "Arm long", "Foot not narrow").
References- Perle, Norell, Chiappe and Clark, 1993. Flightless bird from
the Cretaceous of Mongolia. Nature. 362, 623-626.
Chiappe, 1995. The first 85 million years of avian evolution. Nature. 378, 349-355.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press.
360 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
Unnamed clade
Definition- (Therizinosaurus cheloniformis, Alvarezsaurus calvoi <- Ornithomimus velox, Oviraptor philoceratops, Deinonychus antirrhopus, Passer domesticus)
= Alvarezsauria Bonaparte, 1991
Definition- (Alvarezsaurus calvoi <- Passer domesticus) (Agnolin,
Powell, Novas and Kundrat, 2012)
= Therizinosauria sensu Russell, 1997
Definition- (Alxasaurus elesitaiensis, Enigmosaurus mongoliensis,
Erlikosaurus andrewsi, Nanshiungosaurus brevispinus, Segnosaurus
galbinensis, Therizinosaurus cheloniformis <- Ornithomimus
velox, Troodon formosus, Oviraptor philoceratops) (modified)
= Therizinosauridae sensu Sereno, 1999
Definition- (Erlikosaurus andrewsi <- Ornithomimus velox, Oviraptor
philoceratops, Passer domesticus) (modified)
= Therizinosauroidea sensu Zhang, Xu, Sereno, Kwang and Tan, 2001
Definition- (Therizinosaurus cheloniformis <- Ornithomimus velox,
Oviraptor philoceratops, Velociraptor mongoliensis, Passer
domesticus)
= Alvarezsauroidea sensu Hu, Hou, Zhang and Xu, 2009
Definition- (Alvarezsaurus calvoi <- Ornithomimus edmontonicus,
Passer domesticus) (modified)
= Therizinosauroidea sensu Hu, Hou, Zhang and Xu, 2009
Definition- (Therizinosaurus cheloniformis <- Oviraptor philoceratops,
Passer domesticus) (modified)
= Alvarezsauroidea sensu Choiniere, Xu, Clark, Forster, Guo and Han, 2010
Definition- (Alvarezsaurus calvoi <- Passer domesticus)
Comments- A clade of
therizinosaurs and alvarezsauroids exclusive of ornithomimosaurs or
birds was recovered by Hartman et al. (2019), although alvarezsauroids
could be aractometatarsalians in 4 more steps and therizinosaurs could
be closer to Pennaraptora in 3 steps, so this group requires more
testing. Definitions listed above for alvarezsauroid or
therizinosaur clades were made without considering they could be sister
taxa, so do not exclude therizinosaurs or alvarezsauroids
respectively.
References- Bonaparte, 1991. Los vertebrados f�siles de la Formaci�n
Rio Colorado, de la Ciudad de Neuqu�n y Cercan�as, Cret�cico
Superior, Argentina. Revista del Museo Argentino de Ciencias Naturales "Bernardino
Rivadavia" e Instituto Nacional de Investigaci�n de las Ciencias
Naturales: Paleontolog�. 4(3), 15-123.
Russell, 1997. Therizinosauria. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. 729-730.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Zhang, Xu, Sereno, Kwang and Tan, 2001. A long-necked therizinosauroid
dinosaur from the Upper Cretaceous Iren Dabasu Formation of Nei Mongol, People’s
Republic of China. Vertebrata PalAsiatica. 39(4), 282-290.
Hu, Hou, Zhang and Xu, 2009. A pre-Archaeopteryx troodontid theropod
from China with long feathers on the metatarsus. Nature. 461, 640-643.
Choiniere, Xu, Clark, Forster, Guo and Han, 2010. A basal alvarezsauroid theropod
from the early Late Jurassic of Xinjiang, China. Science. 327, 571-574.
Agnolin, Powell, Novas and Kundrat, 2012 (online 2011). New alvarezsaurid (Dinosauria, Theropoda)
from uppermost Cretaceous of north-western Patagonia with associated eggs. Cretaceous
Research. 35, 33-56.
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
Nqwebasauridae Alifanov and Saveliev, 2015a
Comments- Alifanov and Saveliev (2015a,b) erected this family for Nqwebasaurus
and their supposed ornithomimosaur Lepidocheirosaurus, which is based
on material of the ornithischian Kulindadromeus. Thus the family is monotypic
and rejected, though it may be useful in the future.
References- Alifanov and Saveliev, 2015a. [The most ancient ornithomimosaur (Theropoda,
Dinosauria), with cover imprints from the Upper Jurassic of Russia]. Paleontologicheskii Zhurnal. 2015(6), 71-85.
Alifanov and Saveliev, 2015b. The most ancient ornithomimosaur (Theropoda,
Dinosauria), with cover imprints from the Upper Jurassic of Russia. Paleontological
Journal. 49(6), 636-650.
Nqwebasaurus de Klerk, Forster,
Sampson, Chinsamy and Ross, 2000
N. thwazi de Klerk, Forster, Sampson, Chinsamy and Ross, 2000
Berriasian-Valanginian, Early Cretaceous
Upper Kirkwood Formation, South Africa
Holotype-
(AM 6040) (subadult) maxilla (25 mm), lacrimal, jugal, postorbital,
squamosal, prefrontal,
frontals (30 mm), fused parietals, quadrate, incomplete
parabasisphenoid, prootic, otoccipital, stapes, palatine, pterygoid,
five sclerotic plates, surangular, angular, prearticular, articular,
fifth cervical vertebra
(~14 mm), sixth cervical vertebra (14.5 mm), seventh cervical vertebra
(~15 mm), eighth cervical vertebra (~16 mm), ninth cervical vertebra
(14.7 mm), tenth cervical vertebra (11.3 mm), first dorsal vertebra
(9.6 mm), partial second dorsal centrum, anterior dorsal centrum (10
mm), mid dorsal centrum (15.5 mm), two mid dorsal neural arches, dorsal
rib fragments, several gastralia, proximal chevron (15.9 mm), scapulae
(64.3 mm), coracoids (28 mm), humeri (~59.1, ~58.5 mm), radii (43.2
mm), ulnae (44 mm), scapholunares, semilunate carpals, metacarpals I
(~15.9,
15.5 mm), phalanx I-1 (23 mm), manual ungual I (22.2 mm), metacarpals
II (25.7, 24 mm), phalanx II-1 (11.8 mm), phalanx II-2 (13.8 mm),
manual ungual II (17.6 mm), metacarpals III (20, 19 mm), phalanges
III-1 (8.5 mm), phalanges III-2 (6.5 mm), phalanx III-3 (8 mm), manual
ungual III (18.5 mm), incomplete pubes, partial femora (~118 mm),
tibiae (140.2 mm), fibulae, astragali (15.5 mm trans), distal tarsal
III, metatarsal I (12.8 mm), phalanx I-1 (14.8 mm), pedal ungual I (9.7
mm), metatarsal II (65.3 mm), phalanges II-1 (20 mm), phalanges II-2
(12 mm), pedal ungual II (16.4 mm), metatarsal III (72.8 mm), phalanx
III-1 (22 mm), phalanx III-2 (19.1 mm), phalanx III-3 (12.2 mm), pedal
ungual III (17.4 mm), metatarsal IV (66 mm), phalanx IV-1 (10.9 mm),
phalanges IV-2 (6.1 mm), phalanx IV-3 (4.4 mm), partial phalanx IV-4,
proximal pedal ungual IV (~13 mm), two incomplete metatarsals, twelve
gastroliths
Referred- (AM coll.) femur, tibia, fibula (Sereno, 2017)
Diagnosis- (after de Klerk et al., 2000) ginglymus of metacarpal I very
robust and asymmetrical, with hypertrophied articular surfaces and greatly enlarged
lateral condyle;
metatarsal IV reduced in width to approximately half that of metatarsal III.
(after Choiniere et al., 2012b) ridge on the
lateral margin of the dorsal surface of the distal end of metacarpal I extending
proximally from lateral condyle.
(after Sereno, 2017) well defined, elongate beveled edge on orbital rim
anterior to postorbital; elongate dorsal centra (length approximately 3
times centrum diameter); long manual unguals II and III (the
latter more than twice the length of III-3).
Other diagnoses- Sereno (2017)
stated neither 'manual ungual I elongate (length is four times proximal
depth) and mediolaterally compressed' nor 'fibular shaft reduced
distally to thin splint', listed in de Klerk et al.'s (2000) diagnosis
were autapomorphic.
Choiniere et al.'s (2012b) proposed diagnostic dental characters (unserrated maxillary teeth set into a groove;
straight maxillary tooth crowns; maxillary tooth crowns conical) are typical of alvarezsauroids.
Comments-
The holotype was discovered in 1996. The maxilla was originally
misidentified as a palatine by de Klerk et
al. (2000) before Choiniere et al. (2012b) more fully described the
skull. Radermacher et al. (2021) used microCT scanning to reveal
new elements "including the jugal, lacrimal, postorbital, squamosal,
quadrate, pterygoid, palatine, prootic, parabasisphenoid, surangular,
angular, prearticular, articular; as well as finer details of the
already-known dentition, maxilla, prefrontal, frontal, and
parietal." Contra de Klerk et al., no caudal vertebral material
is
preserved (Sereno, 2017).
This taxon was described as a basal coelurosaur, and found to occupy such a
position in the analyses of Holtz et al. (2004) and Rauhut and Xu (2005). The
latter analyses found the taxon more derived than compsognathids, but outside
Tyrannoraptora and Maniraptora. Holtz et al. furthermore found it to clade with
Ornitholestes,
while Gishlick and Gauthier (2007) noted manual resemblences to
compsognathids and Novas et al. (2012) found it to fall out in that
family. Sereno (2001) noted manual characters shared with
ornithomimosaurs and alvarezsaurids. Dal Sasso and Maganuco
(2011) and Lee et al. (2014) recovered it as sister to alvarezsaurids.
Choiniere et al. (2012b) included new cranial data and found Nqwebasaurus
to be a basal ornithomimosaur, as in Brusatte et al. (2014), though it
was an alvarezsauroid in trees only 4 steps longer. Hartman et
al. (2019) recovered it as an alvarezsauroid intermediate between "Fukuivenator" and Pelecanimimus,
requiring 6 steps to move to Ornithomimosauria. A compsognathid
position requires 10 steps, and a position sister to Pennaraptora 7
steps. Radermacher et al. (2021) incorporated new cranial data into Choiniere's TWiG analysis and still recovered Nqwebasaurus
as a basal ornithomimosaur, while adding some of this data to Hartman
et al.'s analysis results in it being the basalmost member of the
alvarezsaur-therizinosaur clade.
References- de Klerk, Forster, Ross, Sampson and Chinsamy, 1997. New
maniraptoran and iguanodontian dinosaurs from the Early Cretaceous Kirkwood
Formation, South Africa. Journal of Vertebrate Paleontology. 17(3), 42A.
de Klerk, Forster, Ross, Sampson and Chinsamy, 1998. A review of recent dinosaur
and other vertebrate discoveries in the Early Cretaceous Kirkwood Formation
in the Algoa Basin, Eastern Cape, South Africa. Gondwana 10: Event Stratigraphy
of Gondwana. Journal of African Earth Sciences. 27(1A), 55.
de Klerk, Forster, Sampson, Chinsamy and Ross, 2000. A new coelurosaurian dinosaur
from the Early Cretaceous of South Africa. Journal of Vertebrate Paleontology.
20(2), 324-332.
Sereno, 2001. Alvarezsaurids: Birds or ornithomimosaurs? In Gauthier and Gall
(eds.). New Perspectives on the Origin and Early Evolution of Birds. Yale University
Press. 70-98.
Starck and Chinsamy, 2002. Bone microstructure and developmental
plasticity in birds and other dinosaurs. Journal of Morphology. 254,
232-246.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Rauhut and Xu, 2005. The small theropod dinosaurs Tugulusaurus and Phaedrolosaurus
from the Early Cretaceous of Xinjiang, China. Journal of Vertebrate Paleontology.
25(1), 107-118.
Gishlick and Gauthier, 2007. On the manual morphology of Compsognathus longipes
and its bearing on the diagnosis of Compsognathidae. Zoological Journal of the
Linnean Society. 149, 569-581.
Choiniere, 2010. Anatomy and systematics of coelurosaurian theropods from the
Late Jurassic of Xinjiang, China, with comments on forelimb evolution in Theropoda.
PhD thesis. George Washington University. 994 pp.
Dal Sasso and Maganuco, 2011. Scipionyx samniticus (Theropoda: Compsognathidae)
from the Lower Cretaceous of Italy: Osteology, ontogenetic assessment, phylogeny,
soft tissue anatomy, taphonomy, and palaeobiology. Memorie della Societ�
Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano.
281 pp.
Choiniere, Forster and de Klerk, 2012a. New information on Nqwebasaurus thwazi,
a coelurosaurian theropod from the Early Cretaceous (Hauteriverian?) Kirkwood
Formation in South Africa. Journal of Vertebrate Paleontology. Program and Abstracts
2012, 78.
Choiniere, Forster and de Klerk, 2012b. New information on Nqwebasaurus thwazi,
a coelurosaurian theropod from the Early Cretaceous Kirkwood Formation in South
Africa. Journal of African Earth Sciences. 71-72, 1-17.
Novas, Ezcurra, Agnolin, Pol and Ortiz, 2012. New Patagonian Cretaceous theropod
sheds light about the early radiation of Coelurosauria. Revista del Museo Argentino
de Ciencias Naturales. 14(1), 57-81.
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.
Watanabe, Gold, Brusatte, Benson, Choiniere, Davidson and Norell, 2015. Vertebral
pneumaticity in the ornithomimosaur Archaeornithomimus (Dinosauria: Theropoda)
revealed by computed tomography imaging and reappraisal of axial pneumaticity
in Ornithomimosauria. PLoS ONE. 10(12), e0145168.
Sereno, 2017. Early Cretaceous ornithomimosaurs (Dinosauria: Coelurosauria) from Africa. Ameghiniana. 54, 576-616.
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
Radermacher, Fernandez, de Klerk, Chapelle and Choiniere, 2021.
Synchrotron μCT scanning reveals novel cranial anatomy of the enigmatic
Early Cretaceous South African coelurosaur, Nqwebasaurus thwazi. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 213-214.
Therizinosauria
Pennaraptora Foth, Tischlinger and Rauhut,
2014
Definition- (Oviraptor philoceratops + Deinonychus antirrhopus
+ Passer domesticus) (Foth, Tischlinger and Rauhut,
2014)
= Chuniaoae Ji, Currie, Norell and Ji, 1998
Definition- (Caudipteryx zoui + Passer domesticus) (modified
from Martyniuk, 2012)
= Maniraptora sensu Sereno, 1998
Definition- (Oviraptor philoceratops + Passer domesticus)
= Dromavialae Ji and Ji, 2001
= Aviremigia Gauthier and de Quieroz, 2001 vide Martyniuk, 2012
Definition- (Remiges and rectrices [enlarged, stiff-shafted, closed-vaned with
barbules bearing hooked distal pennulae], pennaceous feathers arising from the
distal forelimbs and tail as in Passer domesticus)
Comments- Ji et al. (1998) found a topology where Caudipteryx
was more closely related to Archaeopteryx, alvarezsaurids and ornithothoracines
than Velociraptor and Protarchaeopteryx were. In their online
supplementary information, they call a section "Diagnoses of the Chuniaoae
and the Avialae under alternative optimizations," but go on to list characters
for Avialae and an "Unnamed clade of Caudipteryx + Avialae."
Thus it seems the authors originally intended to name their new clade Chuniaoae,
then decided to leave it unnamed, but didn't catch all the times they used the
name. The concept is invalid, as Caudipteryx is now recognized as an
oviraptorosaur, and two of the proposed chuniaoaen characters (posteriorly extensive
external nares; unserrated teeth) are maniraptoran symplesiomorphies that were
reversed in derived dromaeosaurids and present in Protarchaeopteryx,
while the other one (posteriorly extensive dorsal premaxillary process) is convergent
between some oviraptorosaurs and some birds. A Caudipteryx+Avialae clade
would now include all oviraptorosaurs and paravians, being a subset of Maniraptora
potentially excluding taxa which are placed as basal maniraptorans in some studies
(e.g. therizinosaurs, alvarezsauriods, Ornitholestes). Note Ji and Ji
(2001) later proposed a similar name (Chuniaoia) on a cladogram for a group
containing Protarchaeopteryx, but not birds. Martyniuk (2012) later defined
the clade as Caudipteryx plus Passer, but it has seen almost no
use outside of that volume.
Ji and Ji (2001) erected the taxon Dromavialae in a cladogram for a maniraptoran
group including Protarchaeopteryx, Archaeopteryx and pygostylians,
but not Oviraptor, Troodon or dromaeosaurids. The text suggests
Caudipteryx would be included as well. Dromavialae is invalid content-wise,
since Protarchaeopteryx and Caudipteryx are now recognized as
oviraptorosaurs, which are agreed by most authors to be further from birds than
dromaeosaurids are. The diagnostic feature of the clade is listed as "real
wings with symmetrical feathers of modern concept." This is now known to
be true in oviraptorosaurs, troodontids and dromaeosaurids as well, meaning
Dromavialae could be viewed as a junior synonym of Maniraptora.
Gauthier and de Quieroz (2001) stated that if phylogeny expressed the developmental
origin of feathers, additional clades could be named including Aviremigia, which
they defined using feather apomorphies. Martyniuk (2012) followed them, and
is credited here as his use was definite as opposed to conditional. While ornithomimosaurs
are now known to lack feathers with barbules or retrices (based on Dromiceiomimus),
the condition in therizinosaurs and alvarezsauroids is still uncertain, giving
Aviremigia an uncertain position in the present topology.
Foth et al. (2014) erected Pennaraptora for this clade, which is followed here
due to the lack of use for Chuniaoae, Dromavialae or Aviremigia, the uncertain applicability
of Aviremigia, and the fact Chuniaoae originally had a very different concept.
Paul (2016) creates the term aveairfoilan for taxa "with feather wings
or ancestors with same that are in the clade that includes extant
birds", in his topology including therizinosaurs and what are here
pennaraptorans except for scansoriopterygids. His concept seems
to be that scasnsoriopterygid membrane wings evolved parallel to
feather wings, and members with the latter character are aveairfoilan,
though Paul never uses the technical term 'Aveairfoila' that is
implied. No other publication has used the term either, so that
it remains informal, and difficult to apply given the uncertain
relationships and forelimb feathering in therizinosaurs,
alvarezsauroids and ornithomimosaurs.
References- Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs
from northeastern China. Nature. 393, 753-761.
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.
Gauthier and de Quieroz, 2001. Feathered dinosaurs, flying dinosaurs, crown
dinosaurs, and the name "Aves." In Gauthier and Gall (eds.). New Perspectives
on the Origin and Early Evolution of Birds: Proceedings of the International
Symposium in Honor of John H. Ostrom. Peabody Museum of Natural History. 7-41.
Ji and Ji, 2001. How can we define a feathered dinosaur as a bird? In
Gauthier and Gall (eds.). New Perspectives on the Origin and Early
Evolution of Birds: Proceedings of the International Symposium in Honor
of John H. Ostrom. 43-46.
Xu, Sullivan, Zhang and O'Connor, 2011. A new eumaniraptoran phylogeny and its
implications for avialan origins. Journal of Vertebrate Paleontology. Program
and Abstracts 2011, 217.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Pan Aves. 189 pp.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press.
360 pp.
unnamed pennaraptoran (Gilmore, 1924)
Middle Campanian-Early Maastrichtian, Late Cretaceous
Belly River Group, Alberta, Canada
Material- (CMN 8505) dorsal centrum
Comments- This was described by Gilmore (1924) as distinct from other
coelurosaurs known at the time, though possibly referrable to Chirostenotes
or Dromaeosauridae (neither of which were known from vertebrae at the time).
Currie et al. (1994) commented on a set of vertebrae thought by Gilmore to be
referrable to the same taxon, and noted that the dorsal centrum may be a caenagnathid
but cannot be distinguished from Saurornitholestes either.
Reference- Gilmore, 1924. A new coelurid dinosaur from the Belly River
Cretaceous Alberta. Canada Geological Survey, Bulletin n. 38, geological series
43, 1-13.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
undescribed pennaraptoran (Farke, Letteau Stallings and Andrews, 2020)
Late Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- (RAM 31314) partial pedal phalanx
Comments- Identified as
(?)Avialae by Farke et al., this also resembles pedal phalanges of
caenagnathids and troodontids in general side outline and collateral
pit placement, so is more generally assigned to Pennaraptora here
pending further study.
Reference- Farke, Letteau
Stallings and Andrews, 2020. New vertebrate localities and
biostratigraphic interpretations of the Mesaverde Formation (Campanian,
Late Cretaceous) in northwestern Wyoming. The Society of Vertebrate
Paleontology 80th
Annual Meeting, Conference Program. 136-137.
undescribed pennaraptoran (Larson and Rigby, 2005)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, US
Material- (BHI-5159) incomplete furcula
References- Larson and Rigby, 2005. Furcula of Tyrannosaurus rex.
In Carpenter (ed.). The Carnivorous Dinosaurs. 247-255.
DePalma, Burnham, Martin, Larson and Bakker, 2015. The first giant raptor (Theropoda:
Dromaeosauridae) from the Hell Creek Formation. Paleontological Contributions.
14, 16 pp.
Pennaraptora indet. (Schwimmer, Sanders, Erickson and Weems, 2015)
Late Campanian, Late Cretaceous
Donoho Creek Formation, South Carolina, US
Material- (ChM PV4818) dorsal centrum
Comments- This is approximately square, with little ventral concavity
and no pleurocoel. Based on this, it may be troodontid or related to Avimimus.
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 pennaraptoran (Rodriguez de la Rosa and Cevallos-Ferriz, 1998)
Late Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Material- (IGM-7713) distal phalanx
Comments- This resembles the penultimate manual phalanges of caenagnathids,
troodontids and dromaeosaurids in the dorsal expansion of the distal articulation.
However, it differs in having centrally placed ligament pits, in which it resembles
proximal manual and pedal phalanges. Some manual phalanges (e.g. Hagryphus)
and pedal phalanges (e.g. Sinornithoides) have both characters. It was
assigned to probable Troodontidae by Rodriguez de la Rosa and Cevallos-Ferriz
(1998), but resembles other maniraptorans just as closely.
Reference- 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.
unnamed pennaraptoran (Naish and Sweetman, 2011)
Valanginian, Early Cretaceous
Wadhurst Clay of the Hastings Group, England
Material- (BEXHM: 2008.14.1) posterior cervical vertebra (7.1 mm)
References- Austen, Brockhurst and Honeysett, 2010. Vertebrate fauna
from Ashdown Brickworks, Bexhill, east Sussex. Wealden News. 8, 13-23.
Naish and Sweetman, 2011. A tiny maniraptoran dinosaur in the Lower Cretaceous
Hastings Group: Evidence from a new vertebrate-bearing locality in south-east
England. Cretaceous Research. 32(4), 464-471.
unnamed pennaraptoran (Le Loeuff, Buffetaut, Mechin and Mechin-Salessy,
1992)
Late Campanian-Early Maastrichtian, Late Cretaceous
Gres a Reptiles Formation, Var, France
Material- (MDE-D203) femur (~230 mm)
Comments- Le Loeuff et al. (1992) described a femur (MDE-D203) and anterior
dorsal vertebra (MDE-D01) which they believed was congeneric or at least related
to Elopteryx. Le Loeuff et al. believed these remains were most closely
related to dromaeosaurids, though perhaps deserving their own family or subfamily.
The femur was only stated to share general characteristics with Elopteryx
(reduced fourth trochanter, posterior trochanter, "shape and size")
while differing in having a linear capital ligament fossa and absent fourth
trochanter. It probably belongs to a distinct taxon of pennaraptoran.
Reference- Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992. The
first record of dromaeosaurid dinosaur (Saurichia, Theropoda) in the Maastrichtian
of Southern Europe: palaeobiogeographical implications. Bulletin de la Societe
Geologique de France. 163(3), 337-343.
unnamed possible pennaraptoran (Kessler and Jurcsak, 1984)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania
Material- (MTCO 14422; = MTCO-P 1503) incomplete long bone
References- 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.
Kessler and Jurcs�k, 1984. Fossil bird remains in the bauxite from Cornet
(Bihor county, Romania), Trav. Mus. Hist. Nat. Grigore Antipa, Bucharest. 25,
393-401.
Jurcsak and Kessler, 1986. Evolutia avifaunei pe teritoriul Romanei. Partea
I: Introducere (Evolution of the avifauna in the territory of Romania. Part
I: Introduction). Crisia. 16, 577-615.
Kessler and Jurcs�k, 1986. New contributions to the knowledge of Lower
Cretaceous bird remains from Cornet (Romania), Bucharest, Trav. Mus. Hist. Nat.
Grigore Antipa. 28, 290-295.
Jurcsak and Kessler, 1987. Evolutia avifaunei pe teritoriul Romanei. Partea
II: Morfologia speciilor fosile (Evolution of the avifaune in the territory
of Romania. Part II: Morphology of fossil species). Crisia. 17, 583-609.
Jurcsak and Kessler, 1988. Evolutia avifaunei pe teritoriul Romanei. Partea
III: Filogenie si sistematice (Evolution of the avifauna in the territory of
Romania. Part III: Phylogeny and systematics). Crisia. 18, 647-688.
Jurcsak and Kessler, 1991. The Lower Cretaceous paleofauna from Cornet, Bihor
County, Romania and its importance. Nymphaea. 21, 5-32.
Benton, Cook, Grigorescu, Popa and Tallodi, 1997. Dinosaurs and other tetrapods
in an Early Cretaceous bauxite-filled fissure, northwestern Romania. Palaeogeography,
Palaeoclimatology, Palaeoecology. 130(1-4), 275-292.
Pennaraptora indet. (Nessov, 1984)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Materal- (CCMGE 459/12457) manual ungual (?)I (Nessov, 1995)
(TsNIGRI 45/11915) humeral shaft (~73 mm) (Nessov, 1984)
(TsNIGRI 48/11915) long bone shaft (Nessov, 1984)
(TsNIGRI 49/11915) long bone shaft (Nessov, 1984)
(TsNIGRI 50/11915) long bone shaft (Nessov, 1984)
(ZIN PO 4826) posterior synsacrum (Nessov and Panteleev, 1993)
Comments- The humerus was originally a paratype of Zhyraornis kashkarovi
(Nessov, 1984). Kurochkin (1996) later disagreed, since the nutrient foramen
is located on the ventral shaft, apparently unlike enantiornithines (in which
he included Zhyraornis). The specimen preserves almost no morphological
features, besides being slender and curved with a thin-walled shaft. Scaled
to Ichthyornis, it might measure ~73 mm when complete. It is thus large
enough to come from a deinonychosaur or oviraptorosaur in addition to a bird,
and certainly preserves no characters which could exclude this possibility.
It is here referred to Pennaraptora indet. Isolated shafts of long bones (TsNIGRI
48/11915, 49/11915 and 50/11915) were also made paratypes of Z. kashkarovi.
These were not described or illustrated, and are similarly referred to Pennaraptora
indet..
Nessov and Panteleev (1993) figured and described a partial sacrum they referred
to Kuszholia sp. (ZIN PO 4826). Zelenkov and Averianov (2011) stated
this differs from Kuszholia in "the absence of a pleurocoel in the
posterior vertebra and in the shallow slitlike pleurocoel in the penultimate
vertebra", and while the pleurocoel shape appears similar, the absence
of a pleurocoel in the last sacral is indeed different. They believe the specimen
to be similar to Zanabazar, but I don't see any particular resemblence
and refer it to Pennaraptora incertae sedis here pending further study.
CCMGE 459/12457 was listed in the text as being an oviraptorosaur by Nessov (1995), in
which case it may be referrable to Kuszholia from the same formation.
Nessov also listed the possibility of it being a bird pedal ungual in the figure
caption however.
References- Nessov, 1984. [Upper Cretaceous pterosaurs and birds from
Central Asia]. Paleontologicheskii Zhurnal. 1, 47-57.
Nessov and Panteleev, 1993. On the similarity of the Late Cretaceous ornithofauna
of South America and central Asia. Trudy Zoologicheskogo Instituta, RAN. 252,
84-94.
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.
Kurochkin, 1996. A new enantiornithid of the Mongolian Late Cretaceous, and
a general appraisal of the Infraclass Enantiornithes (Aves). Russian Academy
of Sciences, special issue. 50 pp.
Zelenkov and Averianov, 2011. Synsacrum of a primitive bird from the Upper Cretaceous
of Uzbekistan. Paleontological Journal. 45(3), 314-319.
undescribed pennaraptoran (Turner, Hwang and Norell, 2007)
Berriasian-Barremian, Early Cretaceous
Huhteeg Svita, Mongolia
Holotype- (IGM coll.) proximal femur, proximal tibia, partial pes
Comments- Turner et al. (2007) refer this specimen to Paraves based on
the lateral ridge and posterior trochanter. However, both are also present in
Avimimus, suggesting it cannot be placed more precisely than Pennaraptora
until it is further prepared and described.
Reference- Turner, Hwang and Norell, 2007. A small derived theropod from
Oosh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates. 3557,
27 pp.
undescribed pennaraptoran (Tanaka, Kobayashi, Sasaki and Chiba, 2013)
Santonian, Late Cretaceous
Uge Member of the Taneichi Formation, Japan
Material- feather fragments
Reference- Tanaka, Kobayashi, Sasaki and Chiba, 2013. An isolated feather
in an amber from the Late Cretaceous of northeast Japan. Journal of Vertebrate
Paleontology. Program and Abstracts 2013, 223-224.
undescribed pennaraptoran (Novas, Cladera and Puerta, 1996)
Cenomanian-Early Coniacian, Late Cretaceous
Rio Neuquen Subgroup, Neuquen, Argentina
Material- incomplete skeleton including humerus and pelvis
Comments- This specimen was mentioned in an abstract by Novas et al.
(1996) as having a bird-like humerus (e.g. anteriorly projecting deltopectoral
crest) and a propubic pelvis. It is seemingly not described yet, as Unenlagia's
skeleton is not very complete and its pelvis is mesopubic, while Buitreraptor
is also said to have a mesopubic pelvis and was found in Rio Negro.
Reference- Novas, Cladera and Puerta, 1996. New theropods from the Late
Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 16(3), 56A.
unnamed possible Pennaraptora (Novas, Borges Ribeiro and Souza Carvalho,
2005)
Late Maastrichtian, Late Cretaceous
Serra da Galga Formation of the Bauru Group, Brazil
Material- (CP 659) manual ungual (Novas, Borges Ribeiro and Souza Carvalho,
2005)
(MCT 1718-R) scapula (Machado, Campos and Kellner, 2008)
Comments- In 2021 the Serra da Galga and Ponte Alta Members of the Marilia Formation were recognized as the Serra da Galga Formation.
References- Machado, Kellner and Campos, 2005. On a theropod scapula
from the Late Cretaceous (Bauru Group) of Brazil. Journal of Vertebrate Paleontology.
25(3), 86A.
Novas, Borges Ribeiro and Souza Carvalho, 2005. Maniraptoran theropod ungual
from the Mar�lia Formation (Upper Cretaceous), Brazil. Revista del Museo
Argentino Ciencias Naturales "Bernadino Rivadavia". 7, 31-36.
Machado, Campos and Kellner, 2008. On a theropod scapula (Upper Cretaceous)
from the Mar�lia Formation, Bauru Group, Brazil. Palaeontologische Zeitschrift.
82(3), 308-313.
unnamed pennaraptoran (Benson, Rich, Vickers-Rich and Hall, 2012)
Early-Mid Aptian, Early Cretaceous
Wonthaggi Formation of the Strzelecki Group, Victoria, Australia
Material- (NMV P216672) (adult) mid or posterior dorsal vertebra (17 mm)
Reference- Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna
from southern Australia indicates high polor diversity and climate-driven dinosaur
provinciality. PLOS One. 7(5), e37122.
unnamed Pennaraptora (Britt, 1993)
Late Aptian-Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P186302) dorsal vertebra (23 mm) (Britt, 1993)
(NMV P186323; paratype of Timimus hermani) femur (195 mm) (Rich and Vickers-Rich,
1994)
Comments- Britt (1993) mentions NMV 186303 (the holotype femur of Timimus)
as a dromaeosaurid dorsal vertebra. This may be a typo for NMV 186302. Currie
et al. (1996) first described this vertebrae as oviraptorosaurian, while Salisbury
et al. (2007) placed it in Paraves, and Agnolin et al. (2010) believed it was
dromaeosaurid. Benson et al. (2012) could not distinguish it from paravians
or oviraptorosaurs, and it is here left as Pennaraptora incertae sedis pending
further study.
NMV P186323 was originally a paratype of Timimus hermani,
but was later determined to be maniraptoran (Benson et al., 2012).
Carrano (1998) lists TMP 1991.040.0016 as Tetanurae indet., but the TMP
online catalogue indicates this is a cast of a theropod femur from the
Otway Group, whose length in Carrano (195.1 mm) matches the Timimus paratype.
References- Britt, 1993. Pneumatic postcranial bones in dinosaurs and
other archosaurs. PhD Thesis, University of Calgary. 383 pp.
Rich and Vickers-Rich, 1994. Neoceratopsians and ornithomimosaurs: Dinosaurs
of Gondwana origins? National Geographic Research. 10(1), 129-131.
Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria)
specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia.
Alcheringa. 20(1-2), 73-79.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424
pp.
Salisbury, Agnolin, Ezcurra and Pias, 2007. A critical reassessment of the Creaceous
non-avian dinosaur faunas of Australia and New Zealand. Journal of Vertebrate
Paleontology. 27(3), 138A.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from Southern Australia
indicates high polor diversity and climate-driven dinosaur provinciality. PLOS
One. 7(5), e37122.
Ilerdopteryx Lacasa-Ruiz, 1985
I. viai Lacasa-Ruiz, 1985
Late Berriasian-Early Barremian, Early Cretaceous
La Pedrera de Rubies Lithographic Limestones Formation, Spain
Syntypes - (LP-715 IEI) body feather (27 mm)
(LP-1327 IEI) body feather
(LP IEI coll.) seven body feathers (20-30 mm)
Comments- These feathers have barbules, so are probably from pennaraptorans.
They may be from Noguerornis or the unnamed La Pedrera juvenile enantiornithine
taxon which are from the same locality. Whether all of the nine feathers are
from the same taxon is unknown, and Ilerdopteryx is indeterminate since
feathers are undiagnostic for Mesozoic theropods.
References- Lacasa-Ruiz, 1985. Nota sobre las plumas fosiles del yacimiento
eocretacico de 'La Pedrera-La Cabrua' en la sierra del Montsec. (Prov. Lleida,
Espana). Ilerda. 46, 227-238.
Lacasa-Ruiz, 1986. Nota preliminar sobre el hallazgo de restos keos de un ave
fosil en el yacimiento neocomiense del Montsec. (Prov .Lerida, Espafia). Ilerdu.
47, 203-206.
Lacasa-Ruiz, 1989. An Early Cretaceous fossil bird from Montsec Mountain (Lleida,
Spain). Terra Nova. 1(1), 45-46.
Kellner, 2002. A review of avian Mesozoic fossil feathers. In Chiappe
and Witmer (eds.). Mesozoic Birds - Above the Heads of Dinosaurs. University
of California Press. 389-404.