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. Beiträge zur geschichte der Archosaurier. Geologie und Paläontologie 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 Paläoherpetologie/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, Eichstätt 1984. Freunde des Jura-Museums Eichstätt, Eichstätt. 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.
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 Osmólska (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 Neuquén Group, Río 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 Società 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 Osmólska (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-Maximi­lians-Universität München. 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-Universität 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 Paléontologie. 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" Sayão, Saraiva, Brum, Bantim, Andrade, Cheng, Lima, Paula Silva and Kellner, 2020
"A. museunacionali" Sayão, 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 Sayão 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 Sayão 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" Sayão 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. Sayão et al. added the specimen to Choiniere's coelurosaur analysis, recovering it as sister to Zuolong, the pair being the first branching coelurosaurs.
Reference- Sayão, 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 coelurosaur (USNM online)
Late Aptian, Early Cretaceous
Dinosaur Park / Cherokee-Sanford Brick Clay Pit / Muirkirk Clay Pit USNM 41614, Arundel Formation, Prince George's County, Maryland, US
Material- (USNM 540736) distal pedal phalanx II-2 (~14.0 mm dorsovent distally) (USNM online)
Comments- This was discovered between 2010 and 2013 and identified as a coelurosaur pedal phalanx II-2 by Carrano, probably due to its elongation.

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 für Geologie und Paläontologie 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-Omeñaca 2001)
Tithonian-Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (LC-1) partial tooth (?x15.1x10.2 mm) (Suñer, Santisteban and Galobart, 2005)
(MPZ01/98) partial tooth (Barco and Ruiz-Omeñaca 2001)
References- Barco and Ruiz-Omeñaca, 2001. Primeros dientes de terópodos (Dinosauria, Saurischia) en la Formación Villar del Arzobispo (Titónico-Berriasiense): Yacimientos Cuesta Lonsal y Las Cerradicas 2 (Galve, Teruel). Publicaciones del Seminario de Paleontología de Zaragoza. 5, 239-246.
Suñer, Santisteban and Galobart, 2005. Nuevos restos de Theropoda del Jurásico Superior-Cretácico Inferior de la Comarca de Los Serranos (Valencia). Revista Española 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 terópodos del Cretácico Inferior (Fm. Arcillas de Morella, Aptiense) en los yacimientos del Mas de la Parreta (Morella, Castellón). In Fernández-Martínez (ed.). XXII Jornadas de Paleontología de la Sociedad Española de Paleontología. Libro de resúmenes, 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). Paläontologische Zeitschrift. 89(1), 79-94.

possible Coelurosauria (Jurcsák, 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 Jurcsák (19820 and Jucsak and Popa (1983). They may not belong to the same taxon, and the relationship of either with Aristosuchus is uncertain.
References- Jurcsák, 1982. Occurrences nouvelles des Sauriens mesozoiques de Roumanie. Vertebrata Hungarica. 21, 175-184.
Jurcsák 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-Omeñaca, 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, Métais, 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, Río 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 Neuquén Subgroup, Neuquén, 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 Rincón de los Sauces. Actas de las IV Jornadas Internacionales sobre Paleontología 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, São 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 Técnica de Yacimientos Petrolíferos 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 Società 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. 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. Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it as a basal carcharodontosaurid sister to Neovenator.
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 Osmólska (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.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.

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. Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it as a carcharodontosaurid (when megaraptorans were tyrannosauroids) more derived than Neovenator, but outside Acrocanthosaurus plus carcharodontosaurines (note the supplementary information shows a less precise position was recovered than suggested in Figure 25).
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.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.

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, Río 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 Osmólska. 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 Cretáceo 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, Martínez, 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 Neuquén Subgroup, Neuquén, 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, Martínez, Luna and Ibiricu, 2016)
(UNPSJB-PV 1046) manual ungual I (240 mm) (Casal, Martínez, Luna and Ibiricu, 2016)
....(UNPSJB-PV 1066) incomplete metatarsal III (Casal, Martínez, 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- CCasal, 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 Paleontología de Vertebrados, resumenes. Ameghiniana. 52(4) suplemento, 10-11.
Casal, Martínez, Luna and Ibiricu, 2016. Ordenamiento y caracterización faunística del Cretácico 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 Formación Lago Colhué Huapi, Grupo Chubut, Patagonia Argentina. Reunión de Comunicaciones de la Asociación Paleontológica Argentina 2019. R69.
Lamanna, Casal, Ibiricu and Martínez, 2019. A new peirosaurid crocodyliform from the Upper Cretaceous Lago Colhué 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, Neuquén, 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 terópodos de Auca Mahuevo, provincia del Neuquén (Cretácico tardío, 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 Paleontología de Vertebrados y VII Jornadas Técnicas de Paleontología de Vertebrados, Libro de Resúmenes). R7.
Méndez, Gianechini, Canale and Díaz-Martínez, 2019. A new megaraptorid specimen (Theropoda, Coelurosauria) from Cañadón Amarillo (Anacleto Formation, Campanian,
Upper Cretaceous), Mendoza Province, Argentina. 33.as Jornadas Argentinas de Paleontología de Vertebrados. 60-61.

unnamed Megaraptoridae (Novas, Agnolin, Rozadilla, Aranciaga-Rolando, Brissón-Eli, Motta, Cerroni, Ezcurra, Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento, García-Marsà, 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, Brissón-Eli, Motta, Cerroni, Ezcurra, Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento, García-Marsà, 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. Saurópodes e terópodes da Chapada dos Guimarães, Mato Grosso, Brasil. In Carvalho, Srivastava, Stroschschoen and Lana (eds.). Paleontologia: Cenários de Vida. 4th ed. Interciência. 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, Méndez, 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, Méndez, 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 Osmólska (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 Osmólska (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 Osmólska (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 Neuquén Subgroup, Neuquén, 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 Paleontología de Vertebrados. Santiago de Chile, Octubre del 2002. p.20.
Poblete and Calvo, 2003. Upper Turonian dromaeosaurid teeth from Futalognko quarry, Barreales Lake, Neuquén, 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 Neuquén, 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 Díaz-Martínez and Rábano (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 Apesteguía, 2011. Isolated archosaurian teeth from "La Bonita" locality (Late Cretaceous, Santonian-Campanian), Río 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 Paleontología 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 Neuquén Subgroup, Neuquén, 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 fósiles del Cretácico Superior de Neuquén. Ameghiniana. 38, 6R-7R.
Coria and Currie, 2002. Un gran terópodo celurosaurio en el Cretácico Superior de Neuquén. Ameghiniana. 39, 9R.
Paulina-Carabajal, 2009. El neurocráneo de los dinosaurios Theropoda de la Argentina: Osteología y sus implicancias filogenéticas. 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 Neuquén 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 Paleontología 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, Brissón-Eli, Motta, Cerroni, Ezcurra, Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento, García-Marsà, 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 Formación Pari Aike (Cretácico 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. Sedimentología y modelos deposicionales de la Formación Mata Amarilla, Cretácico 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, Brissón-Eli, Motta, Cerroni, Ezcurra, Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento, García-Marsà, 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
Tratayén, Bajo de la Carpa Formation, Neuquén, 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 right ilium, distal fused pubic boot fragment, partial right ischium
Santonian, Late Cretaceous
Barreales Sur, Bajo de la Carpa Formation, Neuquén, Argentina
Referred- ?(MCF-PVPH-399) distal right metatarsal II (Coria and Arcucci, 2004)
?(MCF-PVPH-418) posterior dorsal centrum (110 mm) (Coria and Arcucci, 2004)
Santonian, Late Cretaceous
Cerro Overo, Bajo de la Carpa Formation, Neuquén, Argentina
?(MAU-Pv-CO-659) several dorsal centra, several caudal centra, distal left humerus, proximal right tibia (Méndez, Paulina-Carabajal, Filippi, Gianechini, Cruzado-Caballero, Previtera, Barrios, Seculi Pereyra, Garrido and Pipo, 2021)
Santonian, Late Cretaceous
isthmus between Lago Barreales and Lago Mari Menuco, Bajo de la Carpa Formation, Neuquén, Argentina
?(MUCPv 1557) incomplete tooth (?x~16.0x~9.8 mm) (Porfiri, Baiano, dos Santos, Gianechini, Pittman and Lamanna, 2024)
Santonian, Late Cretaceous
La Invernada, Bajo de la Carpa Formation, Neuquén, Argentina
?(MAU-Pv-LI-548) two vertebrae (Mendez, Filippi and Garrido, 2015)
Santonian, Late Cretaceous
Bajo de la Carpa Formation, Neuquén?, Argentina
?(MCF-PVPH-654) (~9 m) right tibia (~800 mm) (Coria, Currie, Paulina Carabajal, Garrido and Koppelhus, 2004)
Santonian, Late Cretaceous
La Bonita, Bajo de la Carpa Formation, Río Negro, Argentina
?(MPCA 247) tooth (~21.69x14.53x7.8 mm) (Gianechini, Lio and Apesteguía, 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."
Mendez et al. (2021) note in an abstract in MAU-Pv-CO-659 "The distal end of the humerus exhibits a depression on the anterior surface - above the radial and ulnar condyles - which is also present (although markedly deeper) in Australovenator" and "the ulnar condyle appears to be greater than the radial condyle in distal view" as in Australovenator, both features being absent in Megaraptor. Also, "The development of the tibial cnemial crest is similar to that observed in Aerosteon, being larger than in other megaraptorids such as Australovenator, Fukuiraptor, Murusraptor, Orkoraptor, or Phuwiangvenator." Megaraptorid vertebral charactgers mentioned are "dorsal vertebral centra have large and deep pneumatic foramina on the lateral surfaces" and "caudal vertebral centra also have pneumatic foramina, less pronounced than those of the dorsal vertebrae, which exhibit an internal septum."
Porfiri et al. (2024) describe a tooth (MUCPv 1557) found 203 meters away from the Diuqin type. They state it has "a Crown Base Ratio ... of approximately 0.61", "The mesial carina lacks denticles" and distally "the apicalmost preserved denticles have a density of two per millimeter whereas that of the basal denticles is five per millimeter." As "the specimen strongly resembles the teeth of megaraptorids, particularly in its concave distal margin and lack of serrations on the mesial carina", they refer "the latter tooth to Megaraptoridae indet." Furthermore, they note "MUCPv 1557 shares with MPCA 247 the presence of mesiodistally oriented undulations (enamel wrinkles) extending across the labial and lingual surfaces of the crown" and find they "are closely comparable in all other respects as well." They conclude that based on "its stratigraphic and geographic provenance, we consider it plausible that MUCPv 1557 (and perhaps MPCA 247) could pertain to Tratayenia."
References- Coria and Arcucci, 2004. Nuevos dinosaurios terópodos de Auca Mahuevo, provincia del Neuquén (Cretácico tardío, Argentina). Ameghiniana. 41, 597-603.
Coria, Currie, Paulina Carabajal, Garrido and Koppelhus, 2004. ¿Un nuevo linaje de terópodos en el Cretácico Tardío de Argentina? XX Jornadas Argentinas de Paleontología de Vertebrados. 42R.
Porfiri, Calvo, Juárez Valieri and Santos, 2008. A new large theropod dinosaur from the Bajo de la Carpa Formation (Late Cretaceous) of Neuquén, Patagonia. Actas III Congresso Latinoamericano de Paleontología de Vertebrados. R202.
Gianechini, Lio and Apesteguía, 2011. Isolated archosaurian teeth from "La Bonita" locality (Late Cretaceous, Santonian-Campanian), Río 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 Paleontología 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 Paleontología de Vertebrados y VII Jornadas Técnicas de Paleontología de Vertebrados, Libro de Resúmenes). 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.
Méndez, Paulina-Carabajal, Filippi, Gianechini, Cruzado-Caballero, Previtera, Barrios, Seculi Pereyra, Garrido and Pipo, 2021. Megaraptoran remains from Cerro Overo-La Invernada fossil site (Bajo de la Carpa Formation, Santonian), northern Patagonia, Argentina. III Jornadas de Paleontología de la Cuenca Neuquina, Libro de Resúmenes. R261.
Porfiri, Baiano, dos Santos, Gianechini, Pittman and Lamanna, 2024. Diuqin lechiguanae gen. et sp. nov., a new unenlagiine (Theropoda: Paraves) from the Bajo de la Carpa Formation (Neuquén Group, Upper Cretaceous) of Neuquén Province, Patagonia, Argentina. BMC Ecology and Evolution. 24:77.

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 Osmólska (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.
Jurcsák (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.
Jurcsák, 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 Osmólska, 1996
B. ostromi Osmólska, 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 Osmólska. 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.
Osmólska, 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 Osmólska. 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 Osmólska (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 Società 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. Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover Coelurus as either sister to Maniraptoriformes, a basal ornithomimosaur or a basal therizinosaur.
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 Società 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.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.

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) uniquely 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 Società 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). Mémoires de la Société géologique 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.

Tyrannosauroidea

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 Società 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 Società 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 für Geologie und Paläontologie - 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, Araújo and Choiniere, 2019
= "Neocoelurosauria" Hendrickx, 2015
Definition- (Compsognathus longipes + Passer domesticus) (modified after Hendrickx, Mateus, Araújo 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, Araújo 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. 55, 7-19.

"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 Osmólska (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
Martínez 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 Società 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 Società 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.
Paul (2016) creates the term haplocheirids for a group including only Haplocheirus, though he never uses the technical term 'Haplocheiridae' that is implied. Nelson et al. (2012) used the taxon Haplocheiridae as a misspelling of the amphipod genus Haplocheira, but as there is a tribe Haplocheirini based on that genus (Myers and Lowry, 2003), the latter authors also implicitly created Haplocheiridae due to ICZN Article 36.1 ("A name established for a taxon at any rank in the family group is deemed to have been simultaneously established for nominal taxa at all other ranks in the family group"). Thus that word cannot be used for a family of theropods.
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 Gebrüder 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. 中国最早鸟类化石的发现及鸟类的起源. Chinese Geology. 23(10) (total issue 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.
Myers and Lowry, 2003. A phylogeny and a new classification of the Corophiidea Leach, 1814 (Amphipoda). Journal of Crustacean Biology. 23(2), 443-485.
Holtz, Molnar and Currie, 2004. In Weishampel, Dodson and Osmólska (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.
Nelson, Neill, Farr, Barr, D'Archino, Miller and Stewart, 2012. Rhodolith Beds in Northern New Zealand: Characterisation of Associated Biodiversity and Vulnerability to Environmental Stressors. Ministry for Primary Industries. 102 pp.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press. 360 pp.

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.
Martínez 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.
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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- Chen et al. (1998) list the English title of Ji and Ji (1996) as "On discovery of the earliest bird fossil in China and the origin of birds" while Downs' 2001 translation uses "On the discovery of the earliest fossil bird in China (Sinosauropteryx gen. nov.) and the origin of birds", but the actual paper has no English title. Chen et al.'s is closer, as Google Translate translates the Chinese title as "The discovery of the earliest bird fossils in China and the origin of birds", and it certainly doesn't say "Sinosauropteryx gen. nov." which would be "中华龙鸟属新属".
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. 中国最早鸟类化石的发现及鸟类的起源. Chinese Geology. 23(10) (total issue 233), 30-33.
Ji and Ji, 1997. 中华龙岛(Sinosauropteryx) 化石研究新进展. Chinese Geology. 24(7) (total issue 242), 30-32, 49.
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, Apesteguía, 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 Osmólska (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 Società 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. Über einige im lithographischen Schiefer neu aufgefundene Schildkröten 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 für Mineralogie, Geologie und Paläontologie. 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 Muséum 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 supérieur du Sud-est de la France), position phylogénétique, relation avec Archaeopteryx et implications sur l’origine théropodienne des oiseaux. PhD thesis. Muséum 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 Società 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 München. 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
Comments- Currie and Chen (2001) lists the title as "Advances in the study of the avian Sinosauropteryx prima", but the actual paper has no English title and Google Translate translates the Chinese title to "New progress in the study of Sinosauropteryx fossils" which seems more accurate as the species name is certainly never mentioned.
This was described as a new specimen of Sinosauropteryx prima by Ji and Ji (1997) and photographed on pages 78-79 of Ackerman (1998). 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. 中华龙岛(Sinosauropteryx) 化石研究新进展. Chinese Geology. 24(7) (total issue 242), 30-32, 49.
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 Società 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, Osmólska 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, Araújo 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, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Makovicky, Kobayashi and Currie, 2004. Ornithomimosauria. In Weishampel, Dodson and Osmólska (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, Araújo 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
Laño, 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 Laño 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 Laño 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 terópodos del Cretácico Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Paleontología. 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, Métais, 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-Omeñaca, 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, Métais, 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, Osmólska 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, Eichstätt 1984. Freunde des Jura-Museums Eichstätt, Eichstätt. 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 für Geologie und Paläontologie 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, Osmólska 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 Osmólska 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. Gebrüder 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.
Osmólska and Barsbold, 1990. Troodontidae. In Weishampel, Dodson and Osmólska (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 für Geologie und Paläontologie. 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 für Geologie und Paläontologie 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-Universität 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' which has never been published, and using "all other dinosaurs except oviraptorosaurs" as a specifier would be problematic. In addition 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"), absent 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 fósiles de la Formación Rio Colorado, de la Ciudad de Neuquén y Cercanías, Cretácico Superior, Argentina. Revista del Museo Argentino de Ciencias Naturales "Bernardino Rivadavia" e Instituto Nacional de Investigación de las Ciencias Naturales: Paleontologí. 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 Osmólska (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 Società 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.

Alvarezsauroidea

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. Paul (2024) continues his use of the informal term for maniraptorans excluding scansoriopterygids (and including alvarezsauroids and therizinosaurs), although this time he defines aveairfoilans as "airfoilans with avian shoulder girdles, or ancestors with same that are in the clade that includes extant birds", which seems like a retooling of his 2002 Avepectora. It's debatable where between Maniraptoriformes and Eumaniraptora this evolved , but in any case 'Aveairfoila' is still not used and so is still informal eight years later.
Paul (2024) creates the term airfoilan for "neocoelurosaurs with forelimb-borne airfoils of some form, or ancestors with same that are in the clade that includes extant birds", consisting of traditional maniraptorans (including alvarezsauroids and therizinosaurs). Thus content-wise this should be under Maniraptora, but definition-wise alvarezsauroids and therizinosaurs cannot be scored yet, so the name has an uncertain position similar to Aviremigia. The difference is that Paul includes scansoriopterygid membrane wings as "forelimb-borne airfoils of some form", but if scansoriopterygids were truly outside the clade of taxa with feather-based airfoils as Paul suggests it makes no sense to homologize membrane and feather wings as a shared evolutionary development. In any case, the implied technical term 'Airfoila' is never used and as no other publication has used the term for a theropod clade either, it remains informal.
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 für Geologie und Paläontologie 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.
Paul, 2024. The Princeton Field Guide to Dinosaurs 3rd edition. Princeton University Press. 384 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 Jurcsák, 1984)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania
Material- (MTCO 14422; = MTCO-P 1503) incomplete long bone
References- Kessler and Jurcsák, 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 Jurcsák, 1984. Fossil bird remains in the bauxite from Cornet (Bihor county, Romania), Trav. Mus. Hist. Nat. Grigore Antipa, Bucharest. 25, 393-401.
Jurcsák 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 Jurcsák, 1986. New contributions to the knowledge of Lower Cretaceous bird remains from Cornet (Romania), Bucharest, Trav. Mus. Hist. Nat. Grigore Antipa. 28, 290-295.
Jurcsák 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.
Jurcsák 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.
Jurcsák 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 Neuquén Subgroup, Neuquén, 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 Río 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 Marília 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 Marýlia 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 Marília 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.