Coelurosauria Huene, 1914
Definition- (Ornithomimus velox <- Allosaurus fragilis, Carcharodontosaurus saharicus) (suggested)
Other definitions- (Passer domesticus <- Allosaurus fragilis) (Holtz et al., 2004; modified from Holtz, 1996; modified from Gauthier, 1986)
(Passer domesticus <- Allosaurus fragilis, Sinraptor dongi, Carcharodontosaurus saharicus) (Sereno, in press)
= Coelurosauridae Cope, 1882
= Coelurosauria sensu Gauthier, 1986
Definition- (Passer domesticus <- Allosaurus fragilis) (modified)
= Coelurosauria sensu Sereno, in press
Definition- (Passer domesticus <- Allosaurus fragilis, Sinraptor dongi, Carcharodontosaurus saharicus)
Comments- Sereno's (in press) 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. This wouldn't necessarily be a bad thing, especially as most of the evidence indicates tyrannosauroids are basal to anything else called a coelurosaur nowdays except perhaps Tugulusaurus, Coelurus, Tanycolagreus, Calamosaurus, Proceratosaurus, Bagaraatan and Dryptosaurus. The latter is especially true if carcharodontosaurids have anything to do with tyrannosaurids, as that set of character transformations leaves tyrannosaurids developing their coelurosaurian characters convergently with maniraptoriformes. So the only topology which suffers by using Carcharodontosaurus as an external specifier is Paul's (1988), which would exclude compsognathids, Coelurus, and Ornitholestes from Coelurosauria (in addition to Proceratosaurus and tyrannosauroids). But since Allosaurus would have to be a coelurosaur for any of these latter taxa to be coelurosaurs in Paul's topology, Carcharodontosaurus' inclusion as an external specifier doesn't add any further harm. 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.

Coelurosauria incertae sedis

Iliosuchidae Paul, 1988
Iliosuchus Huene, 1932
I. incognitus Huene, 1932
= Megalosaurus incognitus (Huene, 1932) Romer, 1966
Middle Bathonian, Middle Jurassic
Stonesfield Slate, England
Holotype- (BMNH R83) incomplete ilium
Referred- (OUM J28971) incomplete ilium (Galton and Molnar, 2005)
(OUM J29780) partial ilium (~93 mm) (Galton, 1976)
Comments- This taxon is traditionally allied with Stokesosaurus and thus with tyrannosauroids due to the vertical ilial ridge. However, this character and the concave anterior edge of the pubic peduncle are widespread among basal coelurosaurs. The ilial ridge was lost in coelurosaurs as close to birds as compsognathids though, so Iliosuchus is probably less derived than that family.
References- Huene, 1932. Die fossile Reptile-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monogr. Geol. Palaeontol. (Pt. I and II, Ser. I) 4, 1-361.
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.
Galton and Molnar, 2005. Tibiae of small theropod dinosaurs from Southern England. In Carpenter (ed). The Carnivorous Dinosaurs. pp. 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.

unnamed Coelurosauria (Ostrom, 1970)
Late Aptian, Early Cretaceous
Cloverly Formation, Montana, Wyoming, US
Material- (AMNH coll.) pedal phalangeal fragment
(YPM 5174) incomplete metatarsal II
(YPM 5284) incomplete metatarsal IV
(YPM 5286) incomplete pedal ungual (~35 mm)
Comments- Ostrom (1970) referred these to Ornithomimus sp. based on being supposedly virtually indistinguishable from O. velox, though he did not the ungual was not identical to ornithomimids'. However, Holtz (1992) noted the metatarsals were more robust than ornithomimids', that there are no facets or buttresses for a wedge-shaped metatarsal III, and that metatarsal II resembled Allosaurus and Ornitholestes posteriorly more than it does ornithomimids. Indeed, judging by Ostrom's figure, metatarsal II differs from Ornithomimus in having a less ventrally oriented medial condyle and more proximally placed ventral convexity, while metatarsal IV lacks the marked posterior buttress and has more transversely expanded condyles. The pedal ungual has a more developed posterodorsal process, more obvious side grooves, and less developed side flanges. It resembles Dromiceiomimus in these characters except for the side flange development. Even basal ornithomimosaurs like Harpymimus, Garudimimus and IVPP V12756 seem more like ornithomimids in having distally placed ventral convexity on metatarsal II and less expanded distal condyles on metatarsal IV, so the metatarsals may be from a taxon like Nedcolbertia, Microvenator or another coelurosaur. The ungual does resemble those of ornithomimids, but not those of basal ornithomimosaurs. The specimens did not belong to the same individuals, and may be from different taxa as well.
References- Ostrom, 1970. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Peabody Mus. Nat. Hist., Yale Univ., Bull. 35, 234 pp.
Holtz, 1992. An unusual structure of the metatarsus of Theropoda (Archosauria: Dinosauria: Saurischia) of the Cretaceous. PhD Thesis, Yale University. 347 pp.

undescribed coelurosaur (Kirkland, Lucas and Estep, 1998)
Early Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- 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) stated the tibia of a small theropod is known, 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.
https://scientists.dmns.org/sites/kencarpenter/Cedar%20Mountain%20storage/Dinosaurs.aspx

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.

Gasosaurus Dong and Tang, 1985
G. constructus Dong and Tang, 1985
Bathonian-Callovian, Middle Jurassic
Xiashaximiao Formation, Sichuan, China
Holotype- (IVPP V7264) four cervical vertebrae, seven dorsal vertebrae, (sacrum 279 mm) first sacral vertebra (68 mm), second sacral vertebra (59 mm), third sacral vertebra (41 mm), fourth sacral vertebra (52 mm), fifth sacral vertebra (63 mm), seven caudal vertebrae, humerus (237 mm), ilium (369 mm), pubis (332 mm), ischium (338 mm), femur (425 mm), tibia (370 mm), fibula, metatarsal II, metatarsal III
Paratype- ?(IVPP V7265) three teeth
Referred- material (Holtz, 2000)
Comments- Traditionally associated with megalosauroids, Holtz (2000) recently found it to be a basal coelurosaur in his analyses. This was based on the upturned femoral head, anterior trochanter cleft from the head, and proximal fibula being >75% the proximal tibial width. However, he also indicated new undescribed specimens suggest Gasosaurus is a carnosaur, perhaps a sinraptorid (Currie pers. comm. 1998 to Holtz). Entering Gasosaurus into my theropod supermatrix strengthens Holtz's coelurosaurian identification with further synapomorphies including platycoelous cervical centra, a preacetabular process subequal in length to the postacetabular process, rounded postacetabular process, and anterodorsally concave pubic peduncle. The short tibia may indicate it is more basal than tyrannoraptorans and Tugulusaurus. More detailed descriptions and more information on the new specimens could lead to a different placement, however.
References- Dong and Tang, 1985. A new Mid-Jurasic theropod (Gasosaurus constructus gen et sp. nov.) from Dashanpu, Zigong, Sichuan Province, China. Vertebrata PalAsiatica. 23(1), 77-82.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. Gaia. 15, 5-61.

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 ran Xinjiangovenator in an analysis that resulted with it being placed sister to Bagaraatan inside Paraves. They assigned it to Maniraptora incertae sedis. When added to a modified version of Senter's (2007) matrix, including the characters from Rauhut and Xu (2005), Xinjiangovenator has a highly unstable position. It is at least as derived as Fukuiraptor, Tugulusaurus and tyrannoraptorans, yet can be excluded from Tyrannosauroidea, Ornithomimosauria, Alvarezsauridae, Therizinosauria and Paraves. This makes it most probably an oviraptorosaur or basal coelurosaur.
References- Dong, 1973. Reports of paleontological expediation 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.

Australovenator Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliot, 2009
A. wintonensis Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliot, 2009
Late Albian, Early Cretaceous
Winton Formation, Queensland, Australia
Holotype- (AODF 604) (~4.8 m) dentary (259 mm), two anterior teeth (13.83, 14.62 mm), seven dentary teeth (12.53, 20.85, 17.21, 21.97, 24.91, 23.36 mm), proximal first dorsal rib, proximal second or third dorsal rib, proximal seventh or eighth dorsal rib, dorsal rib shaft fragments, nine gastralial fragments, radius (213 mm), ulnae (269, 268 mm), metacarpal I (78.38 mm), phalanges I-1 (117.82, 117.54 mm), incomplete manual ungual I (150.95 mm straight, 190 mm on curve), metacarpal II (138.42 mm), phalanx II-2 (74.51 mm), distal manual ungual II, phalanx III-3 (43.8 mm), manual ungual III (75.12 mm straight), partial ilium, femur (578 mm), tibiae (569, 564 mm), fibula (538 mm), astragalus (105 mm wide), metatarsal I (66 mm), metatarsal II (284 mm), phalanx II-1 (113.02 mm), phalanx II-2 (81.06 mm), pedal ungual II (71.07 mm), metatarsal III (322 mm), phalanx III-2 (76.94 mm), phalanx IV-2, phalanx IV-3 (40.69 mm), pedal ungual IV (66.29 mm)
Diagnosis- (after Hocknull et al., 2009) eighteen 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 Tugulusaurus and Xiongguanlong.
Comments- 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. When included in a larger saurischian supermatrix, I find Australovenator to be a basal coelurosaur related to Fukuiraptor and Chilantaisaurus. Of the characters Hocknull et al. note as shared with Allosaurus and carcharodontosaurids, a slightly posteriorly forked dentary is present in most coelurosaurs, an elongate mediodistal femoral crest is present in Chilantaisaurus, Fukuiraptor, Stokesosaurus and Guanlong, and a constricted lateral tibial condyle is present in Fukuiraptor, Bagaraatan and tyrannosauroids. Finally, of the two characters placing it in Carcharodontosauridae, a distally extending lateral tibial malleolus (>5% of tibial length) is present in Chilantaisaurus and tyrannosauroids (partially correlated with tibial robusticity and thus with size), and a dorsally directed femoral head is also present in Chilantaisaurus, Gasosaurus, Bagaraatan and tyrannosauroids.
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. doi:10.1371/journal.pone.0006190

"Allosaurus" "robustus" Chure, 2000 vide Glut, 2003
Early Aptian, Early Cretaceous
Wonthoggi Formation of Strzelecki Group, Victoria, Australia
Material- (NMV Pl50070) (~4.8 m) astragalus (~108 mm wide)
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 Tugulusaurus and Coelurus. Differs from Tugulusaurus in plesiomorphically having an anterior transverse condylar groove. 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. based on six characters. Chure (2000) noted these all have a broader distribution. The fibular facet on the ascending process is also found in Torvosaurus, Poekilopleuron and coelurosaurs. The moderately high ascending process is actually higher than Allosaurus and more comparable to coelurosaurs, while its restriction to the lateral astragalar body is plesiomorphic but also found in the basal coelurosaurs Tugulusaurus and Coelurus. The presence of an inflection in the ascending process' medial margin cannot be determined with certainty, but is also present in Fukuiraptor and Australovenator. A medial condyle that is markedly larger than the lateral condyle is found in most carnosaurs and basal coelurosaurs. The supposed calcaneal notch seen in "robustus" is only represented by a small area due to breakage, and was interpreted as ambiguous by Chure as it is not confluent with the anterior condylar groove. Chure notes the calcaneal pit described is also present in a wide variety of tetanurines, also including coelurosaurs such as Fukuiraptor, Coelurus and Appalachiosaurus. The upper horizontal groove extending across the base of the ascending process is also found in basal ceratosaurs, megalosauroids, and most carnosaurs and coelurosaurs. Molnar et al. did note six characters of "robustus" which differ from Allosaurus. While they later (1985) stated these were apomorphies, the four which I can verify are present have a wider distribution. The horizontal condylar groove is shallower, which is also like most coelurosaurs except for Coelurus. The vertical groove on the posterior surface of the ascending process has a wide distribution, being found in such taxa as Torvosaurus, Sinraptor, Fukuiraptor, Australovenator, Coelurus and Appalachiosaurus. The absence of a posteroventral pit on the ascending process is like most theropods, though Acrocanthosaurus and Mapusaurus also have a pit. A laterally shifted anterior ridge defining the fibular surface of the ascending process is also present on those coelurosaurs which show fibular overlap (e.g. Fukuiraptor, Coelurus, Appalachiosaurus). The deeper tibial sulcus is hard to judge from figures. The ventral sillouette does not appear more symmetrical. Welles (1983) disputed the identification as Allosaurus, listing nineteen differences and suggesting a relationship to ornithomimids instead. Most were due to misinterpretation from breakage (Molnar et al., 1985), but additional valid differences he noted include the deeper horizontal groove at the base of the ascending process with sharper anterior edge, which are also present in Fukuiraptor. Chure (2000) additionally notes the ascending process is taller and with more parallel sides than Allosaurus, which are characters similar to coelurosaurs. He also finds "robustus" lacks an extensive anterior depression on the ascending process, which may be a carnosaurian character as it is also present in Sinraptor and Mapusaurus, but those of most coelurosaurs are shallower. The thicker medial edge of the ascending process in Allosaurus is a plesiomorphy lacking in coelurosaurs, as they lack the astragalar buttress on the tibia. Chure (2000) simply referred the astragalus to Avetheropoda based on the tall ascending process, though he noted it was not an allosaurid.
Azuma and Currie (2000) noted this astragalus is very similar to Fukuiraptor, and shares the characters listed above. Hocknull et al. (2009) note that their new taxon Australovenator is extremely similar to Fukuiraptor and "robustus" as well, and referred the latter to Australovenator sp.. Comparing all three, 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.
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.
Hocknull, White, Tischler, Cook, Calleja, Sloan and Elliot, 2009. New Mid-Cretaceous (Latest Albian) dinosaurs from Winton, Queensland, Australia. PLoS ONE. 4(7), e6190. doi:10.1371/journal.pone.0006190

Fukuiraptor Azuma and Currie, 2000
?= "Tsuchikurasaurus" unpublished
F. kitadaniensis Azuma and Currie, 2000
Barremian, Early Cretaceous
Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Holotype- (FPDM 97122, 96082443) (~4.2 m) (subadult) two maxillary fragments, two dentary fragments, premaxillary tooth (>17 mm), four maxillary teeth (12.6->37 mm), three dentary teeth (18.5-34 mm), five teeth, partial fifth cervical vertebra (58 mm), dorsal centrum (77.5 mm), dorsal neural arch, three proximal dorsal ribs, distal caudal vertebra (26.7 mm), coracoid (58 mm deep), humeri (242, 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, distal fibula, 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)
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-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-V9712229) maxillary fragment, tooth (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- (from Azuma and Currie, 2000) narrow dentary (~30% of depth) (also in Eotyrannus); teeth with oblique blood grooves (also in 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. A maxillary fragment, a dentary fragment, nine teeth, the cervical centrum, the dorsal neural arch and the 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. The posterior maxillary fragment figured in Azuma and Currie (2000) is actually a referred specimen (FPDM-V9712229) (Currie and Azuma, 2006). 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 coelurosaurian. 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.
Azuma and Currie (2000) found Fukuiraptor to be a basal carnosaur in their phylogenetic analysis, as did Holtz (2001) and Holtz et al. (2004). My unpublished theropod supermatrix (which incorporates all of Azuma and Currie's data) agrees with Longrich's (2001) analysis in placing the taxon as a very basal coelurosaur, primarily based on the tall astragalar ascending process, which has a broad base and extends medially. Adding it to a modified version of Senter's (2007) coelurosaur matrix results in a similar placement, with additional coelurosaur characters including the concavity at the base of the astragalar ascending process and elongate metatarsus.
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.
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.
http://dml.cmnh.org/2000Dec/msg00399.html
Holtz, 2001. Pedigree of the tyrant kings: New information on the origin and evolution of the Tyrannosauridae. Journal of Vertebrate Paleontology. 21(3), 62A-63A.
Longrich, 2001. Secondarily flightless maniraptoran theropods? Journal of Vertebrate Paleontology. 21(3), 74A.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Currie and Azuma, 2006. New specimens, including a growth series of Fukuiraptor (Dinosauria, Theropoda) from the Lower Cretaceous Kitadani Quarry of Japan. J. Paleont. Soc. Korea. 22(1), 173-193.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.
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. J. Paleont. Soc. Korea. 22(1), 173-193.

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.
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 tarsometatarsus and phalanges. Molnar thought the tibiofemoral ratio was too small for an ornithomimid. Rauhut and Xu (2005) found the taxon to be a coelurosaur more basal than tyrannoraptorans. They referred it to Coelurosauria based on the medial side of metacarpal I forming a sharp edge, the absence of deep extensor pits on its metacarpals, reduced fibular facet on the astragalus, groove at the base of the astragalar acsending process and absence of a groove across the astragalar condyles. It was more primitive than other coelurosaurs in having a laterally restricted astragalar ascending process which is low, and a tibia with a distinct step to brace the ascensing process. When entered into a modified version of Senter's (2007) matrix with Rauhut and Xu's data added, the results agree with theirs. Additionally, the high tibiofemoral ratio groups Tugulusaurus with coelurosaurs more derived than Gasosaurus.
References- Dong, 1973. Reports of paleontological expediation 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, Jefferson, NC. 1076pp.
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.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Tyrannoraptora Sereno, 1999
Definition- (Tyrannosaurus rex + Passer domesticus) (Holtz et al., 2004; modified from Sereno, 1999)

Aristosuchus Seeley, 1887
A. pusillus (Owen, 1876) Seeley, 1887
= Poekilopleuron pusillus Owen, 1876
= Poekilopleuron minor Owen vide Cope, 1878
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (BMNH R178) 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- ?(BMNH R178a) dorsal vertebra (21 mm)
?(BMNH R178b) two caudal vertebrae (28 mm)
?(BMNH R179) partial manual ungual
?(BMNH R899) partial manual ungual
Referred- ?(BMNH R5194) proximal femur (Galton, 1973)
?(BMNH R6426) proximal ischium (Naish, 2002)
?(MIWG 5137) tibia (Naish, 1999)
? incomplete pubis (Seeley, 1887)
References- Owen, 1876. Supplement (No. VII) to the Monograph on the Fossil Reptilia of the Wealden and Purbeck Formations. Palaeontographical Society Monograph. 30, 1-7.
Cope, 1878.
Seeley, 1887. On Aristosuchus pusillus Owen, being further notes on the fossils described by Sir R. Owen as Poikilopleuron pusillus. Owen. Q. J. Geol. Soc. London. 43, 221-228.
Galton, 1973. A femur of a small theropod dinosaur from the Lower Cretaceous of England. Journal of Paleontology. 47, 996-1001.
Naish, 1999. Theropod dinosaur diversity and palaeobiology in the Wealden Group (Early Cretaceous) of England: evidence from a previously undescribed tibia. Geologie en Mijnbouw. 78, 367–373.
Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: theropods. in Martill and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association, p. 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.
A? sp. (Jurcsak, 1982)
Berriasian-Hauterivian, Early Cretaceous
Bauxite of Cornet, Romania
Material- (MTCO 16499) cervical vertebra
(MTCO 17245) incomplete caudal vertebra
Comments- The two specimens may not belong to the same taxon, and the relationship of either with Aristosuchus is uncertain.
References- Jurcsak, 1982. Occurrences nouvelles des Sauriens mesozoiques de Roumanie. Vert. Hungarica. 21, 175-184.
Jurcsak and Popa, 1983. La faune de dinosauriens du Bihor (Roumanie). In: Buffetaut, F., Mazin, J. M., and Salmon, E. (eds.). Actes Syrup. Paleontol. Georges Cuvier. Moutbeliard, France. 325-335.

"Arkansaurus" Sattler, 1983
"A. fridayi" Braden, 1998
Aptian-Albian, Early Cretaceous
Trinity Group, Arkansas, US
Material- (UAM 74-16) metatarsal II (350 mm), phalanx II-1 (105 mm), phalanx II-2 (70 mm), metatarsal III (393 mm), phalanx III-1 (84 mm), phalanx III-2 (70 mm), metatarsal IV (355 mm), phalanx IV-1 (60 mm), three partial pedal unguals
Comments- A partial dorsal vertebra, caudal vertebra, scapular fragment and femoral fragment are also known from the Trinity Group, but were not associated with the specimen.
References- Quinn, 1973. Arkansas dinosaur. Geological Society of America Abstracts with Program. 5, 276-277.
Sattler, 1983. The Illustrated Dinosaur Dictionary. New York: Lothrop, Lee, and Shepard Books.
Braden, 1998. The Arkansas dinosaur "Arkansaurus fridayi". Arkansas Geological Commission.
Kirkland, Britt, Whittle, Madsen and Burge, 1998. A small coelurosaurian theropod from the Yellow Cat Member of the Cedar Mountain Formation (Lower Cretaceous, Barremian) of Eastern Utah. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 239-248.
Hunt, 2002. An Early Cretaceous theropod foot from Southwestern Arkansas as a possible track maker in Central Texas and Southwestern Utah. Journal of Vertebrate Paleontology. 22(3), 68A.
Hunt, 2003. An Early Cretaceous theropod foot from Southwestern Arkansas. Proceedings Journal of the 2003 Arkansas Undergraduate Research Conference. 87-103.

"Dryosaurus" grandis Lull, 1911
= Ornithomimus affinis Gilmore, 1920
= Coelosaurus affinis (Gilmore, 1920) Matthew and Brown, 1922
= Archaeornithomimus affinis (Gilmore, 1920) Russell, 1972
Late Aptian-Early Albian, Early Cretaceous
Arundel Formation, Maryland, US
Syntypes- ?(Goucher College 2609) phalanx
(USNM 5453) pedal phalanx II-1 (82 mm)
?(USNM 5652) partial astragalus (78 mm wide)
?(USNM 5684) distal metatarsal III
?(USNM 5703) pedal phalanx III-2 (69 mm)
?(USNM 5704) distal metatarsal II
?(USNM 6107) pedal ungual II (55.5 mm)
?(USNM 6108) pedal phalanx II-1 (79 mm)
?(USNM 8456) pedal phalanx IV-3 (38 mm)
Referred- ?(USNM 5701) distal caudal vertebra (68.7 mm) (Gilmore, 1920)
?(USNM 6115) distal end of proximal pedal phalanx (Gilmore, 1920)
?(USNM 6116) distal caudal vertebra (67.5 mm) (Gilmore, 1920)
Diagnosis- While the syntypes together provide a unique combination of characters (anterior transverse groove on astragalus; proximally extensive distal articular surface on metatarsal II; metatarsal III distally convex; curved pedal unguals with no side flanges), they may not belong to the same taxon.
Comments- Collected in the late 1880's, Marsh (1888) originally referred hindlimb and pedal elements to his new species Allosaurus medius, including USNM 5453 and 5652 as syntypes. Lull (1911) retained a tooth (USNM 4972) as the lectotype of Allosaurus medius, but noted USNM 5453 was "surely" ornithopod and USNM 5652 was "probably" so, making these and other material (USNM 5684, 5703 [mistyped as 5684 in the materals list], 5704, 6107- then unnumbered, and Goucher College 2609) syntypes of his new species Dryosaurus grandis. He also described USNM 6108 and 8456 as belonging to the taxon. Whether all of these remains were originally associated is unknown, though they are of the right size to be. USNM 5453 and 6108 are both left pedal phalanges II-1, so indicate at least two individuals are present. Gilmore realized they were coelurosaurian, assigning them to Ornithomimidae in a short note (1919) and making the USNM material syntypes of his new species Ornithomimus affinis (since the combination Ornithomimus grandis already existed) in an in depth description (1920). While the species name affinis has been used by everyone since, grandis has priority when not paired with Ornithomimus since it was named nine years earlier. It is not a nomen oblitum, as it was used as a valid name after 1899 (Article 23.9.1.1). If it is to be conserved, it must be referred to the ICZN for a ruling under the plenary power (23.9.3). Gilmore referred four more specimens to his species, but these are not comparable to the syntypes and may belong to other small theropod taxa. Matthew and Brown (1922) referred affinis to "Coelosaurus" (which they thought could be congeneric itself with Ornithomimus) based on geography and incorrectly thinking "Coelosaurus" antiquus lived earlier than Ornithomimus and Struthiomimus. Russell (1972) referred the species to Archaeornithomimus based on the curved pedal unguals, and Paul (1988) went along by placing it in Ornithomimus but the subgenus Archaeornithomimus. Notably both Gilmore and Paul had concepts of Ornithomimus equivalent to Ornithomimidae as used today. However, Smith and Galton (1990) noted the only preserved pedal ungual of Archaeornithomimus asiaticus is straight and that curved unguals from Iren Debasu may be Alectrosaurus instead. Smith and Galton further considered the Arundel ungual to be Theropoda indet. along with the rest of the specimens, though they noted the metatarsals were coelurosaur-grade. USNM 8456 was identified as phalanx IV-2 by Gilmore, but is more likely IV-3 based on its proportions.
"Dryosaurus" grandis differs from basal ornithomimosaurs (Harpymimus, Garudimimus, "Grusimimus" and IVPP V12756) in having a transverse groove on the anterior astragalar body, less bulbous medial condyle, less pointed medial edge of the distal articular surface on metatarsal II, more proximally extensive distal articular surface on the extensor aspect of metatarsal II, more slender pedal phalanges (though III-2 is comparable to "Grusimimus"), less pediculate ventral articular condyle on pedal phalanx II-1 which extends about as proximally as the dorsal peak, less expanded distal metatarsal III (both transversely and anteroposteriorly) with slightly convex distal edge, larger ligament pit on pedal phalanx III-2, and transversely narrower pedal ungual (except for "Grusimimus") with no side flanges. Pedal phalanx III-2 oddly seems to share the proximolateral depression with Garudimimus, which is supposedly an apomorphy of that genus. Yet Caudipteryx and Struthiomimus have it as well, so this may be more variable than Kobayashi and Barsbold believe. Derived ornithomimids like Archaeornithomimus and Struthiomimus are even less similar, with large ventrally extended side flanges on their unguals, though their articular surface on metatarsal II does extend proximally even further than in grandis. Compared to Archaeornithomimus specifically, grandis further differs in having much more concave distal astragalar margin, lacking a large flange medial to the distal condyles of metatarsal III, the distal articular surface of which is proximally peaked, and curved pedal unguals. Archaeornithomimus is similar in having a convex distal articular surface on metatarsal III though.
Among known American Early Cretaceous basal coelurosaurs, Nedcolbertia is difficult to compare, as the apparently complete foot of CEUM 5071 is only photographed articulated in anterior view, while that of CEUM 5072 is fragmentary. Yet it seems more similar in lacking the bulbous medial astragalar condyle, perhaps having a more rounded medial edge of distal metatarsal II, and more slender pedal phalanges and unguals. It still lacks the anterior astragalar groove and comparatively unexpanded distal metatarsal III (transversely at least), and has unguals with a larger flexor tubercle which are lower and straighter. "Arkansaurus" has a more transversely expanded metatarsal II, but the outline of metatarsal III is similar in anterior view, though in lateral view the articular end is less elongated posterodistally and the posterior shaft edge is not convex. Phalanx II-1 is far more expanded transversely and dorsally at its proximal end, while the best preserved ungual is generally similar in proportions and curvature, though it has ornithomimosaur-like side flanges. Ostrom's (1970) Cloverly Ornithomimus sp. cannot be compared except for the highly dissimilar pedal ungual, which is straight and very low with side flanges.
Tyrannosauroids like Appalachiosaurus differ from "Dryosaurus" grandis in similar ways that ornithomimosaurs do, except their metatarsal III is more similar in morphology (less expanded anteroposteriorly and laterally, with a slightly convex distal edge). Metatarsal III is also more similar in having the distal condyles elongated posterodistally with little anterior expansion (also somewhat present in IVPP V12756). The pedal unguals are less tapered in dorsal view and have side flanges though, and pedal phalanges are much stouter. "Dryosaurus" grandis is even more dissimilar to carnosaurs and Fukuiraptor (except in having the astragalar groove), lacks the ginglymoidy and specialized second digit of deinonychosaurs, the derived pedal characters of therizinosaurs, and bears no particular resemblence to basal oviraptorosaur material.
Based on these comparisons, grandis does not belong in Archaeornithomimus, Ornithomimus or "Coelosaurus", and seems more likely to be a basal tyrannosauroid or basal coelurosaur than an ornithomimosaur. Another possibility is that it is a chimaera, with the astragalus being (juvenile?) carnosaurian, the second metatarsal being ornithomimid, third metatarsal tyrannosauroid, pedal phalanges basal coelurosaurian, and the ungual tyrannosauroid or basal coelurosaurian. The Paleobiology Database lists USNM 5453 as the holotype, and it was the specimen assigned to Dryosaurus grandis most definitively by Lull, so perhaps the taxon should be assigned to Coelurosauria incertae sedis for now.
As for the referred specimens, the distal caudal USNM 5701 (referred to Allosaurus medius by Lull) is elongate unlike oviraptorosaurs and derived therizinosaurs, and has a neural spine unlike paravians and some basal coelurosaurs (Coelurus). Gilmore (1920) notes the prezygapophyses are restored too short, and were originally elongate as in carnosaurs, ornithomimosaurs, tyrannosauroids and some basal coelurosaurs (Ornitholestes, Juravenator). The amphicoelous centrum excludes alvarezsaurids from consideration. The caudal is more elongate than those of carnosaurs or tyrannosaurids, so may be ornithomimosaur or basal coelurosaur in nature. It cannot be definitely associated with the syntypes. The other caudal is apparently similar.
The vertebral centrum USNM 8454 was originally referred to the ankylosaur Priconodon by Lull (based on comparison to Stegosaurus), but provisionally reassigned to Ornithomimus affinis by Gilmore (1920), who thought it was a first sacral centrum. It is 69 mm long and from a young individual as the neural arch is unfused. It is certainly not an avetheropod posterior dorsal or sacral centrum, as it is much taller than wide, not very constricted ventrally nor transversely expanded anteriorly, lacks sutures for sacral ribs and has a ventral keel. These features are more similar to caudal vertebrae, but there seem to be no chevron facets. The lack of parapophyses excludes anterior dorsals and cervicals from consideration. Although ankylosaurs sometimes have transversely compressed anterior sacrals (e.g. Polacanthus) and a straighter ventral margin, they also have sacral rib facets and very wide neural canals unlike USNM 8454. Their dorsal vertebrae are universally wide while their caudals have transverse processes located on the centrum. Tenontosaurus has deeper vertebrae which are opisthocoelous (anterior dorsals) or broad (posterior dorsals, sacrals, proximal caudals), though the centra are keeled. Smaller ornithopods have longer centra, which are still far too broad, though more comparable in ventral concavity. Sauropods like Pleurocoelus have pleurocoels or lateral fossae in their centra, except for caudals which resemble those of ankylosaurs in having ventrally placed transverse processes. They further differ in being broader and opisthocoelous (presacrals) with no ventral keel. Thus while no close match can be found at the moment, it doesn't match theropod centra and seems more likely to be a small ornithischian. It is here excluded from grandis.
Neither Goucher College 2609 (which has been recatalogued in the USNM) nor USNM 6115 have been described or illustrated, so comparison is not possible.
References- Marsh, 1888. Notice of a new genus of Sauropoda and other new dinosaurs from the Potomac Formation. American Journal of Science (set 3). 35, 89-94.
Lull, 1911. Systematic paleontology of the Lower Cretaceous deposits of Maryland: Vertebrata. Maryland Geological Survey. Lower Cretaceous volume, 183-211.
Gilmore, 1919. An ornithomimid dinosaur in the Potomac of Maryland. Science. 1295, 394-395.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States National Museum with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. United States National Museum Bulletin. 110, l-154.
Gilmore, 1921. The fauna of the Arundel Formation of Maryland. Proceedings of the United States National Museum. 59(2389), 581-594.
Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History. 46(6), 367-385.
Ostrom, 1970. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Peabody Mus. Nat. Hist., Yale Univ., Bull. 35, 234 pp.
Russell, 1972. Ostrich dinosaurs of the Late Cretaceous of Western Canada. Canadian Journal of Earth Sciences. 9, 375-402.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Smith and Galton, 1990. Osteology of Archaeornithomimus asiaticus (Upper Cretaceous, Iren Dabasu Formation, People's Republic of China). Journal of Vertebrate Paleontology. 10(2), 255-265.

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
Paratype- ?(SAM P18010) pedal phalanx (44 mm)
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 Tugulusaurus, Fukuiraptor 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 arctometatarsalians. 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 based on the anterior ridge extending vertically medial to the ascending process, as seen in Quilmesaurus, Masiakasaurus and Velocisaurus. Kakuru seems to resemble basal arctometatarsalians the most, though the possibility it's another variety of non-maniraptoran tyrannoraptoran cannot be ruled out. It may even be an abelisauroid with an apomorphically elongate ascending process.
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, New York.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". in Weishampel, et al. (eds.), The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. pp. 280-305.
Rauhut, 2005. Post-cranial remains of ‘coelurosaurs’ (Dinosauria, Theropoda) from the Late Jurassic of Tanzania. Geol. Mag. 142(1), 97-107.

Santanaraptor Kellner, 1999
S. placidus Kellner, 1999
Albian, Early Cretaceous
Romualdo member of Santana Formation, Brazil
Holotype- (MN 4802-V) (~1.25 m; subadult) three mid caudal vertebrae, chevrons, ischia (91 mm), femora, tibia, fibula, pes, 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. The individual was probably about 1.25 meters long, assuming it resembled Ornitholestes in proportions, but was a subadult according to unfused vertebral sutures.
The caudal vertebrae appear to be mid caudals and have low, posteriorly oriented neural spines. The chevrons are said to be half the centrum length and expand slightly anteroposteriorly.
The ischia closely resemble Ornitholestes, differing in the longer and more distally placed obturator process, more concave posterior edge, unexpanded distal tip, and less proximally constricted obturator notch. The obturator process is triangular, there are no dorsal processes or proximolateral scar and the shaft is rod-shaped distally.
The proximal femur is figured, but the distal portion is just described. The shaft diameter is 13 mm and the bone thickness is 2.5 mm, so it's typically hollow. The wing-like anterior trochanter is separated from the greater trochanter, which it does not reach, by a cleft. The fourth trochanter is a low crest. The femur has three features unique to this species that are noted above in the diagnosis.
The tibia, fibula and pes are photographed (which doesn't show up well in my photocopy) and not described, although the fibula appears about a third as wide as the tibia. The metatarsus is typically theropod and said to be 70% of the femoral length. Metatarsals II and IV are said to be subequal and it can be seen not to be arctometatarsalian. Digits are also preserved, but no details can be observed.
Soft tissue is found on various parts of the fossil. The epidermis is very thin (~0.04 mm) and formed by irregular quadrangles separated by deep grooves. There are no scales or feathers preserved. Striated muscle fibers were preserved as calcium phosphate, are polygonal in transverse section and 30-50 micrometers in diameter. The bone still preserves channels for blood vessels (diameter 20-25 micrometers) and lacunae for osteocytes (diameter ~5 micrometers). There are also structures preserved that may either be mineralizations filling the bone's capillaries or replacements of the blood vessels. They are rod-like with a rough outer layer and smooth inner layer.
Kellner places this species in the Coelurosauria based on the triangular obturator process and suggests it may be a maniraptoriform based on Sereno's (1999) character "obturator notch U-shaped with slightly divergent sides", which I find highly variable. I agree that the triangular obturator process shows this is a coelurosaur. 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. The elongate and appressed metatarsals suggest it is a tyrannoraptoran, but the wing-like lesser trochantor excludes it from Maniraptora
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.
Agnolin, Apesteguia, and Chiarelli, 2004. The end of a myth: The mysterious ungual claw of Noasaurus leali. Journal of Vertebrate Paleontology. 24(3), 301A.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.

"Tonouchisaurus" Anonymous?, 1994
"T. mongoliensis" Anonymous?, 1994
Hauterivian-Barremian, Early Cretaceous
Shinekhudag Formation, Mongolia
Material- (~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 Huren-duh. 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., 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 over a decade 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", similar to basal coelurosaurs, and unlike ornithomimosaurs, oviraptorids and dromaeosaurids. 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.
References- Anonymous?, 1994. Japanese newspaper article.
Endo, DML 1994. http://dml.cmnh.org/1994Dec/msg00059.html
Holtz, DML 1994. http://dml.cmnh.org/1994Dec/msg00155.html
Olshevsky, DML 1995. http://dml.cmnh.org/1995Nov/msg00158.html

undescribed Coelurosauria (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, 375-402.

unnamed tyrannoraptoran (Holtz, 1992)
Early Campanian, Late Cretaceous
Merchantville Formation, New Jersey, US
Material- (YPM PU 21795) metatarsal II, metatarsal IV
Comments- Based on its age, this specimen is probably not Dryptosaurus. Holtz (1992) mentions it as a referred "Coelosaurus" specimen, noting it was subarctometatarsalian, more robust than ornithomimids, and had only a slight ridge on metatarsal IV to back metatarsal III. He states it does not resemble Dryptosaurus and regards it as a distinct taxon, though he refrains from naming it. Gallagher (1993) later published the specimen as Dryptosaurus sp..
References- Holtz, 1992. An unusual structure of the metatarsus of Theropoda (Archosauria: Dinosauria: Saurischia) of the Cretaceous. PhD Thesis, Yale University. 347 pp.
Gallagher, 1993. The Cretaceous/Tertiary mass extinction event in the northern Atlantic Coastal Plain: The Mosasaur. 5, 75-154.

undescribed possible coelurosaur (Wolfe and Kirkland, 1998)
Middle Turonian, Late Cretaceous
Lower Moreno Hill Formation, New Mexico, US
Material- (~2 m, ~10 kg; Fred) premaxilla, maxilla, partial lacrimal, jugal fragment, anterior dentary, teeth
(~2 m, ~10 kg) twenty-six vertebrae, manual elements, incomplete pubes, femora, tibiae, proximal fibulae, incomplete metatarsus
Comments- The first specimen was discovered in 1997, and reported by Wolfe and Kirkland (1998) as a "new small dromaeosaurid theropod" and a "small (dromaeosaurid?) theropod." An additional specimen was found in 2000, including vertebrae, partial forelimbs and incomplete pelves and hindlimbs. Kirkland and Wolfe (2001) referred to the taxon as a basal coelurosaur, as did Holtz in 2001 (DML). Indeed, casts of a composite skeleton are available to be purchased as the "Zuni coelurosaur". Pringle (2001) photographed much of the skeleton in her popular article, where the specmen was nicknamed Fred. In 2004, Denton et al. referred to it as a basal tetanurine based on several characters.
These characters are also found in some basal coelurosaurs however. The short extent of the antorbital fossa anterior to the antorbital fenestra is seen in Juravenator and most basal tyrannosauroids. Laterally compressed teeth with finely serrated mesial and distal carinae are found in tyrannosauroids. Isodont premaxillary and maxillary teeth are present in Compsognathus, Scipionyx and Huaxiagnathus. Weakly opisthocoelous cervical centra are present in basal tyrannosauroids and Scipionyx, while Compsognathus, eustreptospondylines, spinosauroids and carnosaurs have more strongly opisthocoelous centra. Most non-maniraptoriform coelurosaurs have distally placed anterior trochanters. A prominant fourth trochanter is found in some basal coelurosaurs such as Nedcolbertia and Guanlong. An absent femoral extensor groove is actually more derived than basal tetanurines (e.g. "Szechuanoraptor", Eustreptospondylus, Chilantaisaurus) and is found in Nqwebasaurus and many maniraptorans.
While the taxon may end up being more basal, it seems most likely to be a non-maniraptoriform coelurosaur
References- Wolfe and Kirkland, 1998. Zuniceratops christopheri n. gen. & n. sp., a ceratopsian dinosaur from the Moreno Hill Formation (Cretaceous, Turonian) of west-central New Mexico. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 307-317.
http://dml.cmnh.org/2001Jun/msg00634.html
Kirkland and Wolfe, 2001. First definitive therizinosaurid (Dinosauria; Theropoda) from North America. Journal of Vertebrate Paleontology. 21(3): 410–414.
Pringle, 2001. The creature from the Zuni lagoon. Discover. August, 42-48.
Denton, Nesbitt, Wolfe and Holtz, 2004. A new small theropod dinosaur from the Moreno Hill Formation (Turonian, Upper Cretaceous) of New Mexico. Journal of Vertebrate Paleontology. 24(3), 52A.

Tyrannosauroidea

Avipluma Clarke, Gauthier, de Queiroz, Joyce, Parham and Rowe, 2004
Definition- (hollow-based, branched, filamentous epidermal appendages homologous with Vultur gryphus) (Clarke, Gauthier, de Queiroz, Joyce, Parham and Rowe, 2004)
= Metatheropoda Ji and Ji, 2001
Comments- 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. In any case, Metatheropoda seems similar in suggested content to Clarke et al.'s (2004) Avipluma and Avifilopluma. Evidence increasingly suggests it is an invalid concept however, as Psittacosaurus, Tianyulong and pterosaurs indicate feathers may have originated before Ornithodira and thus be symplesiomorphic for theropods.

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.

Coeluria Marsh, 1881
Coeluroidea Marsh, 1881 sensu Nopcsa, 1928
Coeluridae Marsh, 1881
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) cervical vertebra, 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, four cervical vertebrae, five dorsal vertebrae, five dorsal neural arches, 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, metacarpal II, phalanx II-1, phalanx II-2, phalanx III-1, phalanx III-2, metacarpal IV fragment, ilial fragment, pubes, femora (one proximal) (~210 mm), distal tibia, proximal fibula, astragalus (32 mm wide), distal metatarsal III, metatarsal IV, fragments (Ostrom, 1980)
?(YPM 9162) partial sacral vertebra (Ostrom, 1980)
?(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.
The dorsal vertebrae listed by Carpenter et al. (2005) do not match those illustrated in the paper, as two vertebrae, five centra and five arches are listed. My materials list above contains the five vertebrae and five centra illustrated. In addition, only four cervical vertebrae are listed among the Coelurus specimens, yet five are illustrated. The extra vertebra is added to YPM 2010's materials list above. 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).
Several vertebrae referred to Coelurus fragilis by Gilmore (1920) were provisionally referred to the unnamed (?)enigmosaur 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. Bull. U. S. Nat. Mus. CX 1-154, 36 pls., 78 text-figs.
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, L. (ed.) Aspects of Vertebrate History. Flagstaff, 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, Univ. Copenhagen, 311pp.
Makovicky, 1997. A new small theropod from the Morrison Formation of Como Bluff, Wyoming. Journal of Vertebrate Paleontology. 17, 755-757.
Miles, Carpenter and Cloward, 1998. A new skeleton of Coelurus fragilis from the Morrison Formation of Wyoming. JVP 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.

Eotyrannus Hutt, Naish, Martill, Barker and Newbery, 2001
= “Gavinosaurus” Kelly, 1998
= “Lengosaurus” Kelly, 1998
= “Kittysaurus” Hargreaves, 2001
= “Fusinasus” Hutt, 2002
E. lengi Hutt, Naish, Martill, Barker and Newbery 2001
Early Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (MIWG 1997.550) (~4.5 m; subadult) premaxilla, premaxillary tooth, partial maxilla, nasals (220 mm), incomplete lacrimal, quadrate (94 mm), dentaries, partial surangular, teeth, axial neural arch, three cervical vertebrae (50 mm), six partial dorsal centra (45-55 mm), thirteenth dorsal centrum (64 mm), fourteenth dorsal centrum (52 mm), partial dorsal neural arch, six dorsal rib fragments, fifth sacral centrum (71 mm), six caudal vertebral fragments, scapulae (~285-295 mm), coracoid, humeri (235 mm), proximal radius, partial ulna, distal carpal I, metacarpal I, phalanx I-1, manual ungual I (~115 mm on curve), proximal metacarpal II, phalanx II-1 (70 mm), phalanx II-2 (85 mm), manual ungual II (~108 mm on curve), proximal metacarpal III, phalanx III-1, phalanx III-3, ilial fragments, proximal tibia, incomplete fibula, metatarsal II (250 mm), distal metatarsals III, phalanges III-1 (87 mm), metatarsal IV (~260 mm), phalanx IV-?, pedal phalanges, pedal ungual
Comments- At the Dinosaur Society conference 'British Dinosaurs - Their Lifes and Times' (1998), David Martill spoke about the new Isle of Wight theropod.
At the 1998 Symposium of Vertebrate Palaeontology and Comparative Anatomy, Hutt and Hutt described this theropod in "A new small theropod dinosaur from the Isle of Wight". Naish and Martill (2007) provided a new cranial and skeletal reconstruction that illustrate the known remains.
This is traditionally placed as a non-tyrannosaurid tyrannosauroid (e.g. Hutt et al., 2001), which has been found in subsequent cladistic analyses (Holtz, 2004; Senter, 2007). However, when additional relevent taxa and characters are added to Senter's study, Eotyrannus and several other traditional basal tyrannosauroids are instead resolved as slightly closer to birds than tyrannosaurids. This is based on several characters, such as the transversely compressed axial neural spine, low cervical neural spines, short cervical epipophyses, and elongate forelimb.
References- Kelly, 1998. Is this man our Indiana Jones? The Daily Mail, dated 10-7-1998.
Hutt, Naish, Martill, Barker and Newbery, 2001. A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research. 22, 247-242.
Hargreaves, Richard, 2001. "He's daddy of the dinosaurs," The News, May 10, 2001: 13.
Hutt, 2002. Mr Leng's dinosaur. The Geological Society of the Isle of Wight Newsletter. 2(6), 12–14 [December 2002].
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Naish and Martill, 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society. 164, 493-510.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Tanycolageus Carpenter, Miles and Cloward, 2005
“Tanycolagreus” Carpenter and Miles vide Anonymous, 2001
T. topwilsoni Carpenter, Miles and Cloward, 2005
“T. topwilsoni” Carpenter and Miles vide Anonymous, 2001
Middle-Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, Utah, 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, 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, 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)
Paratypes- (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)
(USNM 5737) pubes (Gilmore, 1920)
(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 Senter (2007). When Senter's analysis is updated with similar taxa and more tyrannosauroid characters, 'coelurids' move from basal Tyrannosauroidea to a position slightly closer to birds. The genus could be synonymous with Stokesosaurus, whose holotype and referred ilia, and referred braincase cannot be compared to Tanycolagreus.
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. Bull. U. S. Nat. Mus. CX 1-154, 36 pls., 78 text-figs.
Madsen, 1974. A new theropod dinosaur from the Upper Jurassic of Utah. Journal of Paleontology. 48, 27-31.
Miles, Carpenter and Cloward, 1998. A new skeleton of Coelurus fragilis from the Morrison Formation of Wyoming. JVP 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.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Calamosaurus Lydekker, 1891
= Calamospondylus Lydekker, 1889 (preoccupied Fox in Anonymous, 1866)
C. foxii (Lydekker, 1889) Lydekker, 1891
= Calamospondylus foxii Lydekker, 1889
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (BMNH R901) (~3.5 m, adult) anterior cervical vertebra (40 mm), anterior cervical centrum
Referred- ?(Dinosaur Farm Museum coll.) (Naish, DML 2002)
Diagnosis- (after Naish et al., 2001) anterior cervical transverse processes square in section.
Comments- Lydekker named this taxon Calamospondylus without realizing Fox had previously used the name for a different specimen. It cannot be compared with Calamospondylus or Aristosuchus, so synonymy with either is inappropriate. The tibia (BMNH R186) often referred to this species cannot come from the holotype individual due to size disparity and is not comparable to the holotype in any case (Naish et al., 2001). Naish (online, 2006) determined Calamosaurus is so similar to Dilong that it is a basal tyrannosauroid. This will be published in at least two papers that are in preparation. It is here moved to a position slightly closer to birds due to Dilong's changed placement following modification of Senter's (2007) matrix (see Dilong entry).
References- Lydekker, 1889. On a coelurid dinosaur from the Wealden. Geol. Mag. (ser. 3)6:119-121.
Lydekker, 1891. On certain ornithisaurian and dinosaurian remains. Q. J. Geol. Soc. London 47: 41-44.
Naish, Hutt and Martill, 2001. Saurichian dinosaurs 2: theropods: In: Dinosaurs of the Isle of Wight. edited by Martill and Naish. The Palaeontological Association, p. 242-309.
http://dml.cmnh.org/2002Apr/msg00529.html
http://dml.cmnh.org/2004Oct/msg00236.html
http://darrennaish.blogspot.com/2006/06/basal-tyrant-dinosaurs-and-my-pet.html
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Dilong Xu, Norell, Kuang, Wang, Zhao and Jia, 2004
D. paradoxus Xu, Norell, Kuang, Wang, Zhao and Jia, 2004
Late Valanginian-Hauterivian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V14243) (1.6 m; subadult) incomplete skull (166 mm), skeleton including mid cervical vertebra (24 mm), posterior dorsal vertebra (30 mm), twelve posterior caudal vertebrae (19-29 mm), scapula (69 mm), coracoid (48 mm), humerus (96 mm), ilium (138 mm), pubis (134 mm), femur (181 mm), tibia (203 mm), fibula (196 mm), pedal phalanx I-1, pedal ungual I, metatarsal II (112 mm), metatarsal III (117 mm), phalanx III-1, metatarsal IV (111 mm)
Paratypes- (IVPP V14242) (1.3 m; subadult) incomplete skull (132 mm), presacral vertebrae including mid cervical vertebra
(TNP01109) partial skull
Early Aptian, Early Cretaceous
Jianshangou Bed of Yixian Formation, Liaoning, China
?(IVPP V11579) (1.5 m; subadult) (skull ~158 mm) maxilla (96 mm), lacrimal, partial jugal, postorbital, squamosal, splenial (69 mm), surangular (95 mm), incomplete dentary, teeth, atlas (22 mm), axis (21 mm), six cervical vertebrae, three cervical ribs, first dorsal vertebra (26 mm), second dorsal vertebra (21 mm), third dorsal vertebra (21 mm), fourth dorsal vertebra (26 mm), fifth dorsal vertebra (26 mm), sixth dorsal vertebra (23 mm), seventh dorsal vertebra (22 mm), three dorsal vertebrae, five dorsal ribs, gastralia, first caudal vertebra (26 mm), second caudal vertebra (25 mm), third caudal vertebra (26 mm), fourth caudal vertebra (26 mm), fifth caudal vertebra (25 mm), sixth caudal vertebra (26 mm), seventh caudal vertebra (23 mm), eighth caudal vertebra (26 mm), ninth caudal vertebra (25 mm), ten caudal vertebrae, nine chevrons, scapula (89 mm), coracoid (~48 mm), three sternal ribs, distal humerus, distal radius, metacarpal I (21 mm), phalanx I-1, manual ungual I (39 mm), metacarpal II (43 mm), phalanx II-1 (28 mm), phalanx II-2 (43 mm), manual ungual II, metacarpal III (33 mm), phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III (24 mm), partial ilium, pubis, ischium, femur, tibiae, partial fibula, metatarsal I (28 mm), metatarsal II (109 mm), metatarsal III (118 mm), metatarsal IV (108 mm), metatarsal V (42 mm), pedal phalanges, feathers
Diagnosis- (modified from Xu et al., 2004) two large pneumatic recesses dorsal to the antorbital fossa on the maxilla; extremely long descending process of the squamosal extending close to the mandibular articulation of the quadrate; lateral projection of the basisphenoid anterior to the basal tuber; very deep, sub-circular interspinous ligamentous fossae on cervical vertebrae; robust scapula with a wide distal end (distal end twice as wide as the proximal scapular blade); hypertrophied coracoid (dorsoventral length about 70% of the scapular length).
Comments- IVPP V11579 may be a separate species. White (2009) illustrated and described a Dilong metatarsus, labeled IVPP V11579 in the text but V14242 in the figure. As V14242 is never claimed to contain appendicular elements in the original description and V15979's metatarsus is seemingly preserved as an impression distally, the figured metatarsus may actually be the holotype's.
Several cladistic analyses have found this to be a basal tyrannosauroid. Xu et al.'s (2004) analysis was a version of Currie et al.'s (2003) cranial-only tyrannosaurid analysis, with Dilong and outgroups added. It is not that useful, as the characters were designed to distinguish Allosaurus from tyrannosaurids, and resolve relationships within Tyrannosauridae, so do not even lead to a plausible outgroup topology (e.g. Citipati is the tyrannosauroid sister group). Xu et al. (2006) added Dilong and Guanlong to Rauhut's (2003) theropod analysis, which was followed by Benson (2008). Both Turner et al. (2007) and Senter (2007) added Dilong to the Theropod Working Group's matrix, though the latter contained more relevent tyrannosauroid characters. While Senter did find it to be a tyrannosauroid, Turner et al. resolved it slightly closer to birds. When I add the relevent taxa and all the characters used by Xu et al. (2006, 2004) to group Dilong with tyrannosauroids to Senter's (2007) matrix, Dilong ends up one node closer to birds than to tyrannosaurids (along with apparently related taxa Eotyrannus, Coelurus and Tanycolagreus). This is based on numerous characters- in lateral view, dorsal border of the antorbital fossa formed by the lacrimal and maxilla; round orbit; anterior cervical epipophyses absent or poorly developed, not extending past posterior rim of postzygopophyses; cervical neural spines dorsoventrally low; short laterotemporal fenestra; posterior dorsal neural spines height <1.5× length; humeral length more than half femoral length; manual phalanx I-1 longer than metacarpal II; feathers present. Constraining Dilong to be a tyrannosauroid results in trees 2 steps longer. It's still in the coelurid clade, but the latter is now the basalmost clade of tyrannosauroids. At least some of the above characters could be size-related, and Senter's matrix certainly has some character correlation problems. Two steps isn't very much, so I don't feel a tyrannosauroid identity can be ruled out. But at the same time, it's not as strongly supported as a blind tally of published analyses would suggest.
References- Currie, Hurum and Sabath, 2003. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontologica Polonica. 48(2), 227-234.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Xu, Norell, Kuang, Wang, Zhao and Jia, 2004. Basal tyrannosauroids from China and evidence for protofeathers in tyrannosauroids. Nature. 431, 680-684.
Xu, Clark, Forster, Norell, Erickson, Eberth, Jia and Zhao, 2006. A basal tyrannosauroid dinosaur from the Late Jurassic of China. Nature. 439, 715-718.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.
Turner, Pol, Clarke, Erickson and Norell, 2007. A basal dromaeosaurid and size evolution preceding avian flight. Science. 317, 1378-1381.
Benson, 2008. New information on Stokesosaurus, a tyrannosauroid (Dinosauria: Theropoda) from North America and the United Kingdom. Journal of Vertebrate Paleontology, 28(3), 732-750.
White, 2009. The subarctometatarsus: intermediate metatarsus architecture demonstrating the evolution of the arctometatarsus and advanced agility in theropod dinosaurs. Alcheringa. 33(1), 1-21.

Nuthetes Owen, 1854
N. destructor Owen, 1854
= Megalosaurus destructor (Owen, 1854) Steel, 1970
Middle Berriasian, Early Cretaceous
Cherty Freshwater Member of Lulworth Formation of Purbeck Limestone Group, England
Holotype- (DORCM G 913) (~2.3 m) (skull ~250 mm) partial dentary, teeth
Referred- (BMNH 48207) dentary fragments, teeth (Owen, 1878)
(BMNH 48208) several teeth (Owen, 1878)
(BMNH R 15870) maxillary tooth (15.5 mm) (Owen, 1879)
(BMNH R 15871) premaxilary tooth (7 mm) (Owen, 1879)
(BMNH R 15872) dentary tooth (5 mm) (Owen, 1879)
(BMNH R 15873) dentary tooth (5 mm) (Owen, 1879)
(BMNH R 15874) maxillary tooth (8.25 mm) (Owen, 1879)
(BMNH R 15875) maxillary tooth (8.5 mm) (Owen, 1879)
(BMNH R 15876) dentary tooth (1.5 mm) (Owen, 1879)
(BMNH R 15877) lateral tooth (5 mm) (Milner, 2002)
(BMNH R 15878) maxillary tooth (16 mm) (Milner, 2002)
Early Berriasian, Early Cretaceous
Marly Freshwater Member of Lulworth Formation of Purbeck Limestone Group, England
(CAMSM J13951) lateral tooth (Milner, 2002)
Berriasian, Early Cretaceous
Purbeck Limestone Group, England
? material (Delair, 1959; Benton and Spencer, 1995)
Description- Initial comparisons are to dromaeosaurids, as Milner (1999, 2002) proposed Nuthetes was a member of this clade. The holotype anterior dentary fragment suggests a cranial length of ~250 mm, and a total length of 2.3 meters, scaling from Deinonychus.
The dentary has an upper row of foramina, like dromaeosaurids and most other theropods, which are circular anteriorly (unlike Sinornithosaurus). A lower row seems to be absent, replaced by a groove that runs posterodorsally until fading around the seventh tooth. This seems unique within dromaeosaurids. Separate interdental plates are present, like Sinornithosaurus (in the anterior dentary at least) and Bambiraptor. The symphysis is represented by horizontal striations. The Mackelian groove is reduced to a poorly defined shallow depression terminating at the fifth tooth, with a foramen directly anterior. In Deinonychus and Saurornitholestes, the groove is shallow but well defined and the foramen located at the third or fourth tooth. Velociraptor's looks similar, but may end a bit more posteriorly, while Dromaeosaurus' ends more anteriorly, at the second tooth or so. Sinornithosaurus' extends to the third tooth, but is deep and narrow.
Teeth (numbering >9) are typically theropod in several respects; they are laterally compressed, recurved and serrated. The tip of some (all?) recurved teeth is located posterior to the alveolar edge, which is seen in dromaeosaurids. Milner reports the extent of serration on mesial carinae varies from the apical third to the whole crown, while the distal serrations are restricted to the apical half when present at all. I'm unaware of this occurring in any other theropod, as most lose their mesial serrations before their distal ones. And indeed one can see in plate 1 (6, 10) that distal serrations extend to the base of the crown in at least some teeth. Thus, I think Milner confused mesial and distal a few times in the text. What I suspect is really the case is that the mesial carinae are serrated for up to half their length (sometimes unserrated) while the distal carinae are completely serrated. This is similar to "velociraptorines" and Sinornithosaurus, but distinct from Dromaeosaurus (whch seemingly always has serrated mesial carinae) and Tsaagan, Microraptor and Cryptovolans (which lack mesial serrations). Heterodonty is observed, as 40% of the teeth are longer and more slender than the rest. This seems more developed than in Saurornitholestes. Constriction between the crown and root is comparable to Saurornitholestes, so is much less than in Microraptor. The DSDI varies from 1.14 to 1.55, which is almost identical to Sinornithosaurus (1.13-1.43), larger than dromaeosaurines (.81-1.18) and comparable to the low end of "velociraptorines" (1.19-2.33). Serrations are smaller than in most dromaeosaurids (4.5-8/mm compared to 2.5-2.8/mm in Dromaeosaurus, 2.4-2.6/mm in Utahraptor, 4-5/mm in Saurornitholestes, 3.2-3.6/mm in Deinonychus, 5/mm in Bambiraptor and Velociraptor; 3-3.6/mm in Achillobator) except for Sinornithosaurus and Microraptor, which have 7-14/mm and 8/mm respectively. Serrations have blunt rounded tips, unlike in "velociraptorine" and some anterior Sinornithosaurus teeth. Dromaeosaurines, Microraptor and most Sinornithosaurus teeth share this plesiomorphy with Nuthetes. Some serrations are asymmetrical, being almost hooked apically. Sinornithosaurus has only perpendicular serrations, while "velociraptorines'" and Microraptor's are hooked. Dromaeosaurus shows similar variation to Nuthetes.
A premaxillary tooth is said to be serrated only mesially, which is also seen in the first premaxillary tooth of Sinornithosaurus' holotype. However, Milner could have reversed the terms again and the distal carina may be the serrated one. In any case, Deinonychus, Dromaeosaurus and probably Utahraptor and Saurornitholestes differ in having both carinae serrated on premaxillary teeth. Most of Sinornithosaurus' anterior teeth are unserrated (as is reported for Microraptor), except for a few serrations on the first one, as mentioned above. The tooth is said to be D-shaped, but dromaeosaurines are often said to exhibit this condition as well, though it is quite different from the tyrannosauroid morphology. This tooth has a few strange characters as well- the enamel is striated, some serrations have a lingual midline ridge, and others have been combined in labial view.
Comments- Discovered in 1852, Owen originally (1854) described this taxon as a lizard, tentatively referring the tibia and fibula DORCM G 914 to Nuthetes as well (later referred to the atoposaurid crocodilian Theriosuchus by Seeley in 1893). Owen later (1879) identified Nuthetes as a crocodilian and referred additional teeth to the taxon, as well as dermal armor called granicones. Nuthetes was first identified as a dinosaur by Lydekker (1888), and as a theropod by Zittel (1911), who placed it in the Coeluridae. Swinton (1934) and Chakravarti (1935) referred it to the Megalosauridae. It was even synonymized with Megalosaurus itself by Romer (1956). Delair (1959) believed the granicones to be from a thyreophoran, and Galton (1986) referred them to Echinodon (now recognized as a heterodontosaurid). They have been recently reidentified as solemydid turtle limb and tail(?) scutes (Barrett et al., 2002). Though most often viewed as a lizard or juvenile megalosauroid or carnosaur last century, Nuthetes was reidentified as a juvenile dromaeosaurid by Milner (1999, 2002).
Milner assigns Nuthetes to the Dromaeosauridae based solely on the high DSDI, though she also compares the Mackelian groove and posterior extent of tooth tips to Deinonychus. She distinguishes it from "velociraptorines" by the absence of apically inclined serrations, despite the fact she earlier describes and illustrates these in the taxon. Although Milner says dromaeosaurines are distinguished from Nuthetes by their low DSDI and that Nuthetes compares most closely to "velociraptorines", she never actually assigns it to the Velociraptorinae. No comparison or mention is made to non-dromaeosaurid dromaeosaurs.
Other taxa besides "velociraptorines" have high DSDI's however. Within coelurosaurs, these include basal coelurid- or tyrannosauroid-like taxa (Guanlong, Dilong, Eotyrannus) and Richardoestesia.
That Richardoestesia is not discussed by Milner is confusing, as she lists it in her table of DSDI values as having a range almost identical to Nuthetes (1.06-1.53). Richardoestesia gilmorei has a Mackelian groove that is shallow like dromaeosaurids and an upper row of circular foramina. Further details are difficult to make out, though an anterior foramen may be present. It also has unfused interdental plates like Nuthetes. Notably, R. gilmorei exhibits similar heterodonty, with some teeth being quite elongate and others being shorter. And like dromaeosaurids, at least some teeth have tips extending posterior to their distal base. Nuthetes falls within the range of R. gilmorei for labiolingual thickness vs. FABL, height vs. FABL. Serration morphology is similar, with blunt rounded tips slightly hooked apically. Richardoestesia also has mesial serrations that vary in extent down the carina, from extending the complete crown edge to being absent. One of the characteristics of the genus is the minute distal serration size (5-12/mm), and this agrees with Nuthetes (4.5-8/mm). The tentatively identified premaxillary tooth of Richardoestesia is only serrated on one carina, with a cross section similar to Dromaeosaurus. Two important differences are that Nuthetes has interdenticle slits and has no significant basal constriction on lateral teeth. Also, Nuthetes shows none of the elongate straight teeth with convex distoapical margins known to exist in the anterior dentary of Richardoestesia, nor the elongate "stacked banana" serrations seen mesially in some teeth of that genus. Although R. isosceles is more similar in that it lacks basal constriction, it is far less recurved than Nuthetes, never as short as Nuthetes dentary teeth, has even smaller interdenticle slits, and never has rounded or pointed serrations.
Eotyrannus differs in having fused interdental plates and larger serrations (2.6/mm), though its Mackelian groove is shallow. Guanlong also has larger serrations (3.8/mm) and is said to have labiolingually thick lateral teeth, unlike Nuthetes. Dilong is more difficult to compare as it has been only preliminarily described. Its dentary tooth apices do extend posterior to the distal base, it has unfused interdental plates, its lateral teeth are highly compressed labiolingually and lack basal constriction, heterodont in the dentary at least, and have serrations which are rounded and sometimes asymmetrical. The serrations are separated by interdenticle slits as in Nuthetes. The premaxillary teeth are D-shaped, as described in Nuthetes, though whether Nuthetes' premaxillary tooth is truly D-shaped is uncertain as noted above. Unfortunately, the serration size, dentary foramina and medial dentary morphology are still undescribed. Nuthetes cannot be distinguished from Dilong from what is currently described of the latter, but this this may change as more material is discovered of Nuthetes and more is described on Dilong. It is provisionally placed by Dilong here.
References- Owen, 1854. On some fossil reptilian and mammalian remains from the Purbecks. Quarterly Journal of the Geological Society of London. 10, 420-433.
Owen, 1961. Monograph on the fossil Reptilia of the Wealden and Purbeck formations. Part V. Lacertilia (Nuthetes, etc.). [Purbeck]. The Palaeontological Society, London. 1858, 31-39.
Owen, 1878. On the fossils called 'granicones'; being a contribution to the histology of the exo-skeleton in 'Reptilia'. Journal of the Microscopical Society. 1, 233-236.
Owen, 1879. Monograph on the fossil Reptilia of the Wealden and Purbeck formations. Supplement no. IX. Crocodilia (Goniopholis, Brachydectes, Nannosuchus, Theriosuchus, and Nuthetes). The Palaeontographical Society. 1879, 1-19.
Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History). Part I. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum (Natural History), London. 1-309.
Seeley, 1893. On a reptilian tooth with two roots. Annals and Magazine of Natural History, s 6. 12, 227-230.
Zittel, 1911. Grundzüge der Paläontologie (Paläozoologie). II. Abteilung. Vertebrata [Fundamentals of Paleontology (Paleozoology). Section II. Vertebrata]. Druck und Verlag von R. Oldenbourg, München. 1-598.
Swinton, 1934. The Dinosaurs. George Allen & Unwin, London. 233 pp.
Chakravarti, 1935. Is Lametasaurus indicus an armored dinosaur?. The American Journal of Science, series 5. 30, 138-141.
Romer, 1956. Osteology of the Reptiles, University of Chicago Press. 1-772.
Delair, 1959. The Mesozoic reptiles of Dorset. Proceedings of the Dorset Natural History and Archaeology Society. 80, 52-90.
Galton, 1986. Herbivorous adaptations of Late Triassic and Early Jurassic dinosaurs. in Padian (ed.). The Beginning of the Age of Dinosaurs. Cambridge Univ. Press. pp. 203-221.
Milner, 1999. Purbeck Limestone meeting.
Milner, 2002. Theropod dinosaurs of the Purbeck Limestone Group, southern England. in Milner and Batten (eds). Life and Environment in Purbeck Times. Special Paper in Palaeontology. 68, 191-201.
Barrett, Clarke, Brinkman, Chapman and Ensom, 2002. Morphology, histology and identification of the 'granicones' from the Purbeck Limestone Formation (Lower Cretaceous: Berriasian) of Dorset, southern England. Cretaceous Research. 23, 279-295.
N. sp. (Mazin, Billo-Bruyat, Pouech and Hantzpergue, 2006)
Berriasian, Early Cretaceous
unnamed unit, Charente, France
Material- (CHEm03.537) tooth
References- Mazin, Billo-Bruyat, Pouech and Hantzpergue, 2006. The Purbeckian site of Cherves-de-Cognac (Beriasian, Early Cretaceous, southwest France): a continental ecosystem accumulated in an evaporitic littoral depositional environment. 9th International Symposium, Mesozoic Terrestrial Ecosystems and Biota, Manchester 2006, Abstracts and Proceedings volume. pp 84-88 and 169.
Pouech, Mazin and Billon-Bruyat, 2006. Microvertebrate biodiversity from Cherves-deCognac (Lower Cretaceous, Berriasian: Charente, France). 9th International Symposium, Mesozoic Terrestrial Ecosystems and Biota, Manchester 2006, Abstracts and Proceedings volume. p. 173.

"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. The constricted root is known in Compsognathus, Pelecanimimus, enigmosaurs, mononykines, troodontids, Richardoestesia, Microraptor and birds. Therefore, chances are pretty good this is a coelurosaur. 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. Pelecanimimus has yet to be described in detail, but has both anterior and posterior carinae unserrated. Therizinosaurs differ in having crowns that are less recurved, more elongate and labiolingually wider, with much larger posterior serrations (8-10 per tooth) and equally sized anterior serrations. Protarchaeopteryx lacks serrations. The highly elongate, needle-like teeth of Caudipteryx lack serrations too, so are very dissimilar. Mononykus has unserrated carinae, more elongate and less recurved crowns. Archaeopteryx has teeth that are completely unserrated, lack distal carinae and are much wider labiolingually. They are similarily proportioned and have unserrated mesial carinae. Some troodontid teeth lack mesial serrations, but have them distally, and are transversely compressed. The teeth of "Beelemodon" differ from troodontids in being less recurved, lacking hooked serrations and having comparatively smaller serrations (compared to 15-20 per tooth). Microraptor is similar in having crowns with unserrated mesial carinae and distal carinae with distally projecting serrations. It differs in having larger serrations in comparison to crown height (20-25 per tooth), longer blood grooves and a wider crown. Ornithurines (sensu Gauthier) have unserrated crowns without carinae and are very wide, so are similar to Archaeornithoides, but dissimilar to "Beelemodon". Dromaeosaurids also sometimes lack anterior serrations, are laterally compressed and have similar amounts of serrations (15-35), but are more recurved and lack basal constriction.
Therefore, the greatest resemblence is to Compsognathus, although a deinonychosaur might also be expected to evolve a similar tooth, judging by comparisons with Microraptor, Saurornitholestes and Morrison 'velociraptorine' teeth (Britt, 1991). Are there any theropods already known from the Morrison Formation that could have "Beelemodon" teeth? Although Morrison compsognathids are not known, both Coelurus and Ornitholestes are close phylogenetically and have poorly described or unknown teeth. Although reported Morrison 'velociraptorine' teeth lack constricted roots (Britt 1991), it is not inconceivable "Paleopteryx" had a mix of avian and dromaeosaur characters in its teeth, like Microraptor. The undescribed Morrison troodontid may have similar teeth, as its serrations are smaller than later troodontids, though the one tooth photographed so far has a more elongate crown. There are therefore several taxa to which "Beelemodon" could be reasonably referred. However, as it is currently impossible to chose one over another, they should be left separate. 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 coelurosaurian nomen nudum until further research is done.
References- Britt, 1991.
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.

"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 abelisaurids, at least some carnosaurs and most coelurosaurs (except those taxa which lack carinae). Abelisaurids and carnosaurs always have mesial serrations, often extending to the base of the crown. Some basal coelurosaurs and dromaeosaurids have anterior teeth which lack 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.

Phaedrolosaurus Dong, 1973
P. ilikensis Dong, 1973
Valanginian-Albian, Early Cretaceous
Lianmugin Formation of Tugulu Group, Xinjiang, China
Lectotype- (IVPP V 4024-1) (~7 m) tooth (31 mm)
Paratype- ?(IVPP V 4024-3) proximal femur
Diagnosis- Provisionally indeterminate.
Comments- Dong originally based this taxon on a tooth, partial hindlimb and proximal femur, all from different localities. Sues (1977) noted since the diagnosis is based on dental characters, the tooth should be the lectotype. Rauhut and Xu (2005) later made the hindlimb the holotype of a new species, Xinjiangovenator parvus, which they found to be a relative of Bagaraatan. Unfortunately, they neither mention the referred femur nor describe the tooth.
The tooth is about twice the size of Deinonychus, which would indicate a theropod about seven meters long.
According to Dong's figure, the tooth's outline is near identical to Deinonychus. It is compressed, recurved and serrated like most theropods. Serrations extend from the base to the tip of the distal carina, with eighteen serrations per five mm. The base of the mesial carina is smooth, but serrations are present starting half way up. It is said to be thicker than Deinonychus teeth.
The proximal femur is briefly mentioned, but not described or illustrated. It was found at a separate site, so should not be regarded as Phaedrolosaurus. Barsbold and Osmolska (1999) note that this femur has a wing-like anterior trochanter. This excludes it from Maniraptora, which have either finger-like or fused anterior trochanters. The lack of further information makes a more exact placement within Neotheropoda impossible.
Both Molnar (pers. comm. to Glut 1989; in Glut, 1997) and Sues (1977) state that Phaedrolosaurus appears dromaeosaurid. Barsbold and Osmolska (1999) say the wing-like lesser trochantor is distinctly non-dromaeosaurid. Besides these opinions, authors have generally just placed this genus in the Dromaeosauridae without question. While a lack of mesial serrations is congruent with a dromaeosaurid identity, various more basal coelurosaurs also exhibit that character. Pheadrolosaurus is probably either a basal coelurosaur or dromaeosaurid, but exact relations cannot be determined at this time. The tooth is said to be thicker than Deinonychus and Velociraptor, but thickness varies with position in the tooth row, and Bambiraptor and Atrociraptor also have generally thicker teeth than the former two genera.
References- Dong, 1973. [Dinosaurs from Wuerho]. Memoirs of the Institute of Vertebrate Paleontology and Paleoanthropolgy, Academia Sinica. 11, 45-52.
Sues, 1977. The skull of Velociraptor mongoliensis, a small Cretaceous theropod dinosaur from Mongolia. Paläontologische Zeitschrift. 51, 173-184.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076pp.
Barsbold and Osmólska, 1999. The skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica. 44, 189-219.
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.

Guanlong Xu, Clark, Forster, Norell, Erickson, Eberth, Jia and Zhao, 2006
G. wucaii Xu, Clark, Forster, Norell, Erickson, Eberth, Jia and Zhao, 2006
Oxfordian, Late Jurassic
Upper Shishugou Formation, Xinjiang, China
Holotype- (IVPP V14531) (~3 m; 7 year old adult) incomplete skeleton including incomplete skull, mandible, anterior dorsal vertebra, sacrum, proximal caudal vertebrae, mid caudal vertebrae, distal caudal vertebrae, humerus, radius, ulna, radiale, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, metacarpal IV, ilium, pubis, ischium, femur, tibia, fibula, astragalus, calcaneum, metatarsal I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, proximal phalanx III-3, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V
Paratype- (IVPP V14532) (<3 m; 6 year old subadult) skull, mandible, cervical vertebrae, cervical ribs, dorsal vertebrae, dorsal ribs, sacrum, proximal caudal vertebrae, scapula, coracoid, humerus, radius, ulna, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanges II-1, phalanx II-2, manual digit III, ilium, partial pubis, femur, tibia, fibula, metatarsals, pedal phalanges, pedal unguals
Diagnosis- (modified from Xu et al., 2006) deep and narrow groove along the anterior margin of the premaxilla; distinct opening on the maxilla close to the premaxilla–maxilla contact; complex, highly pneumatic nasal crest; low, rugose ridge along the midline of the frontals; dorsally flattened parietal with two parallel sagittal crests; transverse ridge within the supratemporal fossa; centropostzygapophyseal lamina on cervicodorsal vertebrae with its dorsal end expanding laterally; deep, longitudinal sulci on dorsal surfaces of the distal caudal vertebrae; ventral part of scapular blade with sub-equilateral triangular cross-section and thick posterior margin; metacarpal II with prominent medioventral and laterodorsal processes proximally; manual phalanx II-2 with prominent medioventral process proximally; femoral greater trochanter much narrower anteroposteriorly than the lesser trochanter; distinct fossa on posterodistal surfaces of astragalus and calcaneum; pedal phalanx II-1 with prominent paired ventral processes proximally.
Comments- Xu et al. (2006) added this and Dilong to Rauhut's (2003) matrix and found them to be basal tyrannosauroids. However, when examined in a version of Senter's (2007) matrix including all of Xu et al.'s data and many additional taxa and characters, it emerges closer to birds. Specifically it is related to Proceratosaurus, Mirischia and Sinocalliopteryx; further from birds than compsognathids, but closer than coelurids. Carr (2006) found Guanlong to be sister to Monolophosaurus within Carnosauria in an unpublished analysis. They were sister taxa based on the shape of the anterior maxillary process, tall pneumatic, fenestrate sagittal cranial crest, and obturator foramen in the ischium. While Carr also noted many characters which differ between the genera, he suggested these may be ontogenetic, as the taxa are from the same geological unit. Adding Guanlong to a theropod supermatrix (unpublished data- http://dml.cmnh.org/2008Mar/msg00195.html) results in a similar position to the modified version of Senter's matrix, but additionally shows that few steps are needed to place Guanlong in Tyrannosauroidea. Placing Guanlong either in Carnosauria or as a sister taxon to Monolophosaurus however, adds so many steps that it is improbable.
References- Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Carr, 2006. Is Guanlong a tyrannosauroid or a subadult Monolophosaurus? Journal of Vertebrate Paleontology. 26(3), 48A.
Xu, Clark, Forster, Norell, Erickson, Eberth, Jia and Zhao, 2006. A basal tyrannosauroid dinosaur from the Late Jurassic of China. Nature. 439, 715-718.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Proceratosaurus Huene, 1926
P. bradleyi (Woodward, 1910) Huene, 1926
= Megalosaurus bradleyi Woodward, 1910
Mid-Late Bathonian, Middle Jurassic
Great Oolite, England
Holotype- (BMNH R4860) ventral skull (260 mm), mandible, hyoids
Diagnosis- (after Rauhut and Milner, 2008) dorsal process of the premaxilla inclined slightly anterodorsally and nasal horn core overhanging the premaxillary internarial bar anteriorly; internarial bar of the premaxilla bifurcating posteriorly into a posteriorly directed ramus and a dorsally directed ramus; anterior end of the maxillary antorbital fossa placed considerably anterior and ventral to the promaxillary fenestra; anteriormost dentary tooth curved anteriorly and with the carinae oriented labiolingually.
Comments- Woodward (1902) named and described the species in 1910 as Megalosaurus bradleyi. It was placed in Megalosaurus as it has four premaxillary teeth, unlike Ceratosaurus - the other 'megalosaurid' mentioned by Woodward. Huene (1926) thought that this species was most closely related to Ceratosaurus because of the nasal crest and therefore separated it from Megalosaurus as Proceratosaurus bradleyi. He distinguished it from Ceratosaurus by the greater amount of premaxillary and maxillary teeth, as well as labially fluted premaxillary teeth. He distinguished it from 'megalosaurs' (Megalosaurus, Eustreptospondylus and Allosaurus) by the shape of the external naris, the shape and breadth of the dorsal maxillary process, the height of the antorbital fenestra and the lower mandibular joint. Huene (1926) later formalized the relationship between Proceratosaurus and Ceratosaurus by placing them both in the Ceratosauridae. Paul (1988) was the first to suggest Proceratosaurus is a coelurosaur, specifically related to Ornitholestes in Ornitholestiinae within the Allosauridae. This was based on their procumbant anterior dentary teeth, conical anterior teeth (both common in basal coelurosaurs), small anterior teeth and nasal horn (both not present in Ornitholestes). Holtz (2000) recovered Proceratosaurus as a coelurosaur more basal than Ornitholestes, though he later (Holtz, 2001; Holtz et al., 2004) recovered it as a sister taxon of Ornitholestes. Rauhut (2003) recovered it as the most basal coelurosaur however. Naish (online, 2006) recently noted his unpublished thesis finds Proceratosaurus to be a basal tyrannosauroid. Rauhut (DML, 1999) reported the right side of the skull has now been prepared. Rauhut and Milner (2008) restudied the skull and referred Proceratosaurus to Tyrannosauroidea based on- short premaxilla; well-developed jugal recess; steeply sloping basisphenoid; premaxillary teeth that are considerably smaller than the maxillary teeth; D-shaped anteriormost premaxillary teeth. While this is quite possible, especially if taxa such as Guanlong and Dilong are also basal tyrannosauroids, unpublished analyses including all the supposed tyrannosauroid characters of these taxa suggest they are slightly closer to birds instead.
References- Woodward, 1910. On a skull of Megalosaurus from the Great Oolite of Minchinhampton (Gloucestershire). Quarterly Journal of the Geological Society of London. 66(262), 111-115.
Huene, 1926. On several known and unknown reptiles of the order Saurischia from England and France. Annal and Magazine of Natural History. ser. 9. 17, 473-489.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 35-167.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus - and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River - Deinonychus, Velociraptor and Saurornitholestes. Hunteria. 2(4), 1-9.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York 1-464.
Rauhut, DML 1999. http://dml.cmnh.org/1999Oct/msg00057.html
Holtz. 2000. A new phylogeny of the carnivorous dinosaurs. GAIA. 15, 5-61.
Rauhut. 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria (second edition). University of California Press, Berkeley. 71-110.
Naish, online 2006. http://darrennaish.blogspot.com/2006/07/war-on-parasites-oviraptorosaurs-eye.html
Rauhut and Milner, 2008. Cranial anatomy and systematic position of the Middle Jurassic theropod dinosaur Proceratosaurus from England. Journal of Vertebrate Paleontology. 28(3), 130A.

Mirischia Naish, Martill and Frey, 2004
M. asymmetrica Naish, Martill and Frey, 2004
Albian, Early Cretaceous
Romualdo Member of Santana Formation, Brazil
Holotype- (SMNK 2349 PAL) (~2.1 m; subadult) posterior twelfth dorsal vertebra, thirteenth dorsal vertebra (26 mm), thefth dorsal rib, gastralia, first sacral vertebra, second sacral vertebra, anterior third sacral vertebra, partial ilia, pubes, 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 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. Naish (online, 2006) noted Mirischia is similar to tyrannosauroids in having an anteriorly concave pubic peduncle and referred the taxon to that clade. When analyzed in a version of Senter's (2007) matrix with additional taxa and characters, Mirischia is resolved as most closely related to Guanlong, Proceratosaurus and Sinocalliopteryx. These taxa are all placed slightly closer to birds than to tyrannosaurids, and have cranially concave pubic peduncles where known. A position for Mirischia in Tyrannosauroidea or Compsognathidae is quite possible though.
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, 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. 2004, 1-14.
http://darrennaish.blogspot.com/2006/06/basal-tyrant-dinosaurs-and-my-pet.html
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Sinocalliopteryx Ji, Ji, Lu and Yuan, 2007
S. gigas Ji, Ji, Lu and Yuan, 2007
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Beds of Yixian Formation, Liaoning, China
Holotype- (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, ulnae, radiale, ulnare, semilunate carpal, distal carpal III, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ilium, pubes, ischia, femora, tibiae, fibulae, astragalus, calcaneum, distal tarsal III, distal tarsal IV, pes, feathers, four gastroliths (15-20 mm)
Comments- Ji et al. (2007) described this as a compsognathid, but when included in an expanded version of Senter's (2007) analysis, it emerges most closely related to Guanlong, Proceratosaurus and Mirischia. All of these taxa are resolved as slightly closer to birds than to tyrannosauroids, though a placement within Tyrannosauroidea is possible.
Reference- 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, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

unnamed clade (Orkoraptor burkei + Aerosteon riocoloradensis)
Diagnosis- postorbital with a wide jugal process, and an orbital margin that is obtuse and devoid of a suborbital flange; anterior caudal vertebrae with pleurocoels.
Comments- Both of these taxa are large theropods from Late Cretaceous Argentina that seem to be basal coelurosaurs. Their postorbitals are a close match and both have proximal caudal pleurocoels, which are otherwise only known in some carcharodontosaurids and maniraptorans. Unfortunately, further comparison is difficult, as atlases and ribs are currently nondiagnostic, and Aerosteon's tibia has yet to be described. When included in a revised version of Senter's (2007) analysis, both clade together, so they are grouped together here as well.
References- Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Aerosteon Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2009
= "Aerosteon" Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2008
A. riocoloradensis Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2009
= "Aerosteon riocoloradensis" Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2008
Early Campanian, Late Cretaceous
Anacleto Formation of the Rio Colorado Subgroup, Argentina
Holotype- (MCNA-PV-3137) (~9-10 m; subadult) prefrontal (68 mm), postorbital (114 mm long), quadrate (163 mm), posterior pterygoid, prearticular, maxillary or dentary tooth (38 mm), 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)
Paratypes- ?(MCNA-PV-3138) metatarsal II
?(MCNA-PV-3139) tibia, incomplete fibula, astragalus, calcaneum
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; posteriormost dorsal vertebra with anterodorsally inclined neural spine; posteriormost dorsal vertebra with a pneumatic canal within the transverse process; medial gastral elements coossified with anterior and posterior flanges; furcula with median pneumatocoel.
Comments- Aerosteon was originally named in the online-only publication PLoS ONE, which does not satisfy the ICZN's requirement (Artcle 8.6) 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 in 2009, making Aerosteon a nomen nudum until that time.
This taxon was originally referred to Carcharodontosauridae by Alcober et al. (1998), and later as a carnosaur most similar to Allosaurus (Sereno et al., 2008). However, several characters are more similar to coelurosaurs. These include the high angle between anterior and ventral postorbital processes, quadrate foramen, possible anterodorsal concavity on the preacetabular process, concave anterior pubic peduncle margin, and tall astragalar ascending process. Furthermore, the postorbital is almost identical to that of Orkoraptor, a basal coelurosaur that shares the presence of caudal pleurocoels with Aerosteon. Including both in a revised version of Senter's (2007) analysis results in a sister group relationship within basal Coelurosauria.
References- Alcober, Sereno, Larsson, Martinez and Varricchio, 1998. A Late Cretaceous carcharodontosaurid (Theropoda: Allosauroidea) from Argentina. Journal of Vertebrate Paleontology. 18(3) 23A.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.
Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2008. Evidence for avian intrathoracic air sacs in a new predatory dinosaur from Argentina. PLoS ONE. 3(9), e3303.
Sereno, Martinez, Wilson, Varricchio, Alcober and Larsson, 2009. Evidence for avian intrathoracic air sacs in a new predatory dinosaur from Argentina. PLoS ONE. 3(9), e3303 [printed version].

Orkoraptor Novas, Ezcurra and Lecuona, 2008
O. burkei Novas, Ezcurra and Lecuona, 2008
Early Maastrichtian, Late Cretaceous
Pari Aike Formation, Santa Cruz, Argentina
Holotype- (MPM-Pv 3457) postorbital, quadratojugal (82 mm), coronoid(?), eight teeth, atlantal intercentrum, atlantal neurapophysis, eight fragmentary ribs, two mid caudal vertebrae (90 mm), three incomplete chevrons, proximal tibia (~700 mm)
Paratype- (MPM-Pv 3458) three teeth
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, but not described until 2008. Novas et al. (2008) found it to be a coelurosaur most probably related to compsognathids or dromaeosaurids in their cladistic analysis. The postorbital is almost identical to Aerosteon, a supposed carnosaur that also has proximal caudal pleurocoels. Including both in a revised version of Senter's (2007) analysis results in a sister group relationship within basal Coelurosauria.
References- Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.
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.

unnamed coelurosaur (Canudo, Filippi, Salgado, Garrido, Cerda, Garcia and Otero, 2009)
Late Coniacian-Early Santonian, Late Cretaceous
Plottier Formation of the Rio Neuquen Subgroup, Patagonia, 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 more basal theropods too.
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.

Nedcolbertia Kirkland, Britt, Whittle, Madsen and Burge, 1998
= "Nedcolbertia" Anonymous, 1996
N. justinhofmanni Kirkland, Britt, Whittle, Madsen and Burge 1998
= "Nedcolbertia whittlei" Anonymous, 1996
= "Nedcobertia justinhofmanni" Anonymous, 1998
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Holotype- (CEUM 5071) (1.5 m; juvenile) fifth sacral vertebra, eleven partial or complete caudal vertebrae, distal humerus, partial pubis, ischial fragments, proximal femora, distal femur, tibiae, proximal fibula, distal fibulae, partial astragali, calcanea, metatarsus, proximal and distal metatarsus, over fourteen pedal phalanges, six pedal unguals
Paratypes- (CEUM 5072) (3 m) dorsal vertebral fragments, several caudal vertebral fragments, four manual phalanges, partial manual ungual I, partial manual ungual II, ilial fragments, pubic fragments, distal pubis, proximal femora, proximal and distal tibia, proximal fibulae, astragalar fragments, proximal and distal metatarsals, several partial and complete pedal phalanges, five partial pedal unguals
(CEUM 5073) (3 m) few complete caudal centra, caudal central fragments, caudal neural arch fragments, several proximal chevrons, coracoid fragments, proximal humerus
Comments- This was originally announced in an abstract by Kirkland et al. (1995), then mentioned in a 1996 news report as "Nedcolbertia whittlei". That report stated it was intermediate between Ornitholestes and "Arkansaurus". Kirkland (1996) listed it as "small coelurosaurid cf. Ornitholestes n. gen." in an abstract. The species name was changed to "justinhofmanni" in January 1998 due to 6 year old Justin Hofmann winning a contest sponsored by Discover Card. It wasn't officially named N. justinhofmanni until October of that year.
References- Kirkland, Britt, Madsen and Burge, 1995. A small theropod from the basal Cedar Mountain Formation (Lower Cretaceous, Barremian) of Eastern Utah. Journal of Vertebrate Paleontology. 15(3), 39A.
Anonymous, 1996. Paleo Horizons. 2(2), 2.
Anonymous, 1998. Young Dino-Whiz Has New Dinosaur Named In His Honor. PRNewswire. Jan. 6.
Kirkland, 1996. Biogeography of western North America's Mid-Cretaceous Dinosaur Faunas: losing European ties and the first great Asian-North American interchange. Journal of Vertebrate Paleontology. 16(3), 45A.
Kirkland, Britt, Whittle, Madsen and Burge, 1998. A small coelurosaurian theropod from the Yellow Cat Member of the Cedar Mountain Formation (Lower Cretaceous, Barremian) of Eastern Utah. in Lucas, Kirkland and Estep (eds). Lower and Middle Cretaceous Ecosystems, New Mexico Museum of Natural History and Science Bulletin. 14, 239-248.

unnamed basal coelurosaur (Kurzanov, 1987)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren dabasu Formation, Inner Mongolia, China
Material- (PIN 2549-100) femur
Comments- Originally referred to Avimimidae by Kurzanov (1987), it was later noted to resemble Bagaraatan by Osmolska (1996).
References- Kurzanov, 1987. Avimimidae and the problem of the origin of birds [in Russian]. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 31, 1-95.
Osmolska, 1996. An unusual theropod dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. Acta Palaeontologica Polonica. 41, 1-38.

Bagaraatan Osmolska, 1996
B. ostromi Osmolska, 1996
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (ZPAL MgD-I/108) (~3 m; adult) (mandible ~230 mm) anterior dentary, posterior mandible, sacral neural spine, twenty-five caudal vertebrae (80, 65, 44, 37 mm), several chevrons, partial ilia, proximal pubis, proximal ischium, proximal and distal femur (~315 mm), tibia (365 mm, 380 mm with tarsus), fibula (~350 mm), astragalocalcaneum, pedal phalanx II-2 (37 mm), pedal phalanx IV-1 (34 mm)
Diagnosis- antarticular present; lateral longitudinal ridge present on proximal caudal prezygopophyses; two large fossae on the lateral postacetabular surface; anterior and greater trochanters with minimal separation; accessory trochanter; posterior trochanter present; tibiofibular crest powerfully developed; tibia broader mediolaterally than long anteroposteriorly in proximal view; tibia, fibula, astragalus and calcaneum fused.
Comments- This taxon was originally placed in the Avetheropoda by Osmolska (1996). She noted resemblences to a supposed avimimid from the Iren dabasu Formation described by Kurzanov (1987). Since then, it has been placed in various positions within Coelurosauria. Csiki and Grigorescu (1998) remarked on similarities between it, several maniraptoriformes (Elopteryx, Heptasteornis, Bradycneme) and an unnamed abelisaurid(?) distal femur (FGGUB R.351). Holtz (2000) placed it more basal than tyrannosauroids, maniraptoriformes and compsognathids, 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. By 2002, it was appearing as a basal tyrannosauroid or basal maniraptoran (sister to enigmosaurs + paravians) in Holtz's unpublished analyses (DML, 2002). 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 (Dilong was not yet known) based on the lateral surangular ridge, shortened retroarticular process and proximoventral pubic flange. Carr (2005) found it to be the sister taxon of NMMNH P-27469, 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. When coded into a modified version of Senter's (2007) analysis, with the maniraptoran characters from Rauhut and tyrannosauroid characters from Holtz added, Bagaraatan ends up as a basal coelurosaur of compsognathid-grade. It is tentatively placed there on this website.
References- Kurzanov, 1987. Avimimids and the problem of the origin of birds. Sovm. Sov.-Mong. Paleontol. Eksped. Trudy 31: 5-95. (In Russian).
Osmolska, 1996. An unusual theropod dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. Acta Palaeontologica Polonica. 41, 1-38.
Csiki and Grigorescu, 1998. Small theropods from the Late Cretaceous of the Hateg Basin (Western Romania) - an unexpected diversity at the top of the food chain. Oryctos. 1, 87-104.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. Gaia 15. 5-61.
Longrich, 2001. Secondarily flightless maniraptoran theropods? Journal of Vertebrate Paleontology. 21(3) 74A.
Coria, Chiappe and Dingus, 2002. A new close relative of Carnotaurus sastrei Bonaparte 1985 (Theropoda: Abelisauridae) from the Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 22, 460-465.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Carr, 2005. Phylogeny of Tyrannosauroidea (Dinosauria: Coelurosauria) with special reference to North American forms. Unpublished PhD dissertation. University of Toronto. 1170 pp.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

Compsognathidae Cope, 1875
Definition- (Compsognathus longipes <- Passer domesticus) (Holtz et al., 2004)
Other definition- (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
= Aristosuchia Seeley, 1901
= Compsognathinae Cope, 1875 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.

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), radiales, distal carpals I, distal carpals II, ulnares, 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, pubes, ischia, femur (163 mm), tibiae (183 mm), fibula, astragali, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsals II, 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, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanges IV-4, pedal unguals IV, metatarsal V, stomach contents
Paratype- (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, tibiae, distal tarsals, metatarsal I, metatarsal II, metatarsal III, metatarsal IV, metataral V
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 Theropod Working Group analysis, including that of Senter (2007).
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, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.

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, radiale (3 mm), distal carpal I (5.6 mm), distal carpal II (1.8 mm), metacarpals I (8.6 mm), phalanges I-1, manual ungual I, metacarpals II (17.1 mm), phalanges II-1, phalanx II-2, manual ungual II, metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3, manual unguals III, 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, 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, phalanges IV-1, phalanges IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V fragment, feathers, viscera, two eggs (37x26 mm) (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)
Diagnosis- (after Currie and Chen, 2001) first manual digit is longer than the humerus or the radius; powerful proximomedial flange on first metacarpal.
Comments- A lizard skeleton is preserved in the gut region of NIGP 127587.
A specimen described by Ji and Ji (1997), NGMC 2124, seems to be a different taxon. This was first suggested by Longrich (DML, 2000), who later wrote an abstract on it in 2002. This is agreed on by Ji et al. (2007) and Gishlick and Gauthier (2007), who label it Sinosauropteryx? sp..
References- Ji and Ji, 1996. On discovery of the earliest bird fossil in China and the origin of birds. Chinese Geology. 233, 30-33.
Ji and Ji, 1997. Advances in the study of the avian Sinosauropteryx prima. Chinese Geology. 242, 30-32.
Morell, 1997. The origin of birds: the dinosaur debate. Audubon Magazine, April, 36-45.
Chen, Dong and Zhen, 1998. An exceptionally well-preserved theropod dinosaur from the Yixian Formation of China. Nature. 391, 147-152.
Longrich, DML 2000. 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, 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.

Compsognathus Wagner, 1859
C. longipes Wagner, 1859
= Compsognathus corallestris Bidar, Demay and Thomel, 1972
Early Tithonian, Late Jurassic
Ober Solnhofen Plattenkalk, 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)
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), radiale, 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) opisthocoelous cervical vertebrae; no fourth trochanter on femur; metacarpal I less than one third as long as metacarpal II; ventral process at the posterior end of premaxillary body; 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.
Dames (1884) described three metapodials and a proximal phalanx (HMN coll.) from the Ober Solnhofen Plattenkalk, 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.
Wagner, 1861. Neue Beitrige zur Kenntis der urweltlichen Fauna des lithographischen Schiefers. V. Compsognathus longipes Wagn. Abh. Bayer. Akad. Wiss. 9: 30-38.
Dames, 1884. Uber Metatarsen eines Compsognathus - ahnlichen Reptils von Solnhofen. Sitz-Ber. Ges. Naturforsch.. 1884, 179-180.
Huene, 1925. Eine neue Rekonstrucktion von Compsognathus longipes. Clb. Mineral. Geol. u. Palaont. Jg. 1925, Abt. 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 Abbandlungen Bayerischen Staatssammlung Paldontol. historische Geol. (Munchen). 4, 73-118.
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).
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.
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.

Juravenator Gohlich and Chiappe, 2006
J. starki Gohlich and Chiappe, 2006
Late Kimmeridgian, Late Jurassic
Schamhaupton, Germany
Holotype- (JME Sch 200; Borsti) (~75-80 cm; juvenile) skull (82 mm), sclerotic ring, mandible (~77 mm), eight to ten cervical vertebrae, cervical ribs, thirteen dorsal vertebrae, dorsal ribs, gastralia, sacrum, forty-four caudal vertebrae, chevrons, scapula (42 mm), coracoids, furcula, humeri (27, 27.5 mm), radii (~19.3 mm), ulnae (20.5, 20.5 mm), metacarpal I (4.5 mm), phalanx I-1 (10.5 mm), manual ungual I (~10 mm), metacarpal II (11.5 mm), phalanx II-1 (8 mm), phalanx II-2 (10, 10 mm), manual ungual II (9, 10 mm), metacarpal III (9 mm), phalanx III-1 (4 mm), phalanx III-2 (4.5, 4.5 mm), phalanx III-3 (5.5 mm), manual ungual III (5.5, 7 mm), ilia (40 mm), ischia, femora (52 mm), tibiae (58.1, 58.1 mm), fibulae (55.3, 56 mm), astragali, calcaneum, metatarsal I (4.6, 4.5 mm), phalanx I-1 (5.8, 6 mm), pedal ungual I (6, 3.5 mm), metatarsal II (26.5 mm), phalanx II-1 (10.4 mm), phalanx II-2 (9, 8 mm), pedal ungual II (10.7, 11.5 mm), metatarsal III (34 mm), phalanx III-1 (11.9, 11.5 mm), phalanx III-2 (8.1, 8 mm), phalanx III-3 (7.4 mm), pedal ungual III (7.4, 6.6 mm), metatarsal IV (29.6, 29.8 mm), phalanx IV-1 (7.4, 7 mm), phalanx IV-2 (5.5, 6.5 mm), phalanx IV-3 (4.5 mm), phalanx IV-4 (4.2, 4 mm), pedal ungual IV (7.2, 5.8 mm), metatarsal V (8, 6.8 mm), scale impressions, caudal musculature impressions
Diagnosis- (modified from Gohlich and Chiappe, 2006) 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- This specimen was given the informal name of Borsti in 2001 news reports.
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.

unnamed coelurosaur (Longrich, 2002)
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Beds of Yixian Formation, Liaoning, China
Material- (NGMC 2124) incomplete skull, 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, pubis, ischia, femora, tibiae, pes, feathers
Comments- This was described as a new specimen of Sinosauropteryx prima by Ji and Ji (1997). Longrich (DML, 2000) noted it differed from Sinosauropteryx in several characters. He published an abstract in 2002 detailing his reasoning, proposing NGMC 2124 was a compsognathid/coelurid-grade coelurosaur, while Sinosauropteryx was a very basal coelurosaur or even a basal carnosaur. Gishlick and Gauthier (2007) refer to the specimen as Sinosauropteryx? sp. and figure the manus. Ji et al. (2007) also agree it does not belong in Sinosauropteryx.
There is a symmetrodont mandible preserved in the gut region of this specimen.
References- Ji and Ji, 1997. Advances in the study of the avian Sinosauropteryx prima. Chinese Geology. 242, 30-32.
Longrich, DML 2000. 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, 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.

Ornitholestes Osborn, 1903
O. hermanni Osborn, 1903
= Coelurus hermanni (Osborn, 1903) Hay, 1930
Middle Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Wyoming, US
Holotype- (AMNH 619) (2.08 m, 12.6 kg) skull (138 mm), mandibles, third cervical vertebra, fourth cervical vertebra, sixth cervical vertebra, first dorsal vertebra, second dorsal vertebra, third dorsal vertebra, seventh dorsal vertebra, eighth dorsal vertebra, ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal vertebra, thirteenth dorsal vertebra, first sacral vertebra, partial second sacral vertebra, partial third sacral centrum, fourth sacral vertebra, fifth sacral vertebra, first through seventh caudal vertebrae, two proximal caudal vertebrae, eighteen distal caudal vertebrae, twelve chevrons, humeri (124 mm), radius (84 mm), radial fragments, fragmentary ulna, metacarpal I, phalanx I-1, manual ungual I, partial phalanx II-2, manual ungual II, ilium (162 mm), incomplete pubess, ischia (152 mm), incomplete femur, proximal fibula, tarsal, pedal ungual I, metatarsal II (109 mm), phalanx II-1, pedal ungual II, metatarsal III (119 mm), phalanx III-1, phalanx III-2, pedal ungual III, metatarsals IV (113 mm), phalanx IV-1, phalanx IV-2, phalanx IV-4, pedal ungual IV
Comments- The holotype skeleton was originally described by Osborn (1903), who also referred a manus (AMNH 587) to the species. A partial skeleton was referred to Coelurus by Miles et al. (1998), also prompting them to refer AMNH 587 to that genus. 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. Ornitholestes has been questionably identified at Quarry 9 in Wyoming (Carrano and Velez-Juarbe, 2006) and Dry Mesa Quarry in Colorado (Britt, 1991), based on small elements that could belong to Coelurus, Tanycolagreus or other Morrison coelurosaurs as well.
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).
Makovicky (1995) described the vertebrae in detail, while Senter (2006) described the manus. Carpenter et al. (2005) described the postcranial skeleton, but the skull will be described by Norell in the future. Carpenter et al. incorrectly state only one humerus is known, do not mention the ulnar or left radial fragments, incorrectly list metacarpal II or III as being preserved, as well as fragments of two other metacarpals, and only mention one of the two manual unguals. The tibia is seemingly unpreserved, contra Paul's (1988) statement it is unusually short. In addition, a tarsal element is listed in the materials list, but not mentioned in the description, while two more pedal phalanges and two more pedal unguals are illustrated than are listed as being preserved.
References- Osborn, 1903. Ornitholestes hermanni, a new compsognathoid dinosaur from the Upper Jurassic. Bulletin of the American Museum of Natural History. 19, 459-464.
Osborn, 1916. Skeletal adaptations of Ornitholestes, Struthiomimus, Tyrannosaurus. Bulletin of the American Museum of Natural History. 35, 733-464.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. Bull. U. S. Nat. Mus. CX 1-154, 36 pls., 78 text-figs.
Hay, 1930. Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution of Washington. 390(II), 1-1074.
Ostrom, 1980. Coelurus and Ornitholestes: are they the same? In Jacobs, L. (ed.) Aspects of Vertebrate History. Flagstaff, Museum of Northern Arizona Press. 245-256.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus - and the sickleclaw theropods of the Cloverly, Djadokhta, and Judith River - Deinonychus, Velociraptor, and Saurornitholestes. Hunteria. 2(4), 1-9.
Britt, 1991. Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri. BYU Geology Studies. 37, 1-72.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Univ. Copenhagen, 311pp.
Miles, Carpenter and Cloward, 1998. A new skeleton of Coelurus fragilis from the Morrison Formation of Wyoming. JVP 18(3) 64A.
Carpenter, Miles, Ostrom and Cloward, 2005. Redescriptions 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. pp. 49-71.
Carrano and Velez-Juarbe, 2006. Paleoecology of the Quarry 9 vertebrate assemblage from Como Bluff, Wyoming (Morrison Formation, Late Jurassic). Palaeogeography, Palaeoclimatology, Palaeoecology. 234(2-4), 147-159.
Senter, 2006. Forelimb function in Ornitholestes hermanni Osborn (Dinosauria, Theropoda). Palaeontology. 49(5), 1029-1034.

Scipionyx Dal Sasso and Signore, 1998
= "Dromaeodaimon" Signore, 1995
S. samniticus Dal Sasso and Signore, 1998
= "Dromaeodaimon irene" Signore, 1995
Albian, Early Cretaceous
Pietraroia Plattenkalk Formation, Italy
Holotype- (Soprintendenza Archeologica coll.) (juvenile) skull, sclerotic ring, mandibles, hyoids, ten cervical vertebrae, nine cervical ribs, thirteen dorsal vertebrae, dorsal ribs and rib fragments, gastralia, first sacral vertebra, fourth sacral neural arch, fifth sacral vertebra, eight caudal vertebrae, ninth caudal neural arch fragment, fourth and fifth chevrons, sixth chevron fragment, scapulae, coracoids, furcula, humeri, radii, ulnae, radiales, distal carpals I, metacarpals I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1, phalanges II-2, manual unguals II (one proximal), metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3 (one distal), manual unguals III, horny manual claws, incomplete ilia, pubis, distal ischia, femora, proximal tibiae, proximal fibulae, tracheal fragment, pectoral musculature, liver remains, intestine, colon, caudifemoralis longus muscle, lateral caudal musculature
Diagnosis- (modified from Dal Sasso and Signore, 1998) accessory transverse postorbital ridge at fronto-parietal contact; compressed nature of the radiale and semilunate carpal.
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. It will be monographed in late 2009 or early 2010 by Dal Sasso and Maganuco (in prep.).
The sternal plate identified by Dal Sasso and Signore is actually the left proximal humerus.
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, Bn). Thesis, Dip. Paleont. Univ. Napoli "Federico II".
Dal Sasso and Signore, 1998. Exceptional soft tissue preservation in a theropod dinosaur from Italy. Nature. 392, 383-387.
Dal Sasso and Signore, 1998. Scipionyx samniticus (Saurischia, Theropoda): the first Italian dinosaur. Third European Workshop on Vertebrate Paleontology, Abstract: 23.
Dal Sasso and Signore, 1998. 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.
Dal Sasso and Maganuco, in prep.

Avepectora Paul, 2002
Definition- (majority of the distal edge of strongly anteriorly facing coracoids articulates with the anterior edge of a broad sternum at an angle of approximately 45-90 degrees from the midline as in Dromaeosaurus albertensis) (modified from Paul, 2002)

Metornithes Perle, Norell, Chiappe and Clark, 1993
Definition- (Mononykus olecranus + Passer domesticus) (modified from Chiappe, 1995)
= Protoavia Paul, 1988
= Bullatosauria Holtz, 1994
Definition- (Ornithomimus velox + Troodon formosus) (modified from Holtz, 1996)
= Maniraptoriformes Holtz, 1995
Definition- (Ornithomimus velox + Passer domesticus) (Maryanska et al., 2002; modified from Holtz, 1996)
Other definitions- (Ornithomimus edmontonicus + Passer domesticus) (Sereno, in press)
= Maniraptoriformes sensu Sereno, in press
Definition- (Ornithomimus edmontonicus + Passer domesticus)
Comments- Contrary to Sereno (in press), 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 Sereno's redefinitions of taxa eponymous with Ornithomimus would not be based on Ornithomimus. Sereno 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.
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, where alvarezsaurids are arctometatarsalians, this results in it being synonymous with Maniraptoriformes.
References- Chiappe, 1995. The first 85 million years of avian evolution. Nature. 378, 349-355.

Yaverlandia Galton, 1971
Y. bitholus Galton, 1971
Late Hauterivian-Early Barremian, Early Cretaceous
Wessex Formation of the Wealden Group, England
Holotype- (MIWG 1530) frontals, postorbital fragment, orbitosphenoid fragment
Diagnosis- frontals domed; frontals with pitted surface.
Comments- This specimen was originally considered to perhaps be referrable to Vectisaurus (Watson, 1930; Swinton, 1936), though Watson also noted resemblence to pachycephalosaurids. Galton (1971) named the taxon Yaverlandia bitholus, and assigned it to the Pachycephalosauridae, which was followed by most authors until the present. When entered into cladistic analyses of Pachycephalosauria, Yaverlandia was resolved at the base of the clade (Williamson and Carr, 2002) or as a basal pachycephalosaurid (Sereno, 2000). However, Hopson (1979) and Giffin (1989) doubted a pachycephalosaurian identity, based on the structure of the endocranium. Sullivan (2000, 2003) noted several characters incongruent with a pachycephalosaurian identity- frontals with broad orbital contact; parietals excluded from domes; supratemporal fenestrae contact frontals. Naish (2004) determined Yaverlandia is not a pachycephalosaur, though its identity remains elusive in his abstract. However, Sullivan (2006) noted that Naish believes it is a theropod, based on bilobed cerebral concavity; narrow olfactory tract; ventrally concave orbital margins; small, closely appressed olfactory bulbs. Indeed, the deep cerebral fossae and posteriorly expanded cerebrum suggest a coelurosaurian identity, while the narrow olfactory lobes are only known in maniraptoriforms. Among maniraptoriforms, Yaverlandia is unlike ornithomimids and dromaeosaurids in having ossified orbitosphenoids.
References- Watson, 1930. Proceedings of the Isle of Wight Natural History Society. 2, 60.
Swinton, 1936. The dinosaurs of the Isle of Wight. Proceedings of the Geologists' Association. 47, 204-220.
Galton, 1971. A primitive dome-headed dinosaur (Ornithischia; Pachycephalosauridae) from the Lower Cretaceous of England and the function of the dome of pachycephalosaurids. Journal of Paleontology. 45(1), 40-47.
Hopson, 1979. Paleoneurology. in Gans, Northcutt and Ulinski (eds.). Biology of the Reptilia (Neurology A). Academic Press, New York. 9, 39-146.
Giffin, 1989. Pachycephalosaur paleoneurology (Archosauria: Ornithischia). Journal of Vertebrate Paleontology. 9(1), 67-77.
Sereno, 2000. The fossil record, systematics and evolution of pachycephalosaurs and ceratopsians from Asia. In Benton, Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, New York. 480-516.
Sullivan, 2000. Prenocephale edmontonensis (Brown and Schlaikjer) new comb. and P. brevis (Lambe) new comb. (Dinosauria: Ornithischia: Pachycephalosauria) from the Upper Cretaceous of North America. New Mexico Museum of Natural History and Science Bulletin 17, Dinosaurs of New Mexico. 177-190.
Naish and Martill, 2001. Boneheaded and horned dinosaurs. pp. 133-146. in Martill and Naish (eds.). Dinosaurs of the Isle of Wight. The Palaeontological Association, London. 433 pp.
Williamson and Carr, 2002. A new genus of derived pachycephalosaurian from western North America. Journal of Vertebrate Paleontology. 22(4), 779-801.
Sullivan, 2003. Revision of the dinosaur Stegoceras Lambe (Ornithischia, Pachycephalosauridae). Journal of Vertebrate Paleontology. 23(1), 181-207.
Naish, 2004. So... what is Yaverlandia? The Annual Symposium of Vertebrate Palaeontology and Comparative Anatomy, Abstracts.
Sullivan, 2006. A taxonomic review of the Pachycephalosauridae (Dinosauria: Ornithischia). New Mexico Museum of Natural History and Science Bulletin. 35, 347-365.

undescribed possible maniraptoriform (Nessov, 1995)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- (N 464/12457) anterior sacrum
Comments- Nessov (1995) referred to this as cf. Gallimimus sp. or Oviraptorosauria.
References- Nessov, 1995. Dinosaurs of Northern Eurasia: new data about assemblages, ecology and paleobiogeography. Scientific Research Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia: 156 pp. + 14 pl. [in Russian with short English, German, and French abstracts].

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. alvarezsaurids, Avimimus, Microraptor).
References- Nessov, 1995. Dinosaurs of Northern Eurasia: new data about assemblages, ecology and paleobiogeography. Scientific Research Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia: 156 pp. + 14 pl. [in Russian with short English, German, and French abstracts].
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.

Arctometatarsalia

Maniraptora Gauthier, 1986
Definition- (Passer domesticus <- Ornithomimus velox) (Maryanska et al., 2002; modified from Padian et al., 1997; modified from Gauthier, 1986)
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)
(Passer domesticus <- Ornithomimus edmontonicus) (Sereno, in press)
= Therizinosauridae sensu Sereno, 1998
Definition- (Erlikosaurus andrewsi <- Ornithomimus velox) (modified)
= Dromavialae Ji and Ji, 2001
Comments- See the comments under Metornithes for why Sereno's (in press) definition of Maniraptora using Ornithomimus edmontonicus is inferior to Maryanska et al.'s (2002) using O. velox.
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 inavlid 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.

Chuniaoae Ji, Currie, Norell and Ji, 1998
= Maniraptora sensu Sereno, 1998
Definition- (Oviraptor philoceratops + 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 enigmosaurs and paravians, being a subset of Maniraptora potentially excluding taxa which are placed as basal maniraptorans in some studies (e.g. alvarezsaurids, Ornitholestes). Note Ji and Ji (2001) later proposed a similar name (Chuniaoia) on a cladogram for a group containing Protarchaeopteryx, but not birds.

unnamed maniraptoran (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. (1993) 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. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 2255-2272.

unnamed maniraptoran (Kessler and Jurcsak, 1984)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania
Material- (MTCO 1503) incomplete humerus (~50 mm)
Comments- This was discovered in 1978 and described by Kessler and Jurcsak (1984 and subsequent publications) as Archaeopteryx sp.. Benton et al. (1997) noted it could equally well be from a non-bird theropod. Indeed, the missing proximal end, deltopectoral crest and distal end leave very little anatomical detail. The general slenderness indicates maniraptoran affinities, as noasaurids and ornithomimosaurs differ in having a straight shaft. Yet it is virtually indistinguishable from not only Archaeopteryx, but also other taxa such as Microvenator and Bambiraptor.
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.
Jurcsak and Kessler, 1986. Evolutia avifaunei pe teritoriul Romanei. Partea I: Introducere (Evolution of the avifauna in the territory of Romania. Part I: Introduction). Crisia. 16, 577-615.
Kessler and 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.
Jurcsak and Kessler, 1987. Evolutia avifaunei pe teritoriul Romanei. Partea II: Morfologia speciilor fosile (Evolution of the avifaune in the territory of Romania. Part II: Morphology of fossil species). Crisia. 17, 583-609.
Jurcsak and Kessler, 1988. Evolutia avifaunei pe teritoriul Romanei. Partea III: Filogenie si sistematice (Evolution of the avifauna in the territory of Romania. Part III: Phylogeny and systematics). Crisia. 18, 647-688.
Jurcsak and Kessler, 1991. The Lower Cretaceous paleofauna from Cornet, Bihor County, Romania and its importance. Nymphaea. 21, 5-32.
Benton, Cook, Grigorescu, Popa and Tallodi, 1997. Dinosaurs and other tetrapods in an Early Cretaceous bauxite-filled fissure, northwestern Romania. Palaeogeography, Palaeoclimatology, Palaeoecology. 130(1-4), 275-292.

Maniraptora indet. (Nessov, 1984)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Materal- (TsNIGRI 45/11915) humeral shaft (~73 mm)
(TsNIGRI 48/11915) long bone shaft
(TsNIGRI 49/11915) long bone shaft
(TsNIGRI 50/11915) long bone shaft
Comments- This 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 Maniraptora 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 Maniraptora indet..
References- Nessov, 1984. [Upper Cretaceous pterosaurs and birds from Central Asia] Paleontologicheskii Zhurnal. 1, 47-57.
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.

unnamed maniraptoran (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), 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 maniraptoran.
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.

undescribed possible maniraptoran (Nessov, 1995)
Late Turonian-Coniacian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- (N 459/12457) manual ungual (?)I
Comments- This was listed in the text as being an oviraptorosaur, in which case it may be referrable to Caenagnathasia from the same formation. Nessov also listed the possibility of it being a bird pedal ungual in the figure caption however.
Reference- Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.

undescribed maniraptoran (Novas, Cladera and Puerta, 1996)
Cenomanian-Early Coniacian, Late Cretaceous
Rio Neuquen Subgroup, 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 perlvis is mesopubic, while Buitreraptor is also said to have a mesopubic pelvis.
Reference- Novas, Cladera and Puerta, 1996. New theropods from the Late Cretacoues of Patagonia. Journal of Vertebrate Paleontology. 16(3), 56A.

unnamed maniraptoran (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, 751-764.

unnamed maniraptoran (Novas, Borges Ribeiro and Souza Carvalho, 2005)
Late Maastrichtian, Late Cretaceous
Marilia Formation of the Bauru Group, Brazil
Material- (CP 659) manual ungual
References- Novas, Borges Ribeiro and Souza Carvalho, 2005. Maniraptoran theropod ungual from the Marý´lia Formation (Upper Cretaceous), Brazil. Revista del Museo Argentino Ciencias Naturales "Bernadino Rivadavia". 7, 31-36.
Candeiro, Martinelli, Avilla and Rich, 2006. Tetrapods from the Upper Cretaceous (Turonian-Maastrichtian) Bauru Group of Brazil: a reappraisal. Cretaceous Research.

undescribed maniraptoran (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 Maniraptora 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. Number 3557, 27 pp.

Bradycnemidae Harrison and Walker, 1975
Bradycneme Harrison and Walker, 1975
B. draculae Harrison and Walker, 1975
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Holotype- (BMNH 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). Martin (1983) suggested it was ornithomimid. Paul (1988) and Osmolska and Barsbold (1990) suggested 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 suggested it was a non-maniraptoran tetanurine. Naish and Dyke (2004) noted the craniocaudally compressed rectangular shape in distal view was similar to alvarezsaurids and maniraptorans, while the astragalar ascending process lacks the alvarezsaurid notched medial margin seen in Heptasteornis. They thus assigned Bradycneme to Maniraptora indet..
References- Lambrecht, 1929. Mesozoische und tertiare Vogelreste aus Siebenburgen. In Csiki (ed.). Xe Congres International de Zoologie. 1262-1275.
Lambrecht, 1933. Handbuch der Palaeornithologie. Berlin: 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, Biol. Sci. 23, 139-228.
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: New York 464 pp.
Osmolska and Barsbold, 1990. Troodontidae. 259-268. in Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press, Berkley, Los Angeles, Oxford. xvi-733.
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.

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 maniraptorans. 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. pp. 389-404. in Chiappe and Witmer, (eds.). Mesozoic Birds – Above the Heads of Dinosaurs. University of California Press, Berkeley, Los Angeles, London.

Praeornithes Rautian, 1978
Praeornithiformes Rautian, 1978
Praeornithidae Rautian, 1978
Praeornis Rautian, 1978
P. sharovi Rautian, 1978
Late Callovian-Kimmeridgian, Middle-Late Jurassic
Balabansai Formation, Kazakhstan
Holotype- (PIN 2585/32) feather?
Comments- Discovered in 1971, Sharov labeled the specimen Praeornis. Rautian later (1978) described this as the feather of a basal bird, naming it Praeornis sharovi. He assigned it its own subclass (Praeornithes), believing it to be more basal than Archaeopteryx due to the lack of barbules, and the medullary cavity in the barbs. Rautian also created the redundant order and family Praeornithiformes and Praeornithidae for the taxon (not to be confused with Kurochkin's 1995 Praeornithurae, which was erected for Protoavis). Bock (1986) stated it was more similar to a cycad leaf, and Burakova and Nessov (in Nessov, 1992) found that it was identical to the contemporaneous cycad species Cycadites saportae. They thus synonymized the species. Doludenko et al. (1990) found it was comparable to the leaves of Paracycas harrisii and similarly concluded it was a cycad. Feduccia (1996) and Wellnhofer (2004) concurred. One of the few recent references defending the feather identity is Glazunova et al. (1991), who used scanning electron microscopy to show it was not a plant and that the microstructure showed some resemblence to ratite feathers. Kurochkin (2001) accepts it as a feather, but says its particular identity remains unsolved. If it is a feather, it corresponds to stage 3A of Prum and Brush, which would perhaps indicate a basal maniraptoran. It is tentatively assigned such a placement here, until Glazunova et al.'s study is translated and evaluated.
References- Rautian, 1978. Unikal'noye pero ptitsy iz otlozheniy yurskogo ozera v khrebte Karatau. Paleontologicheskii Zhurnal. 4, 106-114.
Rautian, 1978. A unique bird feather from Jurassic lake deposits in the Karatau. Paleontological Journal. 4, 520-528.
Bock 1986. The arboreal origin of avian flight. in Padian (ed.). The Origin of Birds and the Evolution of Flight. California Academy of Sciences. Memoir 8, 57-72.
Doludenko, Sakulina and Ponomarenko, 1990. Geology of the unique deposits of the fauna and flora from the Late Jurassic of Aulie (Karatau, South Kazakhstan). Academy of Sciences of the USSR, Geological Institute. Moscow [in Russian].
Glazunova, Rautian and Filin, 1991. Praeornis sharovi: Bird feather or plant leaf? Materialy 10 Vsesoyuznoi Ornitologicheskoi Konferentzii, Vitebsk. Part 2(1), 149-150. [in Russian]
Nessov, 1992. Mesozoic and Paleogene birds of the USSR and their paleoenvironments. in Campbell (ed). Papers in Avian Paleontology Honoring Pierce Brodkorb. Natural History Museum of Los Angeles County Science Series. 36, 465-478.
Kurochkin, 1995. Synopsis of Mesozoic Birds and Early Evolution of Class Aves. Archaeopteryx. 13, 47-66.
Feduccia, 1996. The Origin and Evolution of Birds. Yale University Press, New Haven. 420 pp.
Kurochkin, 2001. Mesozoic birds of Mongolia and the former USSR. in Benton, Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. 533-559.
Wellnhofer, 2004. The plumage of Archaeopteryx: Feathers of a dinosaur. in Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons: Studies on the Transition from Dinosaurs to Birds. 282-300.

Shanyangosaurus Xue, Zhang and Bi, 1996
S. niupanggouensis Xue, Zhang and Bi, 1996
Middle-Late Maastrichtian, Late Cretaceous
Shanyang Formation, Shaanxi, China
Holotype- (NWUV 1111) (1.7 m) uncinate processes(?), partial sacrum, proximal scapula, humeri (116 mm), femur (258 mm), tibia (327 mm), metatarsal IV (137 mm), partial phalanx, pedal ungual
Diagnosis- long triangular cnemial crest on tibia projected forward with apex above posterior apexes of tibial condyles.
Description- The text states the bones are hollow.
The sacral remains consists of two complete fused vertebrae with the anterior section of a third. Both are 32 mm long and shown in ventral view.
The scapula is missing its distal end and is very poorly preserved. It would appear to have a low acromion and there is no forward projecting acromion process visible.
The humerus is 45% of the femoral length and also very poorly preserved, but appears to heve a more proximally placed deltopectoral crest than most other coelurosaurs (apex placed 22% down the shaft). The internal tuberosity is not very prominent.
The femur is in general very similar to "Ingenia", differing in a couple of ways. It's a bit more robust, has less of a neck below the head, which is separated from the greater trochanter by less of a groove. The lateral condyle is broken off, as are any remains of the lesser or accessory trochanters, if they existed. The text states that the fourth trochanter is absent and that the intercondylar groove is deep and wide. The head is elevated.
The tibia is 127% of femoral length and fairly well preserved. It's also very similar to "Ingenia"'s, but differs in a couple of ways as well. The shaft is bowed laterally and the cnemial crest is a different shape, being much longer and more prominent, with the anterodorsal edge almost perpendicular to the shaft. It's the one reason this taxon isn't a nomen dubium. There is a fibular crest and no depression for the ascending process can be observed.
The fourth metatarsal is 53% of femoral length, fairly robust, more so than many other coelurosaurs, and is almost certainly not arctometatarsalian because it's wider transversely than deep. It is narrower transversely when viewed proximally than Deinonychus.
There is also the distal section of a pedal(?) phalanx and another bone, which may be the proximal end of the other fourth metatarsal mentioned as it is roughly similar in proximal view.
The pedal ungual is not as deep as therizinosaurs, but deeper than Nedcolbertia. It's more curved than ornithimimids, but less so than dromaeosaurids.
Of special note is the statement "ribs with horizontal hooks" that is in the description and has fueled thoughts of uncinate processes. Note however, that no ribs are mentioned in the material list, nor are any shown in the plates.
Relationships- Shanyangosaurus was originally identified only to the level of Theropoda and there it has stayed in many people's minds. Unfortunately, the remains are very poorly preserved and described. Shanyangosaurus is obviously some type of theropod with an unfused metatarsus whose fourth metatarsal is wider than deep, a low acromion and an elevated femoral head. This narrows its placement down to basal coelurosaurs, oviraptorosaurs and paravians (excluding derived troodontids and avebrevicaudans).
Reference- Xue, Zhang, Bi, Yue and Chen, 1996. The development and environmental changes of the intermontane basins in the Eastern part of Qinling Mountains. Geological Publishing House, Beijing. ISBN 7-116-02125-6. 179 pages.

Unquillosauridae Powell, 1986
Unquillosaurus Powell, 1979
U. ceibalii Powell, 1979
Campanian, Late Cretaceous
Los Blanquitos Formation, Argentina
Holotype- (PVL 3670-11) ilial fragment, pubis (514 mm)
Description- Novas and Agnolin (2004) determined the supposedly diagnostic proximal sulcus noted by Powel (1979) doesn't exist, and is actually the pubic peduncle of the ilium broken and displaced. This allowed them to identify a ventrally concave pubic peduncle, and the angle between the anterior and ventral edges supports opisthopuby. What's normally seen as the acetabular surface of the pubis is the posterior part of a very long ilial peduncle, leaving a very tiny space for the acetabulum, which is said to resemble maniraptoriformes. These characters support placement in Maniraptora.
Comments- In 1997, Ford suggested on the DML that Unquillosaurus was similar to Unenlagia, which has gone so far as to prompt discussion of synonymy. However, Unquillosaurus is not synonymous with Unenlagia. First, I should note the pubis was described incorrectly, the medial side being lateral and vice versa. The supposed "lateral crest" is really the pubic apron, the "lateral" facets on the pubic boot are really for the pubic symphysis. Compared to Unenlagia- the pubis is more propubic; the ischial peduncle is longer; there was an obturator notch; the shaft is anteroposteriorly thicker distally; the pubic boot projects slightly anteriorly; there is a proximomedial sulcus; the ilial peduncle is less transversly expanded; the pelvic canal is narrower; the proximal shaft expands laterally; the distal end is expanded transversely, not compressed; there is a gap in the symphysis proximal to the pubic boot. Also keep in mind Unquillosaurus is from the Los Blanquitos Formation, while Unenlagia is from the Rio Neuquen Formation.
Novas and Agnolin keep Powell's orientation, remarking on the "prominent external longitudinal ridge" (=pubic apron?), odd medially convex pelvic canal margins (concave if reversed) and lack of an apron or medial symphysis. In addition, the medial end is described as "flattened and lacks marks for the articulation with the opposite bone". I see no reason to doubt the hypothesis they have the pubis reversed.
References- Powell, 1979. Sobre una asociacion de dinosaurios y otras evidencias de vertebrados del Cretacico superior de la region de La Candelaria, Prov. de Salta, Argentina. Ameghiniana. 16, 191-204.
Novas and Agnolin, 2004. Unquillosaurus ceibalii Powell, a giant maniraptoran (Dinosauria, Theropoda) from the Late Cretaceous of Argentina. Rev. Mus. Argentino Cienc. Nat.. 6(1), 61-66.