"Dinosaurus" Lessem and Glut, 1993
Comments- Lessem and Glut (1993) included an entry for a genus named Dinosaurus in their popular book "The Dinosaur Society's Dinosaur Encyclopedia". This was supposed to be a theropod from the Late Cretaceous of India known from fragmentary ribs and a caudal vertebra. However, personal communication with Glut (2000) indicates that the entry was an error by Lessem and does not reflect any real specimen.
Dinosaurus is the name of two other proposed genera though. Dinosaurus murchisoni (Fischer, 1845) Fischer, 1847 is a junior synonym of the ?estemmenosuchid synapsid Brithopus priscus. Contra some sources, Seeley (1895) did not try to name another genus Dinosaurus, but rather described a femur he referred to Dinosaurus that has been subsequently referred to the dinocephalan synapsid Phreatosuchus qualeni. "Dinosaurus gresslyi" was named without a description by Rutimeyer (1856a), so was a nomen nudum. It needed to be renamed due to Fischer's genus anyway, so was officially described as Gresslyosaurus ingens by Rutimeyer later that year (1856b). As "Dinosaurus gresslyi" was invalid, Olshevsky's (2000) claim the species name should still be gresslyi is incorrect. The species is a plateosaurid and is often placed in Plateosaurus itself, though this is a matter of personal preference.
References- Fischer, 1845. Beitrag zur naeheren Bestimmung des von Hrn. Wangenheim von Qualen abgebildeten und beschriebenen Saurier-Schaedels. Bulletin de la Societe Imperiale des Naturalistes de Moscou. 18, 540-543.
Fischer, 1847. Bemerkungen uber das Schadel-Fragment, welches Herr Major Wangenheim von Qualen in dem West-Ural entdeckt und der Gesellschaft zur Beurteilung vorgelegt hat. Bulletin de la Societe Imperiale des Naturalistes de Moscou. 20, 263-267.
Rutimeyer, 1856a. Dinosaurus gresslyi. Bibliotheque Universelle des Sciences Belles-Lettres et Arts, Geneve. September, 53.
Rutimeyer, 1856b. Reptilienknochen aus dem Keuper. Allgemeine Schweizerische Gesellschaft fur de Gesammten Naturwissenschaften. 41, 62-64.
Seeley, 1895. Researches on the structure, organization and classification of the fossil Reptilia. Part IX, section 1. On the Therosuchia. Philosophical Transactions of the Royal Society of London, series B. 185(21), 987-1018.
Lessem and Glut, 1993. The Dinosaur Society's Dinosaur Encyclopedia. New York: Random House.
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings. 3, 1-157.

Macrodontophion Zborzewski, 1834
Lochkovian-Eifelian, Early Devonian?
Dniester Series?, Podolia, Ukraine
Holotype
- shell (25 mm)
Comments- This genus was not given a species name when described by Zborzewski (1834). Macrodontophion's holotype was described as a snake tooth close to ophisaurs (actually anguid lizards) and pythons. It was also said to be similar to rhyncholites (cephalopod mandibles) and Beloptera (a spirulid cephalopod). Because Megalosaurus was mentioned in the same paragraph that the name first appears (the description proper is in the plate explanation), it was assigned to Megalosauridae by general reviews in the 1900's (e.g. Romer, 1956, 1976; Steel, 1970), and stated to be an indeterminate theropod by Weishampel (1990). Zborzewski tentatively referred it to the Jurassic, Molnar (1990) described its age as Late Jurassic or Cretaceous, and Weishampel assigned it to the Jurassic Nikolayevskaya Oblast based on personal comminication from Kurzanov. However, Nessov (1995) suggested its age was Early Devonian based on the geology of the region, Zborzewski's description of the locality as "ancient red sandstone" rich in lizard and snake teeth, comparable to the red strata in the Early Devonian Dniester Series which is rich in Porolepis teeth.
Molnar noted the insignificant distal taper was unlike theropod teeth and could not verify it was dinosaurian, though in pers. comm. to Olshevsky (2000) he stated it could be crocodilian or plesiosaurian, and in 1991 pers. comm. with Glut (1997) stated the abruptly flared base and lateral crests are similar to some shells. Sues (pers. comm. 1988 in Glut, 1997) also believed it was nondinosaurian. Nessov noted several characters which differ from theropod teeth- curvature perpendicular to plane of lateral crests (carinae of theropods); rounded apex; winding crest along inside edge of curve on long axis of tooth. Nessov believed that among Devonion fossils, Macrodontophion was most similar to the distal pectoral spine of an antiarch placoderm. While I agree the specimen is certainly not theropod, it does not resemble antiarch pectoral spines either. These spines are ironically like theropod teeth in being flattened parallel to their plane of curvature, lack the basal flare of Macrodontophion, and consist of numerous plates unlike the latter taxon. Even if most of Macrodontophion is an internal mold, the lateral crests seem to be made of one piece each and are too large to be single plates of an antiarch spine. Another possibility is the cornual process of an osteostracan or heterostracan, which are common in the Dniester Series (Dumbrava and Blieck, 2005; Voichyshyn, 2006). These are made of a single component, but still flattened in the plane of curvature. I believe an internal mold of a mollusk or annelid shell is most likely as an identification. Hyoliths, for instance, can have conical shells which are curved perpendicular to their axis of flattening, as in Macrodontophion. Yet hyoliths seem to lack the basal flare and have a more triangular section than the oval drawn for Macrodontophion. A large number of conical Palaeozoic shells are known which have controversial and problematic identifications, generally as hyoliths, scaphopods, nautiloids, cornulitids, coleolids or even worm tubes. Until Macrodontophion is examined by an expert in these taxa, it is probably best to keep it as Lophotrochozoa incertae sedis.
References- Zborzewski, 1834. Apercu des recherches physiques rationelles, sur les nouvelles curiosites Podoli-Colhyniennes, et sur leurs rapports geologiques aves les autres localites. Bulletin de la Societe Imperiale des Naturalistes de Moscou. 7, 224-254.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". in Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 306-317.
Weishampel, 1990. Dinosaurian distribution. in Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 63-139.
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.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076 pp.
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings. 3, 1-157.
Dumbrava and Blieck, 2005. Review of the pteraspidiform heterostracans (Vertebrata, Agnatha) from the Devonian of Podolia, Ukraine, in the Theodor Vascautanu collection, Bucharest, Romania. Acta Palaeontologica Romianiae. 5, 163-171.
Voichyshyn, 2006. New osteostracans from the Lower Devonian terrigenous deposits of Podolia, Ukraine. Acta Palaeontologica Polonica. 51(1), 131-142.

Kuphus Guettard, 1770
?= Succinodon Huene, 1941
K? putzeri (Huene, 1941) comb. nov.
= Succinodon putzeri Huene, 1941
Late Maastrichtian, Late Cretaceous-Early Danian, Paleocene
Nasilow greensand, Poland

Syntypes- (Museum der Universitat Tubingen coll?) lined tubes (lost) (Huene, 1941)
Referred- (ZPAL Mo. XV/1-25) lined tubes (Pozaryska and Pugaczewska, 1981)
Comments- Succinodon putzeri was originally described by Huene (1941) based on adjacent cylindrical fossils interpreted as titanosaurian (titanosaurine of Huene) sauropod teeth.
These were recognized by Pozaryska and Pugaczewska (1981) as being the calcareous lining of burrowing bivalve tubes, ascribed to the teredinid Kuphus sp.. As a named species cannot be sunk within unspecified 'sp.', this would technically be a new combination Kuphus putzeri. However, this combination has yet to be used, even online. While the authors could not locate Huene's type material in collections, they did describe numerous new specimens (ZPAL Mo. XV/1-25) from the same locality. The tubes are often crushed at one end, corresponding to supposed labiolingual compression noted by Huene, and the inner and outer tube walls are formed of different layers which could be confused with dentine and enamel. Huene described adjacent cylinders as being preserved in mandibular fragments, but it's uncertain whether this was merely an assumption based on the proximity of the "teeth" to each other, a misinterpretation of perhaps reworked sediment, or even fossilized wood as found at other localities.
Abdel-Gawad (1986) doubted their referral to Kuphus, instead assigning Succinodon to Teredinidae indet.. He first stated that Kuphus pallets (stemmed cup-like structures which seal the posterior siphon holes) and valves had never been found as fossils, but this doesn't mean they cannot be found and indeed have definitely been found since in fossil K. melitensis (Zammit Maempel, 1993). He also said that Kuphus differs because it's not found in wood unlike his specimens from the Dziurknow and Kazimierz localities, but the specimens from Nasilow (including Succinodon's types) are preserved in stone. Finally, Abdel-Gawad argued the morphology of the pallets differed, confusing the septa described by Pozaryska and Pugaczewska (perhaps homologous to the camerations described in Kuphus by Zammit Maempel) with pallets (which are as of yet unreported in the Polish material). Abdel-Gawad also stated the tubes are Cretaceous and not Paleocene, contra Pozaryska and Pugaczewska, but Machalski (pers. comm., 2015) confirms they are present in both the Danian layers and as reworked Cretaceous material.
Zammit Maempel refined Kuphus taxonomy to include only K. arenarius and K. melitensis, with other species which are based solely on tubes being indeterminate. The Nasilow material described by Pozaryska and Pugaczewska can still be compared to arenarius and melitensis however. Based on Zammit Maempel's diagnoses, putzeri resembles arenarius in being thinner-shelled, with weaker annulations, and melitensis in being less acute. Only the amount of taper can be determined from Huene's publication, and as his material is lost a neotype (preferrably including pallets and/or valves) would need to be established if putzeri were to be validated.
Another approach is that of Savrda and Smith (1996), who argued bivalve tubes should be covered by ichnotaxonomy instead. In this case the Nasilow material falls under Teredolites longissimus (Kelly and Bromley, 1984), using the revised diagnoses of Pickerill et al. (2003). This is primarily based on its elongation, and other Kuphus would be referred to this ichnospecies as well. As the tubes are secreted by and perminently occupied by the animals however, and consist of multiple calcifications with specialized structure, I think they should be considered body fossils instead of trace fossils.
References- Guettard, 1770. Memories sur differentes parties des Sciences et des Arts. 3, 139-143.
Huene, 1941. Ein obercretacischer Saurierrest aus Polen. Zentralblatt für Mineralogie, Geologie und Paläontologie, Abteilung B: Geologie und Paläontologie. 1941(3), 85-91.
Putzer, 1942. Die oberste Kreide bei Bochotnica a. d. mittleren Weichsel. Zentralblatt für Mineralogie, Geologie und Paläontologie B. 12, 361-377.
Pozaryska and Pugaczewska, 1981. Bivalve nature of Huene's dinosaur Succinodon. Acta Palaeontologica Polonica. 26(1), 27-34.
Kelly and Bromley, 1984. Ichnological nomenclature of clavate borings. Palaeontology. 27, 793-807.
Abdel-Gawad, 1986. Maastrichtian non-cephalopod mollusks (Scaphopoda, Gastropoda and Bivalvia) of the Middle Vistula Valley, Central Poland. Acta Geologica Polonica. 36(1-3), 69-224.
Zammit Maempel, 1993. Kuphus melitensis, a new teredinid bivalve from the Late Oligocene Lower Coralline Limestone of Malta. Contributions to Tertiary and Quaternary Geology. 30(3-4), 155-175.
Savrda and Smith, 1996. Behavioral implications of branching and tube-lining in Teredolites. Ichnos. 4, 191-198.
Pickerill, Donovan and Portell, 2003. Teredolites longissimus Kelly & Bromley from the Miocene Grand Bay Formation of Carriacou, the Grenadines, Lesser Antilles. Scripta Geologica. 125, 1-9.

Gnathostomata Zittel, 1879

Chondrichthyes Huxley, 1880

Elasmobranchii Bonaparte, 1838

Hybodontoidea Owen, 1846

Lonchidiidae Herman, 1977

Lonchidion Estes, 1964
?= Priscavolucris Gomez Pallerola, 1979
L? montsechi (Gomez Pallerola, 1979) new combination
= Priscavolucris montsechi Gomez Pallerola, 1979
= Lissodus palustris Gomez Pallerola, 1992
= Lissodus montsechi (Gomez Pallerola, 1979) Duffin, 2001
Late Berriasian-Early Barremian, Early Cretaceous
La Pedrera de Rubies Lithographic Limestones, Spain
Holotype
- (Gomez Pallerola coll.) (~350 mm) cranium (33 mm), Meckel cartilage, ceratobranchial, teeth, neural arches, scapulocoracoids (55 mm), propterygia, metapterygia, mesopterygia, radials, pectoral fins, scales
Comments- Gomez Pallerola (1979) originally identified this as the skull, forelimbs and feathers of a Jurassic bird, naming it Priscavolucris montsechi (misspelled Priscavulucris in the figure captions). He later (1982) briefly reidentified it as a shark (cf. Selechia), which was apparent from the original photos. The skull was correctly identified, but the supposed humerus is a scapulocoracoid, the supposed radius and ulna are the basal fin elements, the wing itself is the pectoral fin with the feathers being its rays. The supposed metatarsus and pedal digits are a ceratobranchial and perhaps parts of adjacent gill arch elements (hypobranchial, epibranchial), while the supposed ribs are probably neural arches. Gomez Pallerola later (1985, 1988) referred it to the genus Lonchidion. In 1992 he described it in detail as a new species- Lissodus palustris, as Lissodus had been recently synonymized with Lonchidion. Duffin (2001) noted that while it was obviously a shark, the correct species name is still montsechi. This made the correct name for the taxon Lissodus montsechi. Rees and Underwood (2002) revised Lissodus and kept it separate from Lonchidion. They were unable to examine L. montsechi, but felt it was possibly referrable to Lonchidion and could not be distinguished from the contemporary Lonchidion microselachos using information in the literature. However, while the name they used (Lissodus palustris) would be a junior synonym of L. microselachos if the taxa were identical, the species montsechi has priority over microselachos (from Estes and Sanchiz, 1982) if they are found to be synonyms. Neither the combination Lonchidion palustris nor the more correct L. montsechi have been published to my knowledge, though if correctly referred to the genus, this would be the only species known from skeletons.
References- Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Gomez Pallerola, 1979. Un ave y otras especies fosiles nuevas de la biofacies de Santa Maria de Meya (Lerida). Boletin Geologico y Minero. 90(4), 5-18.
Estes and Sanchiz, 1982. Early Cretaceous lower vertebrates from Galve (Teurel), Spain. Journal of Vertebrate Paleontology. 2(1), 21-39.
Gomez Pallerola, 1982. Nuevas aportaciones a la ictiofauna y a la flora del Neocomiense del Montsech de Rubies (Lerida). Boletin Geologico y Minero. 93(3), 199-213.
Gomez Pallerola, 1985. Nuevos hybodontidos del Cretacico Inferior de Santa Maria de Meya (Lerida). Boletin Geologico y Minero. 96(4), 372-380.
Gomez Pallerola, 1988. Nota sobre los peces elasmobranquios de las calizas litograficas del Cretacico Inferior del Montsec (Lerida). Boletin Geologico y Minero. 99(5), 748-756.
Gomez Pallerola, 1992. Nota sobre los tiburones hybodontos de las calizas litograficas del Cretacico Inferior del Montsec (Lerida). Boletin Geologico y Minero. 103(5), 783-813.
Duffin, 2001. Synopsis of the selachian genus Lissodus Brough, 1935. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 221(2), 145-218.
Rees and Underwood, 2002. The status of the shark genus Lissodus Brough, 1935, and the position of nominal Lissodus species within the Hybodontoidea (Selachii). Journal of Vertebrate Paleontology. 22(3), 471-479.

Osteichthyes Huxley, 1880

Sarcopterygii Romer, 1956
Definition- (Homo sapiens <- Perca fluviatilis) (Schultze and Arratia, 2004)

Actinistia Cope, 1872

Indocoelacanthus Jain, 1974
I? sp. (Jain, 1980)
= 'Kota bird' Anonymous?, 1979
Early Jurassic
Kota Formation, India

Material- (GSI coll.; Kota bird) partial skull (~166 mm), mandibles (~133 mm), cleithrum, scales
Comments- In 1979, the Indian popular press reported the discovery of a bird from the Early Jurassic Kota Formation of India. The find was attributed to Yadagiri, who claimed it was the oldest known bird and had a skull 150 mm long. Jain (1980) examined the specimen and identified it as a coelacanth skull. The supposed frontals are gulars, the supposed dentary is an infraorbital, the supposed radius and ulna (which are only a third of the skull's length, so would be far too short for a bird) are frontals, the supposed quadrate is an opercular, the supposed furcula is a cleithtrum and the supposed feathers are scales. Jain noted the skull is comparable to the Kota coelacanth Indocoelacanthus robustus (Jain, 1974) in gular, opercular and cleithral morphology, but differs in having two rows of mandibular denticulation. The absence of a skull roof made it difficult to compare in detail. He believed it could be a new species of Indocoelacanthus or a new genus of coelacanth. The specimen has not been examined since to my knowledge.
References- Jain, 1974. Indocoelacanthus robustus n. gen., n. sp. (Coelacanthidae, Lower Jurassic), the first fossil coelacanth from India. Journal of Paleontology. 48(1), 49-62.
Anonymous?, 1979. India Today. May 16-31. 4(10).
Jain, 1980. The continental Lower Jurassic fauna from the Kota Formation, India. In Jacobs (ed). Aspects of Vertebrate History. Museum of Northern Arizona Press, Flagstaff. 99-123.

Tetrapoda Goodrich, 1930
Definition- (Ascaphus truei + Homo sapiens) (Laurin, 2004)

Amphibia Linnaeus, 1758
Definition- (Ascaphus truei <- Homo sapiens) (Laurin, 2004)

Gymnophionia Rafinesque, 1814

Batrachia Latreille, 1800

Caudata Fischer von Waldheim, 1813

Salientia Laurenti, 1768

Stremmeia Nopcsa, 1930
S. scabra Nopcsa, 1930
Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania

Holotype- (MB coll.) ?tibiale (24.5 mm) fused to ?fibulare (27.3)
Comments- This specimen was discovered in 1910 associated with Giraffatitan's holotype, initially noted by Janensch (1914) as a bird carpometacarpus probably related to Archaeopteryx, then illustrated and described by Stremme (1919). The latter author could not determine what kind of reptile was represented, or if it was a metacarpus or metatarsus. Lambrecht (1933) noted that Stremme only mentioned Archaeopteryx in the context of differences between its metacarpus and the Tendaguru specimen, so dismissed Parkinson (1930) who stated Stremme considered them relatives. His own opinion was that the carpometacarpus resembled Rhea, so might indicate ratite relationships. Nopcsa (1930) meanwhile had identified the specimen as "carpals" of a pelobatid anuran, citing similarity to the tibiofibulare of Macropelobates (as noted by Stipanicic and Reig, 1957), and named it Stremmeia scabra. Among more recent authors, Hecht (1963) merely said that his reexamination of Stremmeia "showed that these bones are not frog remains" and Estes and Reig (1973) followed his interpretation. There has seemingly been no modern reevaluation of Stremmeia, or discussion of what its identity is if it is not a frog tibiofibulare.
Regardless of Hecht's comment, Stremmeia is more similar to e.g. Macropelobates' tibiofibulare than a maniraptoran carpometacarpus in the distinct proximal articular surfaces which form a flat outline instead of a single convex surface, and wide separation of the distal articular surfaces. This latter character is also unlike theropod metatarsals. However, the distal ends are more expanded transversely in anurans, and the distal articular surfaces seem to be simple instead of ginglymoid as in Stremmeia. Further differences from a coelurosaur manus include the large and quadrangular proximal surface of the shorter element (mcIII in most theropods) and sigmoid shape in side view. Whatever Stremmeia turns out to be, it is not theropod.
References- Janensch, 1914. Ubersicht uber die Wirbeltierfauna der Tendaguru-Schichten. Archiv fur Biontologie. 3, 81-110.
Stremme, 1919. Uber die durch Bandverknocherung hervorgerufene proximale Verschmelzung zweier Mittelhand - oder Mittelfussknochen eines Reptils. Wissenschaftliche Ergebnisse der Tendaguru-Expedition. Archiv fur Biontologie. 4, 143-144.
Nopcsa, 1930. Notes on Stegocephalia and Amphibia. Proceedings of the Zoological Society of London. 1930, 979-995.
Parkinson, 1930. The dinosaur in East Africa: An account of the giant reptile beds of Tendaguru, Tanganyika territory. H.F. & G. Witherby. 192 pp.
Lambrecht, 1933. Handbuch der Palaeornithologie. Gebruder Borntraeger. 1024 pp.
Stipanicic and Reig, 1957. El "Complejo Porfírico de la Patagonia extraandina" y su fauna de anuros. Acta Geologica Lilloana. 1, 185-297.
Hecht, 1963. A reevaluation of the early history of the frogs. Part II. Systematic Zoology. 12(1), 20-35.
Estes and Reig, 1973. The early fossil record of frogs: A review of the evidence. In Vial (ed.). Evolutionary biology of the anurans: Contemporary research on major problems. University of Missouri Press. 11-63.

Reptiliomorpha Save-Soderbergh, 1934
Definition- (Homo sapiens <- Ascaphus truei) (Laurin, 2004)

Lepospondyli Zittel, 1888

Amniota Haeckel, 1866
Definition- (Homo sapiens + Chelonia mydas + Sphenodon punctatus + Draco volans + Caiman crocodilus + Vultur gryphus) (Gauthier et al., 2004)
Other definitions- (Mammalia + Reptilia) (Gauthier et al., 1988)
(Homo sapiens + Iguana iguana) (Reisz, 2004)

Synapsida Osborn, 1903
Definition- (Dicynodon lacerticeps <- Crocodilus niloticus) (modified from Kischlat and Timm, 2006)
Other definitions- (Eothyris parkeyi + Varanops brevirostris + Homo sapiens) (modified from Laurin and Reiz, 1995)
(Homo sapiens <- Hylonomus lyelli, Procolophon trigoniceps, Petrolacosaurus kansensis) (modified from Reisz, 2004; modified from Gauthier et al., 1988)
(lower temporal arch as in Homo sapiens) (Rowe et al., 2004)

Eupelycosauria Kemp, 1982

Sphenacomorpha Ivankhnenko, 2003

Sphenacodontia Romer and Price, 1940

Sphenacodontoidea Marsh, 1878

Sphenacodontidae Marsh, 1878

Sphenacodontinae Marsh, 1878 vide Romer, 1936

Bathygnathus Leidy, 1853
= Dimetrodon Cope, 1878
B. borealis Leidy, 1853
= Dimetrodon borealis (Leidy, 1853) Brink, Maddin, Evans and Reisz, 2015
Artinskian, Early Permian
Orby Head Formation, Prince Edward Island, Canada
Holotype
- (ANSP 9524) partial premaxilla, septomaxilla, incomplete maxilla, anterior nasal
Diagnosis- (after Brink et al., 2014) large facial exposure of septomaxilla.
Comments- The holotype was discovered in 1845 and originally briefly described by Leidy (1853) as the mandible of a reptile, and later (1854, 1855) described in more depth as a lizard ("lacertian"). Cope (1868) suggested it was related to Dryptosaurus and provisionally placed it in his Goniopoda (=Theropoda) in 1870. Huene (1902) referred it to his theropod family Zanclodontidae, believing it might be synonymous with Teratosaurus. It was also commonly referred to the Amphisauridae (e.g. Marsh, 1882), and the Anchisauridae which replaced it (e.g. Marsh, 1885) when these were thought to be theropod groups. Owen (1876) correctly identified the specimen as an upper jaw and suggested a relationship to theriodonts (and Lycosaurus in particular) based on the presence of a canine tooth, reduced incisor and other characters. This was not confirmed until Case (1905) and Huene (1905) independently reidentified Bathygnathus as a "pelycosaurian" synapsid, Case believing it to be either Dimetrodon or Naosaurus (a junior synonym of Edaphosaurus that was mistakenly given a Dimetrodon skull). It was referred to Sphenacodontidae by Romer and Price (1940), and Sphenacodontinae in particular by Reisz (1986). Most recently, Brink et al. (2014; published as Brink et al., 2015) reexamined and redescribed Bathygnathus and found it to be sister to Dimetrodon grandis within Dimetrodon based on tooth roots which lack plicidentine, tooth roots equal in length to crowns, postcaniniform teeth with longitudinal distal sulci, and mesial and distal serrations. They note Bathygnathus has precedence over Dimetrodon, so petitioned the ICZN to retain the latter genus (Brink, 2015).
References- Leidy, 1853. [Remarks on Bathygnathus borealis]. Proceedings of the Academy of Natural Sciences of Philadelphia. 6, 404.
Leidy, 1854. On Bathygnathus borealis, an extinct saurian of the New Red Sandstone of Prince Edward Island. Proceedings of the Academy of Natural Sciences of Philadelphia, second series. 2(4), 327-330.
Leidy, 1855. On Bathygnathus borealis, an extinct saurian of the New Red Sandstone of Prince Edward's Island. American Journal of Science, series 2. 19, 444-446.
Cope, 1868. Remarks on extinct reptiles which approach birds. Proceedings of the Academy of Natural Sciences of Philadelphia. 19, 234-235
Cope, 1870. Synopsis of the extinct Batrachia, Reptilia and Aves of North America. Transactions of the American Philosophical Society. 14, 1-252.
Owen, 1876. Evidences of theriodonts in Permian deposits elsewhere than in South Africa. Quarterly Journal of the Geological Society of London. 32, 352-366.
Leidy, 1881. Remarks on Bathygnathus borealis. Journal of the Academy of Natural Sciences of Philadelphia, second series. 8(4), 449.
Marsh, 1882. Classification of the Dinosauria. American Journal of Science. 23, 81-86.
Marsh, 1885. On the classification and affinities of dinosaurian reptiles. Report of the British Association for the Advancement of Science. 1884, 763.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen (Neue Serie). Gustav Fischer Verlag, Jena. 6, 1-84.
Case, 1905. Bathygnathus borealis Leidy and the Permian of Prince Edward Island. Science. 22(550), 52-53.
Huene, 1905. Pelycosaurier im deutschen Muschelkalk. Neues Jahrbuch fur Minerologie, Geologie und Paleontologie. 20, 321-353.
Case, 1907. Revision of the Pelycosauria of North America. 176 pp.
Romer and Price, 1940. Review of the Pelycosauria. Geological Society of America Special Paper. 28, 538 pp.
Langston, 1963. Fossil vertebrates and the Late Palaeozoic Red Beds of Prince Edward Island. National Museum of Canada, Bulletin. 187, 36 pp.
Reisz, 1986. Pelycosauria. Handbuch der Paläoherpetologie. 17A, 102 pp.
Spalding, 1995. Bathygnathus, Canada's first "dinosaur". In Sarjeant (ed.). Vertebrate Fossils and the Evolution of Scientific Concepts. 245-254.
Brink, Evans, Maddin and Reisz, 2014. Phylogenetic assessment of Bathygnathus borealis, a derived species of Dimetrodon from Canada. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 97.
Brink, Maddin, Evans and Reisz, 2015. Re-evaluation of the historic Canadian fossil Bathygnathus borealis from the Early Permian of Prince Edward Island. Canadian Journal of Earth Sciences. 52(12), 1109-1120.
Brink, 2015. Dimetrodon Cope, 1878 (Synapsida, Sphenacodontidae): Proposed conservation by reversal of precedence with Bathygnathus Leidy, 1853. Bulletin of Zoological Nomenclature. 72(4), 297-299.

Sauropsida Goodrich, 1916
Definition- (Crocodylus niloticus <- Homo sapiens) (modified from Kischlat and Timm, 2006; modified from Gauthier et al., 1988)
Other definitions- (Mesosaurus tenuidens + Testudo graeca + Diapsida) (modified from Laurin and Reisz, 1995)
= Procolophonia Seeley, 1888
Definition- (Procolophon trigoniceps + Pareiasaurus serridens + Testudo graeca) (modified from Laurin and Reisz, 1995)
Other definitions- (Owenetta rubidgei + Barasaurus besairei + Procolophon trigoniceps + Lanthanosuchus watsoni + Sclerosaurus armatus + Pareiasaurus serridens + Testudo graeca) (modified from Lee, 1997)
= Testudinomorpha Laurin and Reisz, 1995
Definition- (Procolophon trigoniceps + Testudo graeca) (modified from Laurin and Reisz, 1995)

Parareptilia Olson, 1947
Definition- (Procolophon trigoniceps, Pareiasaurus serridens <- Homo sapiens, Iguana iguana) (Reisz, 2004)
Other definitions- (Testudo graeca <- Diapsida) (modified from Laurin and Reisz, 1995)
(Milleretta rubidgei, Procolophon trigoniceps <- Captorhinus aguti) (Tsuji and Muller, 2009)

Eureptilia Olson, 1947
Definition- (Captorhinus aguti, Petrolacosaurus kansensis <-Procolophon trigoniceps) (Tsuji and Muller, 2009)
Other definitions- (Diapsida <- Testudo graeca) (modified from Laurin and Reisz, 1995)

Romeriida Gauthier, Kluge and Rowe, 1988
Other definitions- (Sauria <- Anapsida) (Gauthier et al., 1988)
(Paleothyris acadiana + Diapsida) (Laurin and Reisz, 1995)

Diapsida Osborn, 1903
Definition- (Petrolacosaurus kansensis, Iguana iguana <- Captorhinus aguti, Procolophon trigoniceps, Paleothyris acadiana) (Reisz, 2004)
Other definitions- (Araeoscelis gracilis + Sauria) (Gauthier et al., 1988)
(Araeoscelis gracilis + Lepidosauria + Archosauria) (Laurin, 1991)
(Araeoscelis gracilis + Youngina capensis) (modified from Laurin and Reisz, 1995)
(Araeoscelis gracilis + Claudiosaurus germaini + Youngina capensis + Sauria) (modified from Braga and Rieppel, 1997)
(two temporal arches/fenestrae of Caiman crocodilus) (Gauthier et al., 2004)
(Araeoscelis gracilis + Crocodylus niloticus) (modified from Kischlat and Timm, 2006)

Eosuchia Broom, 1914
Definition- (Coelurosauravus elivensis + Apsisaurus witteri + Youngina capensis + Lepidosauria + Archosauria) (moidified from Laurin, 1991)
= Avicephala Senter, 2004
Definition- (Coelurosauravus elivensis, Megalancosaurus preonensis <- Neodiapsida) (modified from Senter, 2004)

Weigeltisauridae Kuhn, 1939
= Paleochameleontidae Romer, 1933? (at least Kuhn, 1939)
= Coelurosauravidae Currie, 1981
Other definitions- (Coelurosauravus elivensis + Wapitisaurus problematicus) (modified after Merck, 1997)

Coelurosauravus Piveteau, 1926
= Palaeochamaeleo Weigelt, 1930b (preoccupied De Stefano, 1904)
= Gracilisaurus Weigelt, 1930a
= Weigeltisaurus Kuhn, 1939
= Daedalosaurus Carroll, 1978
References- De Stefano, 1904. I Sauri del Quercy appartenenti alla collezione Rossignol. Atti della Societa italiana di Scienze naturali e del Museo civico di Storia naturale in Milano. 42(4), 382-418.
Piveteau, 1926. Paleontologie de Madagascar, XIII: Amphibiens et reptiles permiens. Annales de Paleontologie. 15, 53-128.
Weigelt, 1930a. Über die vermutliche Nahrung von Protorosaurus und über einen körperlich erhaltenen Fruchtstand von Archaeopodocarpus germanicus aut. Leopoldina (2nd series). 6, 269-280.
Weigelt, 1930b. Palaeochamaeleo jaekeli nov. gen. nov. sp., ein neuer Rhynchocephale aus dem Mansfelder Kupferschiefer. Leopoldina (2nd series). 6, 625-642.
Carroll, 1978. Permo-Triassic "lizards" from the Karoo System. Part II. A gliding reptile from the Upper Permian of Madagascar. Palaeontologia Africana. 21, 143-159.
C. elivensis Piveteau, 1926
= Daedalosaurus madagascariensis Carroll, 1978
Tatarian, Late Permian
Lower Sakamena Formation, Madagascar

Holotype- (IP 1908-11-21a) partial skull, at least thirteen presacral vertebrae, dorsal ribs, fragmentary patagial spurs, sacral vertebrae, at least eight caudal vertebrae, chevrons, pectoral girdles, humerus (29 mm), partial radius, partial ulna, several manual elements, manual ungual, femora (one distal; 31 mm), tibiae (one incomplete; 20 mm), fibula (~20 mm), astragalus, calcaneum, centrale, distal tarsal II, distal tarsal III, distal tarsal IV, distal tarsal V, metatarsal I, metatarsal II, phalanx II-1, phalanx II-2, metatarsal III, phalanx III-1, metatarsal IV, phalanx IV-1, proximal phalanx IV-2, metatarsal V, phalanx V-1, phalanx V-2, phalanx V-3, pedal ungual
Paratype- (IP 1908-11-22a) partial skull, twenty-three presacral vertebrae, cervical ribs, dorsal ribs, scapulocoracoid, clavicles, cleithrum, humeri (31.5 mm), radius, ulnae (one proximal; 23 mm), two metacarpals, two manual phalanges
Comments- Piveteau discovered the holotype in 1926 and described that year it as an ornithosuchid basal to coelurosaurs. It was soon recognized (e.g. Huene, 1930) to be non-dinosaurian, and is currently thought to be a non-reptilian neodiapsid. Obvious non-dinosaurian characters include the highly reduced lacrimal, large pineal foramen, palatine teeth, pleurodonty, intercentra, cleithrum, endo- and ectocondylar foramina in the distal humerus, pedal centrale, five distal tarsals, four phalanges on pedal digit V, etc.. Carroll (1978) described a new specimen as a new genus of coelurosauravid eosuchian- Daedalosaurus madagascariensis. It was later synonymized with Coelurosauravus by Evans (1982), as the supposed acrodont maxillary teeth of Coelurosauravus were actually squamosal horns, and the supposedly absent patagial spurs of Coelurosauravus were present but fragmentary (previously referred to Daedalosaurus by Carroll).
References- Piveteau, 1926. Paleontologie de Madagascar, XIII: Amphibiens et reptiles permiens. Annales de Paleontologie. 15, 53-128.
Huene, 1930. Palaeochamaeleo und Coelurosauravus. Centralblatt fur Geologie und Palaontologie. 1930, 440-441.
Carroll, 1978. Permo-Triassic "lizards" from the Karoo System. Part II. A gliding reptile from the Upper Permian of Madagascar. Palaeontologia Africana. 21, 143-159.
Evans, 1982. The gliding reptiles of the Upper Permian. Zoological Journal of the Linnean Society. 76(2), 97-123.
Evans and Haubold, 1987. A review of the Upper Permian genera Coelurosauravus, Weigeltisaurus and Gracilisaurus (Reptilia: Diapsida). Zoological Journal of the Linnean Society. 90(3), 275-303.
Bulanov and Sennikov, 2015. New data on the morphology of the Late Permian gliding reptile Coelurosauravus elivensis Piveteau. Paleontological Journal. 49(4), 413-423.
C. jaekeli (Weigelt, 1930) Evans and Haubold, 1987
= Gracilisaurus ottoi Weigelt, 1930a
= Palaeochamaeleo jaekeli Weigelt, 1930b
= Weigeltisaurus jaekeli (Weigelt, 1930) Kuhn, 1939a
Tatarian, Late Permian
Kupferschiefer, Germany
Tatarian, Late Permian
Marl Slate, England

Comments- The holotype of C. jaekeli was discovered around 1900 and first thought to be a reptile mixed with a caudal fin of Coelacanthus hassiae (Fritsch, 1901). Weigelt (1930b) described it as Palaeochamaeleo jaekeli, ignoring what he thought to be the caudal fin and referring it to Rhynchocephalia. He described another specimen that same year (1930a) as Gracilisaurus ottoi, also thought to contain rays of coelacanth fins but referred to Protorosauria. Huene (1930) noted the similarity between Coelurosauravus and Palaeochamaeleo. Kuhn (1939) replaced Palaeochameleo with Weigeltisaurus, as the former was preoccupied by what is currently a junior synonym of the agamid squamate Uromastyx. Schaumberg (1976) was the first to recognize the supposed fish fin rays as being patagial spurs. Evans and Haubold (1987) synonymized Weigeltisaurus and Gracilisaurus with Coelurosauravus, although Bulanov and Sennikov (2015) recently maintained generic separation based on several characters. As congenericity is subjective, separation will depend on whether it is needed to maintain monophyly of genera when other weigeltisaurids such as Rautiania are considered.
References- Fritsch, 1901. Exkursion in der Umgebung von Halle unter Fuhrung von Herrn K. v. Fritsch. Anlage. Bericht uber die in Verbindung mit der allgemeinen Versammlung in Halle a. Saale ausgefuhrten Exkursionen. Zeitschrift der Deutschen Geologischen Gesellschaft. 1901(1).
Huene, 1930. Palaeochamaeleo und Coelurosauravus. Centralblatt fur Geologie und Palaontologie. 1930, 440-441.
Weigelt, 1930a. Über die vermutliche Nahrung von Protorosaurus und über einen körperlich erhaltenen Fruchtstand von Archaeopodocarpus germanicus aut. Leopoldina (2nd series). 6, 269-280.
Weigelt, 1930b. Palaeochamaeleo jaekeli nov. gen. nov. sp., ein neuer Rhynchocephale aus dem Mansfelder Kupferschiefer. Leopoldina (2nd series). 6, 625-642.
Kuhn, 1939a. Protorosauria, Mesosauria. Fossilium Catalogus 1. Animalia. 85, 5-6.
Kuhn, 1939b. Schadelbau und systematische Stellung von Weigeltisaurus. Palaontologische Zeitschrift. 21, 163-167.
Schaumberg, 1976. Zwei Reptilneufunde (Weigeltisaurus Kuhn?, Lepidosauria?, Reptilia) aus dem Kuperschiefer von Richelsdorf (Perm, Hessen). Philippia. 3(1), 3-6.
Pettigrew, 1979. A gliding reptile from the Upper Permian of North East England. Nature. 281, 297-298.
Evans, 1982. The gliding reptiles of the Upper Permian. Zoological Journal of the Linnean Society. 76(2), 97-123.
Haubold and Schaumberg, 1985. Die Fossilien des Kupferschiefers. Neue Brehm-Bucherei. 333. 223 pp.
Schaumberg, 1986. Bemerkungen zu einem Neufund von Weigeltisaurus jaekeli (Weigelt) im nordhessischen Kupferschiefer. Palaontologische Zeitschrift. 60, 319-327.
Evans and Haubold, 1987. A review of the Upper Permian genera Coelurosauravus, Weigeltisaurus and Gracilisaurus (Reptilia: Diapsida). Zoological Journal of the Linnean Society. 90(3), 275-303.
Frey, Sues and Munk, 1997. Gliding mechanism in the Late Permian reptile Coelurosauravus. Science. 275(5305), 1450-1452.
Schaumberg, Unwin and Brandt, 2007. New information on the anatomy of the Late Permian gliding reptile Coelurosauravus. Palaontologische Zeitschrift. 81(2), 160-173.
Bulanov and Sennikov, 2015. New data on the morphology of the Late Permian gliding reptile Coelurosauravus elivensis Piveteau. Paleontological Journal. 49(4), 413-423.

Neodiapsida Benton, 1985
Definition- (Youngina capensis + Sauria) (Laurin, 1991)

Reptilia Linnaeus, 1758
Definition- (Testudo hermani + Sphenodon punctatus + Iguana iguana + Crocodylis niloticus) (Reisz, 2004)
Other definitions- (Testudines + Sauria) (Gauthier er al., 1988)
crown(Chelonia mydas + Sphenodon punctatus + Draco volans + Caiman crocodilus + Vultur gryphus) (Gauthier et al., 2004)
(Testudo hermani + Lacerta lacerta) (modified from Kischlat and Timm, 2006)
= Sauria MacCartney, 1802
Definition- crown(Sphenodon punctatus + Draco volans + Caiman crocodilus + Vultur gryphus) (modified from Gauthier et al., 1988; Gauthier et al., 2004)

unnamed possible reptile (Dames, 1884)
Early Tithonian, Late Jurassic
Ober Solnhofen Plattenkalk, Germany

Material- (HMN coll.) metapodial (54 mm), metapodial (60 mm), metapodial (68 mm), proximal phalanx (20 mm)
Comments- Dames (1884) described three metapodials and a proximal phalanx (HMN coll.), 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.
References- 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.
Ostrom, 1978. The osteology of Compsognathus longipes. Zitteliana Abbandlungen Bayerischen Staatssammlung Paldontol. historische Geol. (Munchen). 4, 73-118.

Lepidosauromorpha Benton, 1983
Definition- (Sphenodon punctatus + Squamata <- Crocodylia, Aves) (Gauthier et al., 1988)

Lepidosauria Dumerin and Bibron, 1839
Definition- crown(Sphenodon punctatus + Draco volans) (modified from Gauthier et al., 1988; Gauthier et al., 2004)
Other definitions- (Sphenodon punctatus + Scincus scincus) (modified from Kischlat and Timm, 2006)

Squamata Oppel, 1811
Definition- (Iguana iguana + Dibamus novaeguineae + Gekko gecko + Coluber constrictor) (modified from Kischlat and Timm, 2006)
Other definitions- (Iguana iguana + Autarchoglossa) (modified from Gauthier et al., 1988)
(Iguana iguana + Scleroglossa) (modified from Estes et al., 1988)

Anguimorpha Furbringer, 1900
Definition- (Anguis fragilis, Varanus varius <- Cordylus cordylus, Iguana iguana, Scincus scincus) (Conrad, 2008)

Anguiformes Conrad, 2006
Definition- (Anguis fragilis + Varanus varius) (Conrad, 2008)

unnamed anguiform (Grellet-Tinner, 2005)
Late Barremian, Early Cretaceous
Sao Khua Formation, Thailand
Materia
l- (SK1-1) (embryo) incomplete skull, incomplete mandibles, 57 centra, 81 neural arches, 63 ribs, 8 pectoral elements, 66 limb elements lacking hindlimbs, unidentified elements, incomplete egg (18x11 mm) (Buffetaut et al., 2005)
(SK1-2) (embryo) incomplete skull, mandibles, 7 teeth, 61 centra, 53 neural arches, 65 ribs, 9 pectoral elements, 6 pelvic elements, 72 limb elements, unidentified elements, incomplete egg (Buffetaut et al., 2005)
(SK1-3) (embryo) elements, partial egg (Buffetaut et al., 2005)
(SK1-4) (embryo) elements, partial egg (Buffetaut et al., 2005)
(SK1-5) fragmentary egg (Fernandez et al., 2015)
(SK1-6) (embryo) elements, egg (Fernandez et al., 2015)
(SK1-7) egg (Fernandez et al., 2015)
Diagnosis- (after Fernandez et al., 2015) anterior inferior alveolar foramen solely formed by splenial; broad palatine-groove articulation of prefrontal; autotomy planes present posterior to transverse process.
Comments- Grellet-Tinner (2005 thesis) and Buffetaut et al. (2005) described these as being from a non-ornithothoracine avialan closer to birds than troodontids, but Fernandez et al. (2012) determined they were actually from a squamate. Fernandez et al. (2015) fully redescribed the material based on CT scans to show the embryos are a new taxon of anguimorph, left unnamed due to the embryonic nature of the material. They noted characters congruent with a non-varanoid platynotan, but used a morphological phylogeny which differs from the molecular phylogeny on this site. Determining where the Thai taxon falls within a molecular phylogeny will require mapping morphological characters on to the squamate genetic topology.
References- Buffetaut, Grellet-Tinner, Suteethorn, Cuny, Tong, Košir, Cavin, Chitsing, Griffiths, Tabouelle and Le Loeuff, 2005. Minute theropod eggs and embryo from the Lower Cretaceous of Thailand and the dinosaur-bird transition. Naturwissenschaften. 92, 477-482.
Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case study of saurischian dinosaur relationships and avian evolution. PhD thesis, University of Southern California. 221 pp.
Grellet-Tinner, Chiappe, Norell and Bottjer, 2006. Dinosaur eggs and nesting behaviors: A paleobiological investigation. Palaegeography, Palaeoclimatology, Palaeoecology. 232, 294-321.
Fernandez, Buffetaut, Maire, Adrien, Suteethorn and Tafforeau, 2012. Phase contrast synchrotron microtomography: Improving noninvasive investigations of fossil embryos in ovo. Microscopy and Microanalysis. 18(1), 179-185.
Fernandez, Buffetaut, Suteethorn, Rage, Tafforeau and Kundrát, 2015. Evidence of egg diversity in squamate evolution from Cretaceous anguimorph embryos. PLoS ONE. 10(7), e0128610.

Mosasauridae Gervais, 1852

Russellosaurina Polcyn and Bell, 2005

Tylosaurinae Williston, 1897

Taniwhasaurus Hector, 1874
= "Yezosaurus" Obata and Muramoto vide Muramoto, 1977
= Lakumasaurus Novas, Fernandez, Gasparini, Lirio, Nunez and Puerta, 2002
Comments- There are three or four species in this genus, one of which (T. mikasaensis) was originally believed to be a tyrannosaurid and called Yezosaurus.
References- Hector, 1874. On the fossil Reptilia of New Zealand. Transactions of the New Zealand Institute. 6, 333-358.
Muramoto, 1977. [Road to Dinosaurs - Discovery of Yezosaurus mikasaensis]. North Garden, Inc. 114 pp.
Novas, Fernández, Gasparini, Lirio, Nunez and Puerta, 2002. Lakumasaurus antarcticus, n. gen. et sp., a new mosasaur (Reptilia, Squamata) from the Upper Cretaceous of Antarctica. Ameghiniana. 39, 245-249.
T. oweni Hector, 1874
= Platecarpus oweni (Hector, 1874) Lydekker, 1888
Late Campanian, Late Cretaceous
Conway Siltstone Formation, New Zealand

Comments- Although Caldwell et al. (2005) synonymized Tylosaurus haumuriensis (Hector, 1874) Welles and Gregg, 1971 with Taniwhasaurus oweni, Martin and Fernandez (2007) and Fernandez and Martin (2009) disagreed, the latter finding haumuriensis to group with other Tylosaurus in their phylogenetic analysis.
References- Hector, 1874. On the fossil Reptilia of New Zealand. Transactions of the New Zealand Institute. 6, 333-358.
Lydekker, 1888, Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum of Natural History, London. 309 pp.
Caldwell, Holmes, Bell and Wiffen, 2005. An unusual tylosaurine mosasaur from New Zealand: A new skull of Taniwhasaurus oweni (Lower Haumurian; Upper Cretaceous). Journal of Vertebrate Paleontology. 25, 393-401.
Martin and Fernández, 2007. The synonymy of the Late Cretaceous mosasaur (Squamata) genus Lakumasaurus from Antarctica with Taniwhasaurus from New Zealand and its bearing upon faunal similarity within the Weddellian Province. Geological Journal. 43, 203-211.
Fernanez and Martin, 2009. Description and phylogenetic relationships of Taniwhasaurus antarcticus (Mosasauridae, Tylosaurinae) from the upper Campanian (Cretaceous) of Antarctica. Cretaceous Research. 30(3), 717-726.
T? ivoensis (Persson, 1963) new combination
= Mosasaurus ivoensis Persson, 1963
= Tylosaurus ivoensis (Persson, 1963) Lindgren and Silverson, 2002
Early Campanian, Late Cretaceous
Belemnellocamax mammillatus zone of the Kristianstad Basin, Sweden

Comments- Caldwell et al. (2008) note this taxon's supposedly distinctive teeth are actually characteristic of Taniwhasaurus, though they did not formalize the nomenclature.
References- Persson, 1963. Studies on Mesozoic marine reptile faunas with particular regard to the Plesiosauria. Publications from the Institutes of Mineralogy, Paleontology, and Quaternary Geology, University of Lund, Sweden. 118, 1-15.
Lindgren and Siverson, 2002. Tylosaurus ivoensis: A giant mosasaur from the early Campanian of Sweden. Transactions of the Royal Society of Edinburgh: Earth Sciences. 93, 73-93.
Caldwell, Konishi, Obata and Muramoto, 2008. New species of Taniwhasaurus (Mosasauridae, Tylosaurinae) from the Upper Santonian-Lower Campanian (Upper Cretaceous) of Hokkaido, Japan. Journal of Vertebrate Paleontology. 28(2), 339-348.
T. antarcticus (Novas, Fernandez, Gasparini, Lirio, Nunez and Puerta, 2002) Martin and Fernandez, 2007
= Lakumasaurus antarcticus Novas, Fernandez, Gasparini, Lirio, Nunez and Puerta, 2002
Late Campanian, Late Cretaceous
Santa Marta Formation, Antarctica

References- Novas, Fernández, Gasparini, Lirio, Nunez and Puerta, 2002. Lakumasaurus antarcticus, n. gen. et sp., a new mosasaur (Reptilia, Squamata) from the Upper Cretaceous of Antarctica. Ameghiniana. 39, 245-249.
Martin and Fernández, 2007. The synonymy of the Late Cretaceous mosasaur (Squamata) genus Lakumasaurus from Antarctica with Taniwhasaurus from New Zealand and its bearing upon faunal similarity within the Weddellian Province. Geological Journal. 43, 203-211.
Fernanez and Martin, 2009. Description and phylogenetic relationships of Taniwhasaurus antarcticus (Mosasauridae, Tylosaurinae) from the upper Campanian (Cretaceous) of Antarctica. Cretaceous Research. 30(3), 717-726.
T. mikasaensis Caldwell, Konishi, Obata and Muramoto, 2008
= "Yezosaurus mikasaensis" Obata and Muramoto vide Muramoto, 1977
Late Santonian-Early Campanian, Late Cretaceous
Kashima Formation of the Upper Yezo Group, Japan
Holotype-
(MCM.M0009; specimen of "Yezosaurus mikasaensis") (~10.3 m) (skull ~1.5 m) partial snout, sclerotic plates, partial mandibles
Paratypes- ?(MCM.A1008) two dorsal vertebrae, two proximal caudal vertebrae, distal caudal vertebra
?(MCM.M10) two dorsal vertebrae
Late Santonian-Early Campanian, Late Cretaceous
Haborogawa Formation of the Upper Yezo Group, Japan
Paratype- ?(MCM.A600) jugal, postorbitofrontal, squamosal?, parietal, quadrate, ectopterygoid, coronoid
Comments- The original partial skull of "Yezosaurus mikasaensis" (originally part of Muramoto's private collection, but later designated as MCM.M0009) was discovered in 1976 and thought to be a tyrannosaurid. It has had a confusing citation history, as the West learned about from a two page photocopy in the late 1970's to early 1980's attributing it to Obata and Muramoto, 1977. This authorship and date was used by Olshevsky (1991) and Glut (1997), though neither includes a full citation, and an extensive literature search has failed to locate any such publication. The photocopy originated from a 1977 book by Muramoto and lacks a diagnosis (hence Olshevsky listing the taxon as a nomen nudum), though it's unknown if the book contains further description which could validate the genus. Muramoto wrote a one page article the following month (1977b), which again attributes the taxon to Obata and Muramoto, refers it to Tyrannosauridae, and illustrates it without a diagnosis. Thus a citation based on these references should be Obata and Muramoto vide Muramoto, 1977. Chure and McIntosh (1989) list the authorship merely as Obata, 1977, providing a citation for a short article about Muramoto's collection. This may actually be the correct reference for the name, depending on whether it refers to the taxon and when exactly it was published. Yomiuri (1977) also wrote a book later that year about "Yezosaurus" as a dinosaur (with Osborn's Tyrannosaurus skeletal on the cover), but again the technical content remains unknown. Chure (pers. comm. in Olshevsky, 1991) identified the remains as a mosasaur, which it was later redescribed as by Caldwell et al. (2008). Those authors named it as a new species of Taniwhasaurus, T. mikasaeansis. "Yezosaurus" is not mentioned in their paper due to its unofficial status in the Muramoto article (Konishi, pers. comm. 2010). If any of the 1977 publications include a description which follows ICZN rules, the authorship for mikasaensis will have to change, though "Yezosaurus" would still be a junior synonym of Taniwhasaurus. The holotype is clearly not a theropod, as it lacks an antorbital fenestra and maxillary ascending process (instead having a large prefrontal laterally exposed in that space), and further differs from tyrannosaurids and most other large theropods in having premaxilla-frontal contact, an extremely low snout, and uncompressed fluted teeth with expanded roots and no serrations. While spinosaurines are most similar among theropods, even they lack expanded roots and premaxilla-frontal contact.
References- Muramoto, 1977a. [Road to Dinosaurs - Discovery of Yezosaurus mikasaensis]. North Garden, Inc. 114 pp.
Muramoto, 1977b. Discovery of the fossil of a large reptile skull. Kaseki no Tomo. 16, 2.
Obata, 1977. [Home country collection room of the city of Mikasa and Muramoto's collection]. Natural Science and Museum. 44(1), 40-43.
Yomiuri, 1977. [Yezomikasa-Ryu, Testimomy by Dinosaurs: 100 Million Years of Japanese Archipelago]. Green Arrow Publishing.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076 pp.
Caldwell, Konishi, Obata and Muramoto, 2008. New species of Taniwhasaurus (Mosasauridae, Tylosaurinae) from the Upper Santonian-Lower Campanian (Upper Cretaceous) of Hokkaido, Japan. Journal of Vertebrate Paleontology. 28(2), 339-348.

Archosauromorpha Huene, 1946
Definition- (Archosauria <- Lepidosauria) (Gauthier et al., 1988)
Other definitions- (Protorosaurus speneri <- Lepidosauria) (modified from Dilkes, 1998)
(Rhynchosaurus articeps + Protorosaurus speneri + Caiman crocodilus) (Gauthier et al., 2004)
?= Basitheropoda Olshevsky, 1991
?= Theropodomorpha Olshevsky, 1991
Definition- (Megalancosaurus preonensis + Longisquama insignis + Megalosaurus bucklandii, - Passer domesticus) (modified from Olshevsky, 1991)
?= Archelosauria Crawford, Parham, Sellas, Faircloth, Glenn, Papenfuss, Henderson, Hansen and Simison, 2014
Definition- (Testudo graeca + Crocodylus niloticus) (Crawford et al., 2014)

unnamed Archosauromorpha (Kirby, 1993)
Rhaetian, Late Triassic
Owl Rock Member of the Chinle Formation, Arizona, US
Material
- (MNA V6729) distal tibia, limb shaft fragment
Comments- Kirby (1993) referred to cf. Staurikosauridae? in a faunal list. Spielmann et el. (2007) found two bone fragments catalogued as Staurikosauridae? indet. in the MNA collections, which they felt were too poorly preserved to place any more exactly than Archosauromorpha indet.. If the one is indeed a distal tibia as identified, it differs from Staurikosaurus and other herrerasaurids in being elongate in distal view instead of round.
References- Kirby, 1993. Relationships of Late Triassic basin evolution and faunal replacement in the southwestern United States: Perspectives from the upper part of the Chinle Formation in northern Arizona. In Lucas and Morales (eds.). The Nonmarine Triassic. New Mexico Museum of Natural History and Science Bulletin. 3, 233-242.
Spielmann, Lucas and Heckert, 2007. Tetrapod fauna of the Upper Triassic (Revueltian) Owl Rock Formation, Chinle Group, Arizona. In Lucas and Spielmann (eds.). The Global Triassic, New Mexico Museum of Natural History and Science Bulletin. 41, 371-383.

Euryapsida Colbert, 1945
Definition- (Ichthyosaurus communis + Plesiosaurus dolichodeirus) (modified from Merck, 1997)

Ichthyosauromorpha Motani, Jiang, Chen, Tintori, Rieppel, Ji and Huang, 2015
Definition- (Ichthyosaurus communis + Hupehsuchus nanchangensis) (Motani, Jiang, Chen, Tintori, Rieppel, Ji and Huang, 2015)

Ichthyosauriformes Motani, Jiang, Chen, Tintori, Rieppel, Ji and Huang, 2015
Definition- (Ichthyosaurus communis <- Hupehsuchus nanchangensis) (Motani, Jiang, Chen, Tintori, Rieppel, Ji and Huang, 2015)
Other definitions- (Ichthyosaurus communis + Mixosaurus cornalianus) (modified after Motani, 1997)

Grippiidae sensu Maisch and Matzke, 2000
Definition- (Grippia longirostris + Chaohusaurus geishanensis)

Ichthyopterygia Owen, 1840
Definition- (Ichthyosaurus communis + Utatsusaurus hataii + Parvivenator wapitiensis) (Motani, 1999)
= Eoichthyosauria Motani, 1999
Definition- (Grippia longirostris + Ichthyosaurus communis)

Ichthyosauria Blainville, 1835
Definition- (Ichthyosaurus communis <- Grippia longirostris) (Motani, 1999)
Other definitions- (Ichthyosaurus communis + Utatsusaurus hataii + Grippia longirostris) (modified after Merck, 1997)
(Thaisaurus chonglakmanii + Utatsusaurus hataii + Ophthalmosaurus icenicus) (Maisch and Matzke, 2000)

Longipinnati Huene, 1948
Definition- (Cymbospondylus petrinus + Ophthalmosaurus icenicus) (Maisch and Matzke, 2000)

Rachitrema Sauvage, 1883
R. pellati Sauvage, 1883
Rhaetian, Late Triassic
limestone at Conches-les-Mines, Autun, Saône-et-Loire, France

Lectotype (proposed)- (Pellat coll.) (~6.5 m) posterior dorsal neural arch (140 mm tall)
Paralectotypes- ?(Pellat coll.) scapula (220 mm)
(Pellat coll.) (Amniota incertae sedis) skull fragment, three dorsal rib fragments, proximal ?humerus, distal ?ilium or ?fibula, partial ?pelvic element
Comments- Sauvage (1883) described this material as a new taxon of dinosaur, though an oddly primitive one which he does not favorably compare to any other species. The specimen Sauvage based the name on is a neural arch he assigned to the caudal series, while he referred additional bones ("undoubtedly to the same animal") identified as a braincase fragment, at least three dorsal rib fragments, a scapula, proximal humerus, proximal radius and distal pubis. He notes the neural arch was unfused to the centrum, citing this as a primitive character. The structure of the prezygapophyses and ligament pits are compared to Actinodon (now recognized as a junior synonym of the temnospondyl Onchiodon), while the cranial fragment, scapula and humerus are compared to crocodilians and the latter two at least are said to differ from Megalosaurus and Cetiosaurus. Boulenger (1883) attempted to respell the name Racheotrema without comment. It has also sometimes been misspelled Rhachitrema, starting with Boettger (1884). Zittel (1890) stated it was a possible zanclodontid theropod (though insufficiently characterized), though he later (1895, 1902) stated it was a megalosaurid close to Zanclodon instead. Pocta (1905) also placed it in Megalosauridae, while Simroth (1907) placed it in Zanclodontidae. Nopsca (1901) assigned it to Anchisauridae, which was thought to be a theropod family at the time. Huene (1902) identified Rachitrema's type neural arch as that of an ichthyosaur and synonymized the genus with Shastasaurus without comment. Sauvage (1903) agreed Rachitrema was an ichthyosaur and synonymized it with the contemporaneous Ichthyosaurus? rheticus, since he believed that species to be more similar to Shastasaurus than the also contemporaneous I? carinatus. He regarded the scapula to be more similar to Toretocnemus (as Leptocheirus) than Ichthyosaurus, and the reidentified distal ilium (previously thought to be a distal radius) to be like Toretocnemus and Shastasaurus. The supposed proximal humerus and distal pubis could not be compared well. Merriam (1908) kept Rachitrema as a synonym of Ichthyosaurus? rheticus and believed the type neural arch was more primitive than Jurassic plesiosaurs. Huene (1908) synonymized it with Ichthyosaurus, crediting his 1902 paper despite the fact that work had it synonymized with Shastasaurus. Huene (1922) now listed Rachitrema as a junior synonym of Leptopterygius? rheticus (Leptopterygius has since been replaced by Leptonectes), though referring to not only the type neural arch but also a "supposed ischium". He later (1951) listed Rachitrema as a possible synonym of Merriamia (now a synonym of Toretocnemus), as did Romer (1976). Bardet and Cuny (1993) accepted some of the material as possibly ichthyosaurian, but assigned the rest to Reptilia indet.. McGowan and Motani (2003) consider Rachitrema to be probably dinosaurian without justification.
When evaluating Rachitrema, it should first be noted that no neural arches have been described for rheticus, so the two taxa cannot be shown to be synonymous. Thus rheticus can be ignored in the following discussion. In addition, there is no evidence the material described as Rachitrema belongs to one individual or taxon. It was all found isolated and by two different collectors. The neural arch is here proposed to be the lectotype, as the genus name refers to it and Sauvage states "The remarkable characters present in this neural arch make us think they indicate a dinosaurian of unknown type which we will indicate under the name Rachitrema." The neural arch is not dinosaurian, as it differs from dinosaur presacrals and proximal caudals in lacking transverse processes and having reduced zygapophyses which are placed near the midline. Nor is it from a distal caudal, which have reduced neural spines. The neural arch does strongly resemble ichthyosaurs in these characters however, so Huene's identification was correct. Within Ichthyopterygia, Rachitrema is outside Neoichthyosauria based on its divided postzygapophyses (Maisch and Matzke, 2000), as expected for a Triassic taxon. This indicates it is not synonymous with Ichthyosaurus or Leptonectes, though it still may be Shastosaurus or Toretocnemus. It is probably a member of Ichthyosauria, as no more basal ichthyosaurs are known from the Late Triassic. Pending further study, Rachitrema is considered Ichthyosauria incertae sedis.
The scapula has a short contact surface for the coracoid (~23% of scapular length), as in Cymbospondylus and non-longipinnatin taxa. The moderately sized anterior blade expansion is similar to Cymbospondylus, Shastasaurus and Shonisaurus, unlike the large flange of non-longipinnatins or the parvipelvians, Besanosaurus, Californosaurus, Mikadocephalus and Callawayia. Similarly, the low posterior blade expansion is unlike non-longipinnatins and neoichthyosaurs. The referred scapula is thus from a longipinnatin ichthyosaur that seems most similar to Cymbospondylus.
The supposed distal ilium could indeed belong to an ichthyosaur like Cymbospondylus, Toretocnemus or Californosaurus, or it could be a partial long bone shaft of a non-ichthyosaur (e.g. choristodere or dinosaur distal fibula) as originally described.
The other paratype fragments deserve detailed comparison to a wide variety of Triassic taxa, though the supposed humerus is certainly not an ichthyosaur humerus based on its elongation and the supposed distal pubis (which could also be a choristodere ventral ilium for instance) resembles parvipelvian pubes in rough shape but not Triassic ichthyosaurs.
References- Boulenger, 1883. Reptilia and Batrachia. in Rye (ed.). The Zoological Record for 1883. Record of Zoological Literature. 20, 24 pp.
Sauvage, 1883. Recherches sur les reptiles trouves dans l'etage Rhetien des environs d'Autun. Annales des Sciences Geologiques. 14(6, Article 3), 1-44.
Boettger, 1884. Bericht über die Leistungen in der Herpetologie während des Jahres 1883. Archiv Fur Naturgeschichte. 50(2), 379-434.
Zittel, 1890. Handbuch der Palaeontologie. Volume III. Vertebrata (Pisces, Amphibia, Reptilia, Aves). 900 pp.
Zittel, 1895. Grundzüge der Palaeontologie (Palaeozoologie). 971 pp.
Nopcsa, 1901. Synopsis und Abstammung der Dinosaurier. Földtani Közlöny. 31, 247-288.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen (Neue Serie). Gustav Fischer Verlag, Jena. 6, 1-84.
Zittel, 1902. Text-book of palaeontology, Volume 2. 283 pp.
Sauvage, 1903. Note sur les reptiles de letage Rhetien des environs d'Autun. Bulletin Societe d'Histoire Naturelle d'Autun (France). 16, 309-318.
Pocta, 1905. Rukovet Palaeozoologie. II. Cast: Vertebrata. 310 pp.
Simroth, 1907. Die Pendulationstheorie. Leipzig Konrad Grethlein's Verlag. 564 pp.
Huene, 1908. Die Dinosaurier der europäischen Triasformation mit Berücksichtiging der aussereuropäischen Vorkommnisse [The dinosaurs of the European Triassic Formation, with consideration of non-European occurrences]. Geologische und Paläontologische Abhandlungen Supplement-Band. 1, 419 pp.
Merriam, 1908. Triassic Ichthyosauria: With special reference to the American forms. 196 pp.
Huene, 1922. Die Ichthyosaurier des Lias und ihre Zusammenhänge. Jahresversammlung der palaeontologischen Gesellschaft. Verlag Bornträger, Berlin. 114 pp.
Huene, 1951. Eine neue Ichthyosaurier-Gattung der mitteleren Trias. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen. 94, 80-92.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Bardet and Cuny, 1993. Triassic reptile faunas from France. Paleontologia Lombarda. 2, 9-17.
McGowan and Motani, 2003. Handbook of Paleoherpetology: Ichthyopterygia, Part 8. 175 pp.

Sauropterygia Owen, 1860
Definition- (Placodus gigas + Plesiosaurus dolichodeirus) (modified from Rieppel, 1994)

Eosauropterygia Rieppel, 1994
Definition- (Pachypleurosaurus edwardsii + Corosaurus alcovensis + Plesiosaurus dolichodeirus) (modified from Rieppel, 1994)
Other definitions- (Nothosaurus mirabilis + Plesiosaurus dolichodeirus) (modified from Merck, 1997)
(Pachypleurosaurus edwardsii + Corosaurus alcovensis + Nothosaurus mirabilis + Plesiosaurus dolichodeirus) (modified from Rieppel, 1998)
(Pachypleurosaurus edwardsii + Nothosaurus mirabilis + Pistosaurus longaevus) (modified from Rieppel, 2000)

Eusauropterygia Tschanz, 1989
Definition- (Cymatosaurus fridericianus + Simosaurus gaillardoti + Lariosaurus balsami + Ceresiosaurus calcagnii + Nothosaurus mirabilis + Pistosaurus longaevus + Plesiosaurus dolichodeirus + Pliosaurus brachydeirus) (modified from Rieppel, 1994)
Other definitions- (Simosaurus gaillardoti + Nothosaurus mirabilis) (modified after Merck, 1997)
(Nothosaurus mirabilis + Pistosaurus longaevus) (modified from Rieppel, 2000)
= Pistosauroidea sensu Rieppel, 1998
Definition- (Corosaurus alcovensis + Pistosaurus longaevus) (modified)
= Pistosauria sensu Rieppel, 1998
Definition- (Cymatosaurus fridericianus + Pistosaurus longaevus + Plesiosaurus dolichodeirus + Pliosaurus brachydeirus) (modified)

Pistosauria Baur, 1890
Definition- (Augustasaurus hagdorni + Pistosaurus longaevus + Plesiosaurus dolichodeirus) (modified after Rieppel et al., 2002)
Other definitions- (Cymatosaurus fridericianus + Pistosaurus longaevus + Plesiosaurus dolichodeirus + Pliosaurus brachydeirus) (modified after Rieppel, 1998)

Plesiosauria Gray, 1825
Definition- (Plesiosaurus dolichodeirus + Pliosaurus brachydeirus) (modified from O'Keefe, 2001)
Other definitions- (Plesiosaurus dolichodeirus + Peloneustes philarchus) (Druickenmiller and Russell, 2008)

Pliosauroidea Seeley, 1874 vide Welles, 1943
Definition- (Pliosaurus brachydeirus <- Plesiosaurus dolichodeirus) (Ketchum and Benson, 2010)
Other definitions- (Thalassiodracon hawkinsi + Eurycleidus arcuatus + Attenborosaurus conybeari + Rhomaleosaurus cramptoni + Pliosaurus brachydeirus) (modified after O'Keefe, 2001)
(Rhomaleosaurus victor + Pliosaurus brachyspondylus) (Druickenmiller and Russell, 2008)

Pliosauridae Seeley, 1874
Definition- (Pliosaurus brachydeirus <- Rhomaleosaurus victor, Polycotylus latipinnis, Leptocleidus superstes) (Ketchum and Benson, 2010)
Other definitions- (Macroplata tenuiceps + Brachauchenius lucasi) (modified from O'Keefe, 2001)
(Simolestes vorax + Pliosaurus brachyspondylus) (Druickenmiller and Russell, 2008)

Thalassophonea Benson and Druckenmiller, 2013
Definition- (Pliosaurus brachydeirus <- Marmornectes candrewi) (Benson and Druckenmiller, 2013)

unnamed thalassophonean (Hahnel, 1988)
Middle Kimmeridgian, Late Jurassic
La Caja Formation, Mexico
Material
- (UANL-FCT-R2; The Monster of Aramberri) (~15 m) jaw fragment (lost), cranial fragments, nine cervical vertebrae, seven partial pectoral vertebrae (90-105 mm), rib fragments, gastralium?, axial material, incomplete scapula, incomplete coracoids, humerus, pelvis, femora (~1.2 m), epipodials
Comments- Hahnel (1988) initially referred this specimen to Theropoda based on the large size and carnivorous teeth. Buchy et al. (2003) reidentified it as Pliosauridae indet. based on the pectoral section and lost snout. Frey et al. (2006) announced the recovery of much of the rest of the specimen, preliminary results which are given in Buchy's (2007) thesis. Buchy retained it as Pliosauridae indet., which can be narrowed to Thalassophonea indet. given its late age. Once more of the specimen is prepared, it may be possible to assign further.
References- Hahnel, 1988. Hallazgo de restos de dinosaurio en Aramberri, N.L., Mexico. Actas de la. Facultad de Ciencias de la Tierra, U.A.N.L. 3, 245-250.
Buchy, Frey, Stinnesbeck and Lopez-Oliva, 2003. First occurrence of a gigantic pliosaurid plesiosaur in the Late Jurassic (Kimmeridgian) of Mexico. Bulletin de la Societe Geologique de France. 174(3), 271-278.
Frey, Stinnesbeck and Buchy, 2006. The Monster of Aramberri. German Research. 27(3), 4-7.
Buchy, 2007. Mesozoic marine reptiles from north-east Mexico: Description, systematics, assemblages and palaeobiogeography. PhD thesis, Universitat Karlsruhe. 89 pp.

Pliosaurus Owen, 1841 sensu Owen, 1842
?= Spondylosaurus Fischer de Waldheim, 1845
?= "Tapinosaurus" Lennier, 1887
= Stretosaurus Tarlo, 1959
= Strongylokrotaphus Novozhilov, 1964
Comments- This was originally erected as a subgenus of Plesiosaurus, spelled Pleiosaurus (Owen, 1841). As Benson et al. (2013) state, ICZN Article 33.3.1 indicates Pliosaurus is an incorrect subsequent spelling but should be retained as it has been "in prevailing usage and is attributed to the publication of the original spelling." Owen (1841) raised it to genus level. See Knutsen (2012) and Benson et al. (2013) for differing views on species validity.
References- Owen, 1841. Odontography; or a treatise on the comparative anatomy of the teeth, I Part 11. Dental system of reptiles. Hippolyte Bailliere, London. 179-295.
Owen, 1842. Report on British fossil reptiles. Part II. Report of the Eleventh Meeting of the British Association for the Advancement of Science. 60-204.
Fischer de Waldheim, 1845. Notice sur le Spondylosaurus genre de saurien fossile de l'oolithe de Moscou. Aus dem Bulletin de la Societe Imperiale des Naturalistes de Moscou. 18, 343-351.
Lennier, 1887. Études paléontologiques. Description des fossiles du Cap de la Hève. Bulletin de la Société Géologique de Normandie. 1886(12), 17-98.
Tarlo, 1959. Stretosaurus gen. nov., a giant pliosaur from the Kimmeridge Clay. Palaeontology. 2, 39-55.
Knutsen, 2012. A taxonomic revision of the genus Pliosaurus (Owen, 1841a) Owen, 1841b. Norwegian Journal of Geology. 92, 259-276.
Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013. A giant pliosaurid skull from the Late Jurassic of England. PLoS ONE. 8(5), e65989.
P. brachydeirus (Owen, 1841) Owen, 1842
= Plesiosaurus brachydeirus Owen, 1841
?= Pliosaurus evansi Seeley, 1869
?= Plesiosaurus sterrodeirus Seeley, 1869
Early Kimmeridgian, Late Jurassic
Kimmeridge Clay Formation, England

? Late Callovian-Early Oxfordian, Middle-Late Jurassic
? Oxford Clay Formation, England

References- Owen, 1841. Odontography; or a treatise on the comparative anatomy of the teeth, I Part 11. Dental system of reptiles. Hippolyte Bailliere, London. 179-295.
Owen, 1842. Report on British fossil reptiles. Part II. Report of the Eleventh Meeting of the British Association for the Advancement of Science. 60-204.
Seeley, 1869. Index to the fossil remains of Aves, Ornithosaurier and Reptilia in the Woodwardian Museum of the University of Cambridge. 143 pp.
P. archiaci (Deslongchamps vide Lennier, 1870) Sauvage, 1894
= Polyptychodon archiaci Deslongchamps vide Lennier, 1870
Kimmeridgian, Late Jurassic
Kimmeridge Clay Formation, France
References
- Lennier, 1870. Etudes géologiques et paléontologiques sur l'embouchure de la Seine et les Falaises de la Haute-Normandie. Imprimerie Eugène Costey, Havre. 245 pp.
Sauvage, 1894. Les Reptiles du terrain jurassique supérieur du Boulonnais. Comptes rendus hebdomadaires des séances de l'Academie des sciences. 119, 926-927.
P. brachyspondylus (Owen, 1839) Eichwald, 1868
= Plesiosaurus brachyspondylus Owen, 1839
Late Kimmeridgian, Late Jurassic
Kimmeridge Clay Formation, England

References- Owen, 1839. Report on British fossil reptiles Part 1. Report of the Ninth Meeting for the British Association for the Advancement of Science, Birmingham. 43-126.
Eichwald, 1868. Lethaea Rossica ou Paléontologie de la Russie. Stuttgart, Germany. 1304 pp.
P. carpenteri Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013
Late Kimmeridgian, Late Jurassic
Kimmeridge Clay Formation, England

Reference- Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013. A giant pliosaurid skull from the Late Jurassic of England. PLoS ONE. 8(5), e65989.
P? frearsi (Fischer de Waldheim, 1845) Bogoljubow, 1912
= Spondylosaurus frearsi Fischer de Waldheim, 1845
Kimmeridgian, Late Jurassic
Kimmeridge Clay, Russia
References
- Fischer de Waldheim, 1845. Notice sur le Spondylosaurus genre de saurien fossile de l'oolithe de Moscou. Aus dem Bulletin de la Societe Imperiale des Naturalistes de Moscou. 18, 343-351.
Bogoljubow, 1912. Die russischc oberjurassische Plesiosaurierfauna. Annuaire géologique et minéralogique de la Russie. 14, 1-7.
P. funkei Knutsen, Druckenmiller and Hurum, 2012
Tithonian, Late Jurassic
Slottsmøya Member of the Agardhfjellet Formation, Norway
Reference
- Knutsen, Druckenmiller and Hurum, 2012. A new species of Pliosaurus (Sauropterygia: Plesiosauria) from the Middle Volgian of central Spitsbergen, Norway. Norwegian Journal of Geology. 92, 235-258.
P? gamma Phillips, 1871
Late Callovian-Early Oxfordian, Middle-Late Jurassic
Oxford Clay Formation, England

Reference- Phillips, 1871. Geology of Oxford and the Valley of the Thames. The Clarendon Press, Oxford, UK. 523 pp.
P? grandis (Owen, 1839) Owen, 1841
= Plesiosaurus grandis Owen, 1839
Kimmeridgian-Early Tithonian, Late Jurassic
Kimmeridge Clay Formation, England
References- Owen, 1839. Report on British fossil reptiles Part 1. Report of the Ninth Meeting for the British Association for the Advancement of Science, Birmingham. 43-126.
Owen, 1841. Report on British fossil reptiles Part 2. Report of the Eleventh Meeting for the British Association for the Advancement of Science, Plymouth. 60-204.
P. irgisensis (Novozhilov, 1948) Tarlo, 1960
= Peloneustes irgisensis Novozhilov, 1948
= Strongylokrotaphus irgisensis (Novozhilov, 1948) Novozhilov, 1964
Tithonian, Late Jurassic
Savel-evsk Mine No. 1, Russia
References
- Novozhilov, 1948. Two new pliosaurs from the Lower Volga beds Povolzhe (right bank of Volga). Doklady Akademii Nauk SSSR. 60, 115-118.
Tarlo, 1960. A review of Upper Jurassic pliosaurs. Bulletin of the British Museum (Natural History), Geology. 4, 145-189.
Novozhilov, 1964. Order of Sauropterygia. Osnovy Paleontologii. 12, 309-332.
P. kevani Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013
Early Kimmeridgian, Late Jurassic
Kimmeridge Clay Formation, England

Reference- Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013. A giant pliosaurid skull from the Late Jurassic of England. PLoS ONE. 8(5), e65989.
P. macromerus Phillips, 1871 (as Pleiosaurus macromerus)
= Stretosaurus macromerus (Phillips, 1871) Tarlo, 1959
= Liopleurodon macromerus (Phillips, 1871) Halstead, 1989
Early Tithonian, Late Jurassic
Kimmeridge Clay Formation, England
References- Phillips, 1871. Geology of Oxford and the Valley of the Thames. The Clarendon Press, Oxford, UK. 523 pp.
Tarlo, 1959. Stretosaurus gen. nov., a giant pliosaur from the Kimmeridge Clay. Palaeontology. 2, 39-55.
Halstead, 1989. Plesiosaur locomotion. Journal of the Geological Society. 146, 37-40.
P? nitidus Phillips, 1871
Kimmeridgian-Early Tithonian, Late Jurassic
Kimmeridge Clay Formation, England
Reference- Phillips, 1871. Geology of Oxford and the Valley of the Thames. The Clarendon Press, Oxford, UK. 523 pp.
P. patagonicus Gasparini and O'Gorman, 2014
Middle Tithonian, Late Jurassic
Vaca Muerta Formation of the Mendoza Group, Neuquen, Argentina

Reference- Gasparini and O'Gorman, 2014. A new species of Pliosaurus (Sauropterygia, Plesiosauria) from the Upper Jurassic of northwestern Patagonia, Argentina. Ameghiniana. 51(4), 269-283.
P? planus Hulke, 1883
Kimmeridgian-Early Tithonian, Late Jurassic
Kimmeridge Clay Formation, England
Reference- Hulke, 1883. The anniversary address of the president. Quarterly Journal of the Geological Society of London. 39, 38-65.
P. portentificus Noe, Smith and Walton, 2004
Late Kimmeridgian, Late Jurassic
Kimmeridge Clay Formation, England
Reference- Noe, Smith and Walton, 2004. A new species of Kimmeridgian pliosaur (Reptilia; Sauropterygia) and its bearing on the nomenclature of Liopleurodon macromerus. Proceedings of the Geologists' Association. 115, 13-24.
P. rigauxi (Sauvage, 1874) new comb.
= Cetiosaurus rigauxi Sauvage, 1874
Middle Tithonian, Late Jurassic
Portel, France
Holotype
- (MHNL coll.; = MHNB 233) posterior cervical centrum (85 mm)
Comments- Sauvage (1874) initially described this as a posterior cervical centrum of a new Cetiosaurus species, notable for its short proportions. In 1895 he referred it to Pliosaurus sp. in a brief note, which would technically make it Pliosaurus rigauxi. That combination has never been used to my knowledge, however. Sauvage (1902) later referred the specimen to Pliosaurus grandis without stated justification. P? grandis is based on a scapula and three unassociated propodials from the Kimmeridgian of England which were poorly described and lost, so there's no reason to connect them to rigauxi. No more recent studies have been published, and the specimen has never been illustrated. The specimen number is taken from Fossilworks' website, though the Museum d'Histoire naturelle de Boulogne-sur-Mer (MHNB) closed in 2003 and transferred its collections to the Musée d'Histoire naturelle de Lille. Thus rigauxi presumedly has a MHNL number now instead.
Based on the original description, rigauxi differs from sauropods such as Cetiosaurus in the short centrum (52% of height, compared to no less than 154% in Cetiosaurus), with even the short-necked Brachytrachelopan having subequal proportions at best. Furthermore, all gravisaurs have strongly opisthocoelous cervicals, whereas rigauxi's is weakly amphicoelous. The parapophyses are 55% of centrum height, while they are much smaller in sauropods (e.g. 34% in cervical 12 of Cetiosaurus). All of these features are seen in pliosaur posterior cervicals however (e.g. length/width ratio of 51-58% and parapophysis/centrum height ratio 70% in P. brachydeirus' holotype). The undivided parapophysis located on the centrum indicates Sauvage had its position in the vertebral column correct despite assigning it to the wrong group. Notably, Sauvage states the ventral surface is "cut into a peak", which suggests a median keel like that which characterizes P. brachydeirus. The latter species' holotype is from the Early Kimmeridgian, but it's possible rigauxi is part of a long-lived P. brachydeirus, or that the French species will be shown to be distinct from P. brachydeirus once it is studied in more detail. rigauxi is not officially synonymized here though, pending verification of the keeled morphology and modern description of the specimen.
References- Sauvage, 1874. Mémoire sur les dinosauriens et les crocodiliens des terrains jurassiques de Boulogne-sur-Mer. Mémoires de la Société Géologique de France, série 2. 10(2), 1-57.
Sauvage, 1895. Les dinosauriens du terrain jurassique supérieur du Boulonnais. Bulletin de la Société Géologique de France, 3e série. 22, 465-470.
Sauvage, 1902. Note sur quelques Reptiles du Jurassique supérieur du Boulonnais. Bulletin de la Societe Academique de l‘Arrondissement de Boulogne-sur-Mer. 6, 380-398.
Fossilworks http://fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&taxon_no=65160
P. rossicus Novozhilov, 1948
= Liopleurodon rossicus (Novozhilov, 1948) Halstead, 1971
Tithonian, Late Jurassic
Buinsk Mine oil shales, Russia
References
- Novozhilov, 1948. Two new pliosaurs from the Lower Volga beds Povolzhe (right bank of Volga). Doklady Akademii Nauk SSSR. 60, 115-118.
Halstead, 1971. Liopleurodon rossicus (Novozhilov) - a pliosaur from the Lower Volgian of the Moscow basin. Palaeontology. 14, 566-570.
P? simplex Phillips, 1871
Kimmeridgian-Early Tithonian, Late Jurassic
Kimmeridge Clay Formation, England
Reference- Phillips, 1871. Geology of Oxford and the Valley of the Thames. The Clarendon Press, Oxford, UK. 523 pp.
P? suprajurensis Sauvage, 1879
Tithonian, Late Jurassic
Boulogne-sur-Mer, France

Reference- Sauvage, 1879. Prodrome des Plesiosauriens et des Elasmosauriens des formations Jurassiques superieures de Boulogne-sur-Mer. Annales des Sciences Naturelles, 6 Serie. 8(13), 1-38.
P. westburyensis Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013
Late Kimmeridgian, Late Jurassic
Kimmeridge Clay Formation, England

Reference- Benson, Evans, Smith, Sassoon, Moore-Faye, Ketchum and Forrest, 2013. A giant pliosaurid skull from the Late Jurassic of England. PLoS ONE. 8(5), e65989.
P. wosinskii Fischer de Waldheim, 1846
Kimmeridgian, Late Jurassic
Kimmeridge Clay, Russia
Reference
- Fischer de Waldheim, 1846. Notice sur quelques sauriens fossiles du Gouvernement de Moscou. Bulletin de la Societe Imperiale des Naturalistes de Moscou. 19, 90-107.
P? sp. (Lennier, 1887)
= "Tapinosaurus" Lennier, 1887
Late Kimmeridgian, Late Jurassic
lower Argiles d'Ecqueville Member of the Argiles d'Octeville Formation, France
Material
- (Muséum d’Histoire naturelle du Havre; destroyed) anterior cervical centrum (70 mm) (Lennier, 1887)
(Muséum d’Histoire naturelle du Havre; destroyed) three cervical centra (80, 78, 65 mm), two cervical ribs, two partial cervical or anterior dorsal neural arches, four incomplete dorsal neural arches, six dorsal ribs (four partial, one fragmentary; 1 m) (Rabeck, 1925)
?...(Lepage coll. 15.8.31.E1) posterior cervical centrum (91 mm) (Lepage et al., 2009)
?...(Muséum du Havre IIFD358) incomplete dorsal rib (Lepage et al., 2009)
Early Kimmeridgian, Late Jurassic
Marnes de Bleville, France

(Muséum d’Histoire naturelle du Havre; destroyed) proximal dorsal rib (Lennier, 1887)
Comments- Lennier (1887) referred an unassociated cervical centrum and dorsal rib (both originally in the Muséum d’Histoire naturelle du Havre, but destroyed in 1944) to Tapinocephalus, then thought to be dinosaurian but now known to be a dinocephalian synapsid. This was seemingly due to their large size. The figure caption of Lennier's plate illustrating the centrum mistakenly said "Tapinosaurus sp?", clearly a misspelling of Tapinocephalus and not previously suggested to be a valid genus.
Rabeck (1925) described a specimen found in 1923 as the dinosaur "Tapinosaurus sp?", based on resemblence to Lennier's specimens, unaware that "Tapinosaurus" was not an accepted genus. This specimen (consisting of partial vertebrae and ribs) was also held at the Muséum d’Histoire naturelle du Havre and similarly destroyed in WWII. Stiegelmann (1925) provided measurements of the specimen.
After this, "Tapinosaurus" was seldomly mentioned. Kuhn (1939) includes Rabeck's material questionably under Omosaurus, which is treated as Saurischia indet. by Kuhn despite being stegosaurid (a replacement name for Dacentrurus). Steel (1970) realized Lennier's article was supposed to reference Tapinocephalus, but stated Rabeck's material "seemingly pertains to a large dinosaur, but is indeterminable", placing "Tapinosaurus" in Sauropoda incertae sedis. Rabeck's specimen is not a sauropod, as it has short amphicoelous cervical centra without pleurocoels, short laterally oriented cervical ribs, dorsal neural arches lacking laminae, and single-headed dorsal ribs. Buffetaut et al. (1991) first noticed the discrepency between the article and plate caption in Lennier's work, and identified both this centrum and Rebeck's "Tapinosaurus" as sauropterygians. Similarly, Molnar (pers comm. in Olshevsky, 1991) stated these specimens were probably plesiosaurian. Finally, Lepage et al. (2009) published detailed overview of "Tapinosaurus"' history (forming the basis of most of this entry), and redescribed the specimens as Pliosaurus sp. for Lennier's and Pliosaurus cf. macromerus for Rabeck's. They also described three new specimens from the lower Argiles d'Ecqueville, two of which might be referrable to the same individual as Rabeck's specimen as they are from the same layer and of similar size.
"Tapinosaurus" is Pliosaurus?- Unfortunately, the alpha level taxonomy of Pliosaurus is currently controversial, and most proposed distinguishing characters are cranial. The only axial character currently used to distinguish Pliosaurus species is the median ventral keel on cervical centra of P. brachydeirus. Rabeck's specimen lacks this, as do P. brachyspondylus (both current and proposed neotypes), P. funkei (paratype), P. macromerus (lectotype), P. rossicus (holotype) and P. westburyensis. Yet this morphology is plesiomorphic, also being found in e.g. Brachauchenius, Simolestes and "P." andrewsi. Pliosaurus archiaci is based on a mandible (MNHN 24.1) also discovered in the Kimmeridgian deposits of Le Havre, but cannot be compared to "Tapinosaurus". Another method would be to correlate the "Tapinosaurus" specimens stratigraphically with known Pliosaurus species. According to Lepage et al., at least Rabeck's specimen derives from the Aulacostephanus mutabilis zone of the Late Kimmeridgian. This corresponds to CAMSM J35990, a partial skeleton initially referred to a broad concept of P. macromerus (Pliosaurus without ventral cervical keels) but more recently found to be closest to P. kevani (in an analysis that did not include the P. macromerus lectotype or proposed neotype). CAMSM J35990 is similar Rabeck's specimen in being larger than most and lacking ventral cervical keels, so there may be a real large A. mutabilis zone species of Pliosaurus that is currently undiagnosed. This may correspond to the large P. portentificus, although that has been considered a nomen dubium and may belong to the more recent A. euxodus zone. Perhaps notable is that P. portentificus has the same number of symphyseal alveoli (8) as P. archiaci, whereas other species have more (>11 in P. brachydeirus, 9 in P. carpenteri, ~14-15 in P. kevani) or less (6 in P. rossicus and P. patagonicus). In any case, both P. macromerus' lectotype and proposed neotype are from more recently deposited sediments, so Lepage et al.'s assignment seems unlikely. Pending further studies, the "Tapinosaurus" material is best retained as Pliosaurus sp.. Lennier's dorsal rib is from a different locality, whose age corresponds to P. brachydeirus and P. kevani, though it is near certainly indeterminate.
References- Lennier, 1887. Études paléontologiques. Description des fossiles du Cap de la Hève. Bulletin de la Société Géologique de Normandie. 1886(12), 17-98.
Rabeck, 1925. Notes sur la découverte d'ossements de dinosaurien dans les Argiles supérieures Kimméridgiennes du Cap de la Hève (Octeville-sur-Mer). Bulletin de la Société Géologique de Normandie. 1916/1923(34), 72-74
Stiegelmann, 1925. Note additionnelle [à Notes sur la découverte d'ossements de dinosaurien dans les Argiles supérieures Kimméridgiennes du Cap de la Hève (Octeville-sur-Mer) de G. Rabeck]. Bulletin de la Société Géologique de Normandie.1916/1923(34), 75.
Kuhn, 1939. Saurischia. In Fossilium Catalogus I. Animalia. 87. 124 pp.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp.
Buffetaut, Cuny and Le Loeuff, 1991. French dinosaurs: The best record in Europe? Modern Geology. 16, 17-42.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Lepage, Buffetaut and Lepage, 2009. Qu'est-ce que Tapinosaurus? Lennier, Rabeck et les grands Sauroptérygiens du Kimméridgien supérieur de la région havraise (Normandie, France). Bulletin de la Société géologique de Normandie et des amis du Muséum du Havre. 96(1), 27-59.

Brachaucheninae Williston, 1925 vide Benson and Druckenmiller, 2014

unnamed probable brachauchenine (Knoll, Collete, Dubus and Petit, 2000)
Early Albian, Early Cretaceous
Sables verts, France
Material
- (Petit coll.) posterior dorsal centrum (107 mm)
Comments- Knoll et al. (2000) originally described this as a possibly brachiosaurid sauropod proximal caudal, but it was reidentified by Buffetaut et al. (2005) as a pliosaurid dorsal. Given its age, it is probably referrable to Brachaucheninae, but is likely to be indeterminate.
References- Knoll, Collete, Dubus and Petit, 2000. On the presence of a sauropod dinosaur (Saurischia) in the Albian of Aube (France). Geodiversitas. 22, 389-394.
Buffetaut, Collete, Dubus and Petit, 2005. The "sauropod" from the Albian of Mesnil-Saint-Père (Aube, France): A pliosaur, not a dinosaur. Carnets de Géologie. 2005/01, 1-5.

Choristodera Cope, 1876
Definition- (Pachystropheus rhaeticus + Cteniogenys antiquus + Lazarussuchus inexpectatus + Champsosaurus annectens) (modified from Merck, 1997)
Other definitions- (Cteniogenys antiquus + Lazarussuchus inexpectatus + Champsosaurus annectens) (modified from Dilkes, 1998)

Actiosaurus Sauvage, 1883
A. gaudryi Sauvage, 1883
Rhaetian, Late Triassic
limestone at Conches-les-Mines, Autun, Saône-et-Loire, France
Lectotype (proposed)
- (University of Leuven coll.) proximal humerus (~105 mm)
Paralectotypes- (University of Leuven coll.) (Amniota incertae sedis) centrum (65 mm), humerus (90 mm), proximal ?humerus
Comments- Sauvage described this as a probable dinosaur most similar to Palaeosaurus (based on Thecodontosaurus material) within the Megalosauridae (in which he placed all theropods), though differing in the more projecting femoral head, more poorly developed greater trochanter, and marked tuberosity on the proximal humerus. Sauvage based the taxon on two supposed proximal femora of different sizes, a humerus "found with them" and a more questionably referred centrum supposedly from the dorsal series. While the femoral morphology suggested something intermediate between crocodiles and lizards, Sauvage believed the vertebral centrum would indicate Actiosaurus was dinosaurian if properly referred (due to the oblique articular surfaces). The species name was misspelled A. gaudrii by Boulenger (1883). Later authors often placed it in whichever theropod family they included Paleosaurus in- Mayer (1886) in Amphisauridae, Nopcsa (1901) in Anchisaurinae within Megalosauridae, and Simroth (1907) and Zittel (1890) in Zanclodontidae. Zittel was provisional in his assignment though and stated Actiosaurus was insufficiently characterized. Huene (1902) listed it provisionally under Lycosauria (containing gorgonopsids and theriocephalans), and Werner (1906) listed it under Thecodontia. Sauvage (1903) disagreed with Huene's identification, noting that while resemblences between Actiosaurus and some "anomodonts" such as Deuterosaurus (now recognized as a dinocephalan) existed, the form was more similar to crocodiles. Note Stache's (1889) reference to Actiosaurus tommasinii is a misspelling of the ophidiomorph squamate Acteosaurus. Huene (1908) referred it to Ichthyosaurus, crediting his 1902 paper despite the fact that was not his conclusion then.
The various described elements were found by two different collectors, belonged to at least two individuals based on the two sizes of "femur" and have no record of association, so could easily belong to different taxa. The bone described as the smaller proximal femur is here taken to be the type as it is illustrated, described in most detail and first, and it is the supposed femora that are said to indicate "the presence of a reptile close to Palaeosaurus." Why Actiosaurus has been referred to Ichthyosauria is not at all apparent. The limb elements are more elongate than any ichthyosaur bone, while ichthyosaur vertebrae are famously amphicoelous unlike the weakly biconvex centrum referred to Actiosaurus. The limb elements are also more elongate with better developed articular surfaces than sauropterygians. Compared to contemporaneous dinosaurs, the humerus has a much smaller and proximally located deltopectoral crest and more greatly expanded distal end, while the supposed femur would differ in having a basically proximally projecting elongated and flattened head. However, the "femur" is extremely similar to the humerus of the choristodere Pachystropheus, which is common in Rhaetian beds of England. Points of similarity include the head structure just noted, as well as the deltopectoral crest (greater trochanter of Sauvage) which is an oval knob placed in the center of the anterior face, at the same distance from the head. It is also within the size range of Pachystropheus specimens (greatest transverse width 25 mm compared to 13-35 mm). Interestingly, Actiosaurus has priority over Pachystropheus (named by Huene in 1935). However, Pachystropheus' proximal humeri exhibit so much individual variation and are not distinct from other choristoderes (e.g. Irenosaurus). Thus Actiosaurus is best left as Choristodera indet.. Interestingly, Godefriot (pers. comm. 2012) has rediscovered Actiosaurus' remains in the University of Leuven and confirms my 2010 identification. Whether the additional proximal "femur" is also a choristodere humerus is unknown, as it is not illustrated and only stated to be larger (though with oddly different bredth vs. depth proportions given). The element described as a humerus seems to be correctly identified, but it differs from Pachystropheus in being more abruptly expanded proximally and straighter, with a more proximally placed deltopectoral crest. Its affinities require a broader look at amniote humeri. The centrum is not from a choristodere, as it is weakly biconvex to amphiplatyan, taller than long, has oblique articular faces and a longitudinal ventral groove (as in Pachystropheus caudals, but those further differ in being fused to their neural arches at a much smaller size than the Actiosaurus centrum). Among dinosaurs, only caudal vertebrae usually have ventral grooves (common in theropods at least), though only cervicals usually have oblique faces. Plateosaurus ingens does have oblique faces on its caudal centra though, making it a potential candicate for the Actiosaurus centrum. Yet with such a brief description and no illustration, further comparison to other amniotes is warranted before any judgement is made.
References- Boulenger, 1883. Reptilia and Batrachia. in Rye (ed.). The Zoological Record for 1883. Record of Zoological Literature. 20, 24 pp.
Sauvage, 1883. Recherches sur les reptiles trouves dans l'etage Rhetien des environs d'Autun. Annales des Sciences Geologiques. 14(6, Article 3), 1-44.
Mayer, 1886. Herausgegeben von der Zoologischen Station zu Neapel. IV. Abtheilung: Tunicata, Vertebrata. 413 pp.
Stache, 1889. Die Liburnische Stufe und deren Grenzhorizonte, eine Studie uber die Schichtenfolgen der cretacisch-eocanen oder protocanen Landbildungs-Periode im Bereich der Küstenländer von Oesterreich-Ungarn mit einer einleitenden Uebersicht der geologischen Verhältnisse dieses Gebietes. Abhandlungen der geologischen Bundesanstalt. 13, 170 pp.
Zittel, 1890. Handbuch der Palaeontologie. Volume III. Vertebrata (Pisces, Amphibia, Reptilia, Aves).
Nopcsa, 1901. A dinosaurusok atnezete es szarmazasa. Földtani Közlöny. 31, 193-224.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen (Neue Serie). Gustav Fischer Verlag, Jena. 6, 1-84.
Sauvage, 1903. Note sur les reptiles de letage Rhetien des environs d'Autun. Bulletin Societe d'Histoire Naturelle d'Autun (France). 16, 309-318.
Werner, 1906. Reptilia und Amphibia fur 1903. Archiv fur Naturgeschichte. 1-70.
Simroth, 1907. Die Pendulationstheorie. Leipzig Konrad Grethlein's Verlag. 564 pp.
Huene, 1908. Die Dinosaurier der europäischen Triasformation mit Berücksichtiging der aussereuropäischen Vorkommnisse [The dinosaurs of the European Triassic Formation, with consideration of non-European occurrences]. Geologische und Paläontologische Abhandlungen Supplement-Band. 1, 419 pp.
Mortimer, online 2010. http://theropoddatabase.blogspot.com/2010/06/actiosaurus-is-choristodere-not.html

Patricosaurus Seeley, 1887
P. merocratus Seeley, 1887
Late Albian, Early Cretaceous
Gault Clay (reworked into Cambridge Greensand), England
Lectotype
- (SMC B58401) (~3 m) proximal femur (~107 mm)
Comments- Patricosaurus was based on two unassociated specimens, a sacral vertebra (SMC B58402) found before 1859 and a proximal femur discovered in the 1880's. Seeley (1887) believed both belonged to the same taxon since he thought "there was little chance of any remains of two lizards occurring" in the Cambridge Greensand, but stated the femur could remain as the type if the vertebra was found to not belong to the same taxon. Barrett and Evans (2002) redescribed the specimens and referred the vertebra to Archosauria, which makes the femur the lectotype. Although they called Patricosaurus indeterminate, they also listed several features which differed from all lepidosaurs they examined.
Seeley named the taxon as a new genus of lizard, more closely related to modern taxa than any other Cretaceous lizard known at the time, but outside the crown group. Patricosaurus was referred to Lacertilia incertae sedis without comment by most later authors as well. Barrett and Evans compared it to several modern lizard taxa, but found no characters that were phylogenetically useful in the proximal femur. Additionally, they could not distinguish it from rhynchocephalians, leaving them to classify Patricosaurus as Lepidosauria incertae sedis. Evans later (2003) listed Patricosaurus as a possible anguimorph without comment, perhaps because she and Barrett felt it was phenetically closest to terrestrial varanids. Barrett and Evans noted that the lepidosaur-like characters of Patricosaurus were symplesiomorphic for diapsids, correctly excluding it from turtles, plesiosaurs, ichthyosaurs and archosaurs. Choristoderes share the same femoral plesiomorphies as lepidosaurs, but were said to differ from Patricosaurus in having "narrower, more elongate, internal trochanter and an absence of obvious proximal muscle scars or strong muscle ridges." Yet the near contemporaneous Khurendukhosaurus does not have a narrower internal trochanter, has a rough insertional surface for the m. pubischiofemoralis externus (the dorsal surface is worn, so cannot be judged), and the length of Patricosaurus' internal trochanter is unknown as it is broken distally (unless they mean length in proximal view away from the head, in which case Khurendukhosaurus is in the range of modern lepidosaurs). Thus a relationship with choristoderes needs to be considered further. Of the characters listed by Barrett and Evans as variable in lepidosaurs, Khurendukhosaurus is less similar to Patricosaurus than some lizards in having almost no anterior curvature of its femoral head (instead the entire head is projected anteriorly as in Xantusia and Elgaria), is similar to the examined lepidosaurs in having weaker trochanteric crests (and thus a shallower intertrochanteric fossa), is more similar than most examined lepidosaurs (except Tupinambis and Cyclura) in having a small amount of separation between the internal trochanter and head, but is more similar to Patricosaurus than any examined lepidosaur in having a teardrop-shaped head in proximal view and in having the anteroposterior width be >95% of the dorsoventral width in proximal view. Cteniogenys (perhaps the choristodere Evans compared, as she previously redescribed it) is dissimilar in the latter three ways, while Pachystropheus is quite different in the proximally flattened head, distally placed interior trochanter and other features. Pending comparison to a greater variety of lepidosaurs (including fossil taxa), Patricosaurus is provisionally considered more likely to be a choristodere, perhaps closer to neochoristoderes than Cteniogenys. Evans (pers. comm. from Barrett, 2010) believes the taxon to be a lizard regardless of the details described here.
Olshevsky (1991) stated Patricosaurus was probably an indeterminate small theropod, citing personal communication from Molnar. However, Molnar stated (pers. comm., 2001) that while he was not convinced it is a lizard, he did not think it was a theropod or even an archosaur. Thus there was a misunderstanding, and no one actually ever considered Patricosaurus to be dinosaurian based on evidence. As Barrett and Evans stated, Patricosaurus cannot be an archosaur because members of that clade have lost the intertrochanteric fossa and internal trochanter. Similarly, there is no anterior or greater trochanter, unlike theropods.
References- Seeley, 1887. On Patricosaurus merocratus, Seeley, a lizard from the Cambridge Greensand, preserved in the Woodwardian Museum of the University of Cambridge. Quarterly Journal of the Geological Society of London. 43, 216-220.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Barrett and Evans, 2002. A reassessment of the Early Cretaceous reptile ‘Patricosaurus merocratus’ Seeley from the Cambridge Greensand, Cambridgeshire, UK. Cretaceous Research. 23, 231-240.
Evans, 2003. At the feet of the dinosaurs: The early history and radiation of lizards. Biological Reviews. 78(4), 513-551.

Pantestudines Joyce et al., 2004
Definition- (Testudo graeca <- Homo sapiens, Draco volans, Caiman crocodilus, Vultur gryphus) (proposed)
Other definitions- (Chelonia mydas <- Homo sapiens, Draco volans, Caiman crocodilus, Vultur gryphus) (modified from Joyce et al., 2004)
= Anapsida Osborn, 1903
Definition- (Testudines <- Sauria) (Gauthier et al., 1988)

Testudinata Klein, 1760
Definition- (carapace formed from costal bones with fused ribs, neural bones with fused thoracic vertebrae, and marginal bones; plastron formed from interclavicle, clavicle, and three to five paired bones sutured together; carapace and plastron articulated at lateral margin and enclosing shoulder girdle and pelvic girdle as in Chelonia mydas) (Joyce et al., 2004)

Testudines Batsch, 1788 (not Linnaeus, 1758)
Definition- crown(Chelus fimbriatus + Testudo graeca) (proposed)
Other definitions- (Chelonia mydas <- Captorhinus aguti) (modified from Gauthier et al., 1988)
(Proganochelys quenstedti + crown turtles) (modified from Laurin and Reisz, 1995)
crown(Chelus fimbriatus + Chelonia mydas) (Joyce et al., 2004)
= Chelonii Latreille, 1800
Definition- (Chelus fimbriatus + Testudo hermani) (modified from Kischlat and Timm, 2006)
= Chelonia Ross and Macartney, 1802 (preoccupied Sonnini and Latreille, 1801)

Trionychia Hummel, 1929
Definition- crown(Trionyx triunguis + Carettochelys insculpta) (Joyce et al., 2004)

Trionychidae Gray, 1825
Definition- crown(Trionyx triunguis + Cyclanorbis senegalensis) (Joyce et al., 2004)

Trionychinae Gray, 1825 vide Lydekker, 1889

Axestemys Hay, 1899
= Axestus Cope, 1872 (preoccupied Dejean, 1835)
= Temnotrionyx Hay, 1908
= Paleotrionyx Schmidt, 1945
Definition- (Axestemys byssina <- Aspideretoides foveatus, Apalone spinifera, Aspideretes gangeticus, Rafetus euphraticus, Trionyx triunguis, Cyclanorbis senegalensis, Plastomenus thomasii, Chitra indica, Pelodiscus sinensis) (Vitek, 2012)
Comments- Axestus was originally used for a curculionid beetle (Dejean, 1835).
References- Dejean, 1835. Catalogue des Coléoptères de la collection de M. le Comte Dejean. [Livraison 4]. Méquignon-Marvis. 257-360.
Cope, 1872. Descriptions of some new Vertebrata from the Bridger Group of the Eocene. Proceedings of the American Philosophical Society. 12, 460-465.
Hay, 1899. On the nomenclature of certain American fossil vertebrates. The American Geologist. 24, 345-349.
Hay, 1908. The fossil turtles of North America. Carnegie Institution of Washington Publication. 75, 568 pp.
Schmidt, 1945. A new turtle from the Paleocene of Colorado. Fieldiana: Geology. 10, 1-4.
Vitek, 2012. Giant fossil soft-shelled turtles of North America. Palaeontologia Electronica. 15(1), 13A.
A. splendida (Hay, 1908) Vitek, 2012
= Aspideretes splendidus Hay, 1908
= Trionyx splendidus (Hay, 1908) Hummel, 1932
= Aspideretes granifer Gardiner, Russell and Brinkman, 1995
= Aspideretes planus Gardiner, Russell and Brinkman, 1995
= Aspideretoides splendidus (Hay, 1908) Gardiner, Russell and Brinkman, 1995
= Eugenichelys splendida (Hay, 1908) Chkhikvadze, 2000
Late Campanian, Late Cretaceous
Judith River Formation, Montana

Holotype- (AMNH 3952) incomplete carapace
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, North Dakota, US

Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US

Referred- (KUVP 152429; paratype of Dakotaraptor steini) entoplastron
(NCSM 13170; paratype of Dakotaraptor steini) entoplastron
(PBMNH.P.10.113.T; holotype of Dakotaraptor steini in part) partial entoplastron
Diagnosis- (after Vitek, 2015) dorsomedian carapacial sculpture of larger individuals consists of high, narrow ridges bordering wavy, flat-bottomed, reticulate lacunae or troughs; unsutured median contact between xiphiplastra; epiplastral projections approximately one-fourth maximum hypoplastral width; anterior projections of epiplastra wider and not tapered; gracile entoplastron; sculptured, unfused hyo-hypoplastral callosities.
References- Hay, 1908. The fossil turtles of North America. Carnegie Institution of Washington Publication. 75, 568 pp.
Hummel, 1932. Pars 52: Trionychia fossilia. In Quenstedt (ed.). Fossilium Catalogus. I: Animalia. W. Junk. 106 pp.
Gardiner, Russell and Brinkman, 1995. Systematics and taxonomy of soft-shelled turtles (family Trionychidae) from the Judith River Group (Mid-Campanian) of North America. Canadian Journal of Earth Sciences. 32(5), 631-643.
Ckhikvadze, 2000. O sistematicheskom polozheniye nekotorikh vimershikh trekhkogotnikh cherepakh Severnoi Ameriki i Azii. Trudy Tbilisskogo gosudarstvennogo pedagogicheskogo universiteta. 7, 199-213.
Vitek, 2012. Giant fossil soft-shelled turtles of North America. Palaeontologia Electronica. 15(1), 13A.
Arbour, Zanno, Larson, Evans and Sues, 2015. The furculae of the dromaeosaurid dinosaur Dakotaraptor steini are trionychid turtle entoplastra. PeerJ PrePrints. https://doi.org/10.7287/peerj.preprints.1570v1
DePalma, Burnham, Martin, Larson and Bakker, 2015. The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation. Paleontological Contributions. 14, 16 pp.
A. montinsana Vitek, 2012
Selandian-Thanetian, Middle-Late Paleocene
Melville Formation, Montana, US

Danian, Early Paleocene
Fort Union Formation, North Dakota, US
Danian, Early Paleocene
Denver Formation, Colorado, US

Reference- Vitek, 2012. Giant fossil soft-shelled turtles of North America. Palaeontologia Electronica. 15(1), 13A.
A. cerevisia Vitek, 2012
Ypresian-Lutetian, Early Eocene
Bridger Formation, Wyoming, US

Reference- Vitek, 2012. Giant fossil soft-shelled turtles of North America. Palaeontologia Electronica. 15(1), 13A.
A. quinni (Schmidt, 1945) Hutchison and Holroyd, 2003
= Paleotrionyx quinni Schmidt, 1945
Thanetian-Ypresian, Late Plaeocene-Early Eocene
Plateau Valley beds, Colorado, US

References- Schmidt, 1945. A new turtle from the Paleocene of Colorado. Fieldiana: Geology. 10, 1-4.
Hutchison and Holroyd, 2003. Late Cretaceous and Early Paleocene turtles of the Denver Basin. Rocky Mountain Geology. 38, 121-142.
A. byssinus (Cope, 1872) Hay, 1899
= Axestus byssinus Cope, 1872
= Temnotrionyx manducans Hay, 1908
= Eugenichelys robertemryi Chkhikvadze, 2008
Ypresian-Lutetian, Early Eocene
Bridger Formation, Wyoming, US
Ypresian, Early Eocene
Wasatch Formation, Wyoming, US

References- Cope, 1872. Descriptions of some new Vertebrata from the Bridger Group of the Eocene. Proceedings of the American Philosophical Society. 12, 460-465.
Hay, 1899. On the nomenclature of certain American fossil vertebrates. The American Geologist. 24, 345-349.
Hay, 1908. The fossil turtles of North America. Carnegie Institution of Washington Publication. 75, 568 pp.
Chkhikvadze, 2008. Trekhkogotniye cherepakhi (Trionychidae) Azii i Severnoi Ameriki: Morfologiya, filogeniya, sistematika, terminologiya elementov karapaksa. Problemy paleobiologii, Tbilisi. 3, 85-95.

Chelonioidea Gray, 1825 vide Agassiz, 1857
Definition- (Chelonia mydas + Dermochelys coriacea) (Joyce et al., 2004)

Protostegidae Cope, 1889

Pneumatoarthrus Cope, 1870
P. peloreus Cope, 1870
Early Maastrichtian, Late Cretaceous
Mount Laurel Formation, New Jersey, US
Holotype
- (ANSP 9225) (~2.8 m) fifth dorsal centrum (~128 mm), sixth dorsal centrum (~129 mm), seventh dorsal centrum (~127 mm), eighth dorsal centrum (~112 mm) (Leidy, 1865)
Comments- This specimen was originally described as a juvenile Hadrosaurus sacrum by Leidy (1865) before Cope (1870) named it as a new taxon. Cope believed it to be "more Megalosaurian than Iguanodontine", comparing it favorably to Anchisaurus (his Megadactylus), Efraasia (his Palaeosaurus) and Clepsysaurus (a parasuchian), and less closely to Ornithopsis and Dryptosaurus (his Laelaps). He also stated it may belong to Ornithotarsus or a turtle, and in his 1872 description of the sea turtle Protostega considered Pneumatoarthrus quite likely to be a turtle as well. Cope (1875) later listed it under Testudinata with sea turtles. However, most authors were unaware of Cope's reassignment and continued to refer it to a dinosaur, either a hadrosaurid (Hay, 1903 [his Trachodontidae], 1930; Russell, 1930; Kuhn, 1964; Chapman and Brett-Surman, 1990) or theropod (Huene, 1932; Lull and Wright, 1942; Miller, 1955; Romer, 1956, 1976; Steel, 1970; White, 1973). Of the latter, Huene illustrated it as a Dryptosaurus aquilunguis sacral, Lull and Wright stated it resembled Anchisaurus, while White placed it as Megalosauridae indet.. The sacrals of Anchisaurus are roughly similar, but it and other dinosaurs have much smaller intervertebral foramina (so-called pneumatic foramina of Cope). The seemingly large foramina on Anchisaurus' caudals are spaces where transverse processes have broken. The similarity to Efraasia and Clepsysaurus noted by Cope involves the alternating expansion and contraction of the neural canal, which is common in reptiles including Archelon. Baird independantly recognized Pneumatoarthrus as a protostegid turtle a century after Cope's identification, which was first noted in personal communication in Gillette (1978), and later described in detail by Baird (1979, 1984) as a protostegine. He considered it extremely similar to Protostega and indistinguishable from Archelon, though he thought it to be generically undiagnostic and thus not a senior synonym of the latter genus. No rationale for referring it to Protosteginae instead of (the paraphyletic) Chelospharginae was given, and today we know of several more basal protostegids as well. It was listed as a protostegid by Spamer et al. (1995) and as a possible synonym of Archelon by Glut (1997). Another possibility is that Pneumatoarthrus is a junior synonym of the protostegid Atlantochelys mortoni, which is also known from Late Cretaceous New Jersey. Unfortunately, protostegid dorsals have not been closely studied for systematic variation so Pneumatoarthrus is retained here as Protostegidae indet..
References- Leidy, 1865. Memoir on the extinct reptiles of the Cretaceous formations of the United States. Smithsonian Contributions to Knowledge. 14(6), 135 pp.
Cope, 1870. Observations on the Reptilia of the Triassic formations of the Atlantic region of the United States. Proceedings of the American Philosophical Society. 11, 444-446.
Cope, 1872. A description of the genus Protostega, a form of extinct Testudinata. Proceedings of the American Philosophical Society. 12, 422-433.
Cope, 1875. The Vertebrata of the Cretaceous formations of the West. Report of the United States Geological Survey of the Territories. 2, 303 pp.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America. United States Geological Survey Bulletin. 179, 868 pp.
Gilmore, 1924. A new species of hadrosaurian dinosaur from the Edmonton Formation (Cretaceous) of Alberta. Canada Department of Mines, Geological Survey Bulletin. 38, 13-26.
Hay, 1930. Second bibliography and catalogue of the fossil Vertebrata of North America. Carnegie Institution of Washington Publication. 390(2), 1074 pp.
Russell, 1930. Upper Cretaceous dinosaur faunas of North America. Proceedings of the American Philosophical Society. 69(4), 133-159.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Lull and Wright, 1942. Hadrosaurian dinosaurs of North America. Geological Society of America Special Papers. 40, 242 pp.
Miller, 1955. A check-list of the Cretaceous and Tertiary vertebrates of New Jersey. Journal of Paleontology. 29(5), 903-914.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Kuhn, 1964. Pars 105. Ornithischia (Supplementum I). in Westphal (ed.). Fossilium Catalogus. I: Animalia. 80 pp.
White, 1973. Catalogue of the genera of dinosaurs. Annals of Carnegie Museum. 44(9), 117-155.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Gillette, 1978. Catalogue of type specimens of fossil vertebrates Academy of Natural Sciences, Philadelphia. Part IV: Reptilia, Amphibia, and tracks. Proceedings of the Academy of Natural Sciences of Philadelphia. 129, 101-111.
Baird, 1979. Pneumatoarthrus Cope, 1870, not a dinosaur but a sea-turtle. Proceedings of the Academy of Natural Sciences of Philadelphia. 129, 71-81.
Baird, 1984. Evidence of giant protostegid sea-turtles in the Cretaceous of New Jersey. The Mosasaur. 2, 135-140.
Chapman and Brett-Surman, 1990. Morphometric observations on hadrosaurid ornithopods. in Carpenter and Currie (eds.). Dinosaur Systematics: Approaches and Perspectives. Cambridge University Press, Cambridge. 163-177.
Spamer, Daeschler and Vostreys-Shapiro, 1995. A study of fossil vertebrate types in the Academy of Natural Sciences of Philadelphia: Taxonomic, systematic and historical perspectives. The Academy of Natural Sciences of Philadelphia. Special Publication 16, 435 pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076 pp.

Simiosauria Senter, 2004
Definition- (Drepanosaurus unguicaudatus <- Coelurosauravus elivensis, Sauria) (modified from Senter, 2004)
= Drepanosauromorpha Renesto, Spielmann, Lucas and Spagnoli, 2010

Drepanosauridae Olsen and Sues, 1986
Definition- (Drepanosaurus unguicaudatus + Megalancosaurus preonensis + Dolabrosaurus aquatilis) (modified from Dilkes, 1998)
= Megalancosauridae Olshevsky, 1991
Definition- (Drepanosaurus unguicaudatus + Megalancosaurus preonensis) (modified from Merck, 1997

Megalancosaurus Calzavara, Muscio and Wild, 1980
M. preonensis Calzavara, Muscio and Wild, 1980
Middle Norian, Late Triassic
Dolomia di Forni Formation, Italy
Holotype
- (MFSN 1769) skull (30 mm), mandibles (18 mm), hyoid, several cervical vertebrae (third 7 mm, sixth 9 mm), second dorsal neural spine, fused third and fourth dorsal neural spines, two dorsal ribs, dorsal rib fragments, scapula (23 mm), coracoid fragment, humerus (22 mm), radius (15 mm), ulna (15 mm), intermedium, ulnare, centrale, distal carpal I, distal carpal II, distal carpal III, metacarpal I (1.5 mm), phalanx I-1 (6 mm), partial manual ungual I, metacarpal II (3 mm), incomplete phalanx II-1 (6 mm), metacarpal III (3.5 mm), partial phalanx III-1 (6 mm), manual ungual III (4 mm), metacarpal IV (3 mm), partial phalanx IV-1 (3 mm), phalanx IV-2 (5 mm), manual ungual IV (4.5 mm), metacarpal V (2 mm), partial phalanx V-1 (3 mm), phalanx V-2 (4 mm), manual ungual V (4.5 mm)
Referred- (MFSN 1801) caudal vertebrae 8-38 fused to chevrons (Pinna, 1987)
(MFSN 18443a) caudal vertebrae 13-38 (Renesto, 2000)
Middle Norian, Late Triassic
Zorzino Limestone Formation, Italy
(MBSN 26; paratype of Drepanosaurus unguicaudatus) cervical vertebrae (third 4.5 mm, sixth 6 mm), anterior dorsal vertebrae (second 4 mm), anterior dorsal ribs, posterior dorsal vertebrae fused to dorsal ribs (eighteenth 3 mm), supraneural element, caudal vertebrae fused with chevrons (tenth 5.5 mm), scapulae (21 mm), humerus (18.5 mm), radius, ulna, pelvis, femur (20 mm), tibia (11.5 mm), fibula, pes (Pinna, 1980)
(MPUM 6008; = P 11 24) cervical vertebrae (third 7.5 mm, fifth 9.5 mm, sixth 10 mm), anterior dorsal vertebrae (second 6 mm), anterior dorsal ribs, posterior dorsal vertebrae fused to dorsal ribs, supraneural element, sacral vertebrae, caudal neural spines, scapula (23 mm), coracoids fused to sternal plates, furcula, humerus (22.5 mm), radius (14 mm), ulna (14 mm), proximal carpal, four distal carpals, phalanx I-1 (6 mm), manual ungual I (4 mm), metacarpal II (3 mm), phalanx II-1 (6 mm), manual ungual II (4 mm), metacarpal III (3 mm), phalanx III-1 (6 mm), manual ungual III (4 mm), metacarpal IV (2.5 mm), phalanx IV-1 (3 mm), phalanx IV-2 (5 mm), phalanx V-1 (2.5 mm), phalanx V-2 (4 mm), manual ungual V (3.5 mm), three manual phalanges, two manual unguals, partial pelvis (Renesto, 1994)
(MPUM 8437; = CCSR 63115) posterior skull, incomplete mandible, eight cervical vertebrae (fifth 7 mm, sixth 7.5 mm), five anterior dorsal vertebrae, anterior dorsal ribs, eighteen posterior dorsal vertebrae fused to dorsal ribs, supraneural element, three sacral vertebrae, thirty-nine caudal vertebrae fused to chevrons, partial scapula, partial coracoid, furcula, distal humerus (~21 mm), radius (11 mm), ulna (12 mm), intermedium, ulnare, two centrales, distal carpal I, distal carpal II, distal carpal III, distal carpal IV, distal carpal V, metacarpal I, phalanx I-1, partial manual ungual I, metacarpal II, incomplete phalanx II-1, incomplete manual ungual II, metacarpal III (4 mm), phalanx III-1 (5 mm), manual ungual III, metacarpal IV, phalanx IV-1, phalanx IV-2, manual ungual IV, metacarpal V, phalanx V-1, incomplete phalanx V-2, manual ungual V, partial pelvis, femora (27 mm), tibiae (17.5 mm), fibulae (16 mm), astragali, calcanea, centrale, distal tarsal I, distal tarsal II, distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (4 mm), phalanx III-1 (3 mm), phalanx III-2, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx V-1, phalanx V-2, pedal ungual (Renesto, 2000)
Late Triassic?
Italy?

(MCSNB 7833) (Senter, 2004)
Comments- Megalancosaurus preonensis was discovered in 1980 and originally assigned to Pseudosuchia sensu Huene (Calzavara et al., 1980). Although Carroll (1988) also placed it in Thecodontia (in the traditional paraphyletic sense), generally only those workers who reject cladistics and a dinosaurian origin for birds have continued to call Megalancosaurus an archosaur (e.g. Feduccia and Wild, 1993; Feduccia, 1996). This is almost exclusively based on its supposed antorbital fenestra (considered near certainly absent by Renesto and Dalla Vecchia, 2005), and presumably the need to have birds derive from archosaurs. Megalancosaurus is currently placed in the larger clade Simiosauria, which has a highly uncertain placement among eosuchians. Various analyses place them outside Neodiapsida, sister to Euryapsida, in Lepidosauromorpha, as non-thecodont archosauromorphs or as 'protorosaurs', with the latter three possibilities sometimes including a close relationship with pterosaurs. Determining their relationships will require a larger diapsid phylogenetic analysis than those currently published.
Megalancosaurus a theropod? Olshevsky (1991) believed Megalancosaurus to be a basal theropod (or in his taxonomy, a basitheropod theropodomorph), but this was based only on the holotype. Of the theropodomorph characters he lists, carnivorous dentition is primitive for gnathostomes, while new specimens show Megalancosaurus lacks erect limbs and a reduced calcaneum. Of Olshevsky's basitheropod characters, an antorbital fenestra is primitive for archosauriforms and probably lacking in Megalancosaurus, "generally avian appearence of the skull" is vague and unlike basal theropods, relatively large forelimbs are primitive for tetrapods and unlike basal theropods, "clavicles, fused clavicles, or primitive furcula" covers every possibility though Megalancosaurus' are fused as in theropods, and pentadactyl manus and pes are plesiomorphic for tetrapods and not found in basal theropods. The tarsus is not even incipiently mesotarsal and as noted above the calcaneum is not reduced. Megalancosaurus does share the presence of at least three sacral vertebrae with dinosaurs, but this is present in pterosaurs and some other taxa as well. Furthermore, the more basal Vallesaurus and Drepanosaurus only have two sacrals. While Megalancosaurus and theropods both have manus capable of grasping, in theropods digit I is angled towards II and III due to an asymmetrical metacarpal I articulation and twisted phalanx I-1, whereas in Megalancosaurus half the digits oppose the other half due merely to a lack of articulation between the metacarpals. This suggests the grasping abilities are convergent. Megalancosaurus lacks numerous characters expected in a basal theropod, such as subnarial fenestra, external mandibular fenestra, thecodont dentition, more than eight cervicals, cervical epipophyses, vertebral laminae, dicephalous dorsal ribs, reduced manual digits IV and V, perforated acetabulum, dorsally angled preacetabular process, elongate postacetabular process, elongate pubis and ischium, inturned femoral head, anterior and fourth trochanter, mesotarsal ankle, reduced calcaneum, centrale absent, less than three distal tarsals, and reduced pedal digit V. These make it virtually impossible to assign Megalancosaurus to Dinosauria, let alone Theropoda, despite its furcula.
Megalancosaurus a bird ancestor? Feduccia and Wild (1993) first suggested Megalancosaurus was more closely related to birds than theropods were, though the absence of other taxa in their cladogram leaves one uncertain exactly where in Archosauria they place the the Megalancosaurus+bird clade. The expanded braincase is also present in pterosaurs and coelurosaurs. Large orbits are found in those two groups as well, and many other small tetrapods. A pointed snout is also present in many maniraptoriforms, pterosaurs, most 'protorosaurs', choristoderes, thalattosaurs, and some basal lepidosauromorphs. The "large, oval preorbital fenestra" is actually the external naris. The reduced premaxillary dentition is not present in basal birds, though the authors state it "may be important, because when modern birds loose their teeth they loose the maxilla that houses them, and in the Cretaceous toothed birds, the teeth are borne only on the maxilla." Even ignoring the fact Aves does retain a maxilla, and subsequently discovered birds like omnivoropterygids and longipterygids have toothed premaxillae and toothless maxillae, mere propensity for a group to exhibit a character is not a synapomorphy. The dentary is said to be birdlike, but this is too vague to evaluate. The foramen magnum is claimed to be posteroventrally oriented, but this is based solely on the angle of the quadrate and ventral squamosal process. The latter is also true in Icarosaurus, the basal squamate Tamaulipasaurus, the basal choristodere Lazarussuchus, Cosesaurus and pterosaurs. Feduccia and Wild state "the six to seven elongate cervical vertebrae create a highly movable birdlike neck." The high mobility is caused by heterocoelous centra, which are similar to ornithuromorphs but not basal birds. Elongate cervical centra are present in coelurosaurs, 'protorosaurs' and other taxa, while birds have at least nine cervicals. Though the authors claim the "anterior limbs, without manus, are proportionately similar to those of modern birds and Archaeopteryx; and dissimilar to theropods," the radiohumeral ratio of 68% is shorter than most maniraptorans (even taxa known in 1993, e.g. Oviraptor 87%, Deinonychus 76%) and Archaeopteryx (84-96%), while modern birds usually have ratios of more than 100%. The large manual ungual flexor tubercles are said to be birdlike, but these are absent in the more basal Hypuronector, and also found in most theropods and pterosaurs. Contra Feduccia and Wild, the forelimbs minus manus are not longer than the hindlimbs minus pes, being 72% as long instead. Finally, the straplike scapula is indeed superficially like Aves in being extremely slender, bowed and having a tapered distal end. Yet basal birds lack these features, and only have scapulae as slender as pterosaurs and most theropods.
Feduccia later (1996) refers to Megalancosaurus' "tiny isodont teeth set in sockets", but they are actually subthecodont as in most basal diapsids, while isodonty is also plesiomorphic.
After the mid-1990s, Feduccia and other Birds Are Not Dinosaurs supporters seem to have reduced their emphasis on Megalancosaurus' relationship with birds. Geist and Feduccia (2000) repeat the 1993 observations, but then state "though probably not the avian ancestor, Megalancosaurus represents a chronologically and biophysically plausible model for a gliding stage through which birds must have passed." Martin (2004) incorrectly stated Megalancosaurus has a furcula, but did not explicitly link it to bird origins.
Besides those characters noted above that exclude Megalancosaurus from Theropoda, it is less similar to basal birds than even basal coelurosaurs are in many other ways. These include the absence of pleurocoels, less than five sacral vertebrae, absence of a transition point in the tail, small distal carpal I, metacarpal III longer than II, presence of manual digits IV and V, absent pubic boot, absent obturator process, obturator foramen in pubis, astragalus without tall ascending process, robust metatarsus, metatarsal I reaching tarsus, and phalanx V-1 present.
References- Calzavara, Muscio and Wild, 1980. Megalancosaurus preonensis, n. g., n. sp., a new reptile from the Norian of Friuli. Gortania. 2, 49-63.
Pinna, 1980. Drepanosaurus unguicaudatus, nuovo genere e nuova specie di lepidosauro del Trias alpino. Atti della Società Italiana di Scienze Naturali - Museo civico di Storia Naturale di Milano. 121, 181-192.
Pinna, 1987. Un nuovo esemplare giovanile di Drepanosaurus unguicaudatus del Norico di Val Preone (Udine) [A new juvenile specimen of Drepanosaurus unguicaudatus from the Norian of the Preone Valley, Udine]. Atti della Società Italiana di Scienze Naturali - Museo civico di Storia Naturale di Milano. 128, 80-84.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Feduccia and Wild, 1993. Birdlike characters in the Triassic archosaur Megalancosaurus. Naturwissenschaften. 80, 564-566.
Renesto, 1994. Megalancosaurus, a possibly arboreal archosauromorph (Reptilia) from the Upper Triassic of northern Italy. Journal of Vertebrate Paleontology. 14(1), 38-52.
Feduccia, 1996. The Origin and Evolution of Birds. Yale University Press. 420 pp.
Padian and Chiappe, 1998. The origin and early evolution of birds. Biological Reviews. 73, 1-42.
Ruben, 1998. Gliding adaptations in the Triassic archosaur Megalancosaurus. Journal of Vertebrate Paleontology. 18(3), 73A.
Geist and Feduccia, 2000. Gravity-defying behaviors: Identifying models for protoaves. American Zoologist. 40, 664-675.
Renesto, 2000. Bird-like head on a chameleon body: New specimens of the enigmatic diapsid reptile Megalancosaurus from the Late Triassic of northern Italy. Rivista Italiana di Paleontologia e Stratigrafia. 106(2), 157-180.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica. 50(6), 978-990.
Senter, 2004. Phylogeny of the Drepanosauridae (Reptilia: Diapsida). Journal of Systematic Palaeontology. 2, 257-268.
Renesto and Dalla Vecchia, 2005. The skull and lower jaw of the holotype of Megalancosaurus preonensis (Diapsida, Drepanosauridae) from the Upper Triassic of Northern Italy. Rivista Italiana di Paleontologia e Stratigrafia. 111(2), 247-257.
Renesto, Spielmann, Lucas and Spagnoli, 2010. The taxonomy and paleobiology of the Late Triassic (Carnian-Norian: Adamanian-Apachean) drepanosaurs (Diapsida: Archosauromorpha: Drepanosauromorpha). New Mexico Museum of Natural History and Science Bulletin. 46, 1-81.
Castiello, Renesto and Bennett, 2015. The role of the forelimb in prey capture in the Late Triassic reptile Megalancosaurus (Diapsida, Drepanosauromorpha). Historical Biology. DOI: 10.1080/08912963.2015.1107552
M. endennae Renesto, Spielmann, Lucas and Spagnoli, 2010
Middle Norian, Late Triassic
Zorzino Limestone Formation, Italy

Comments- MBSN 25 was first described as a juvenile specimen of Drepanosaurus (Pinna, 1980), then referred to Megalancosaurus (Renesto, 1994). Renesto et al. (2010) have recently redescribed it as a new species.
References- Pinna, 1980. Drepanosaurus unguicaudatus, nuovo genere e nuova specie di lepidosauro del Trias alpino. Atti della Società Italiana di Scienze Naturali - Museo civico di Storia Naturale di Milano. 121, 181-192.
Renesto, 1994. Megalancosaurus, a possibly arboreal archosauromorph (Reptilia) from the Upper Triassic of northern Italy. Journal of Vertebrate Paleontology. 14(1), 38-52.
Renesto, Spielmann, Lucas and Spagnoli, 2010. The taxonomy and paleobiology of the Late Triassic (Carnian-Norian: Adamanian-Apachean) drepanosaurs (Diapsida: Archosauromorpha: Drepanosauromorpha). New Mexico Museum of Natural History and Science Bulletin. 46, 1-81.

Longisquamidae Sharov, 1970
Longisquama Sharov, 1970
L. insignis Sharov, 1970
Ladinian, Middle Triassic
Madygen Formation, Kyrgyzstan
Holotype
- (PIN 2584/4) incomplete skull (~23 mm), incomplete mandibles, eight cervical vertebrae, several anterior dorsal vertebrae, dorsal ribs, scapula (12 mm), coracoid (5 mm), interclavicle? (9 mm), furcula, sternum?, humerus (13 mm), radii, ulnae (11 mm), radiale, intermedium, ulnare, pisiform, four distal carpals, metacarpals I, phalanx I-1, manual ungual I, metacarpals II (one proximal), metacarpals III (one proximal), proximal phalanx III-1, metacarpals IV (one proximal), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, manual ungual IV, metacarpals V, phalanx V-1, phalanx V-2, phalanx V-3, manual ungual V, scales, parafeathers
Paratypes- (PIN 2584/5) partial parafeather
(PIN 2584/6) two partial parafeathers
(PIN 2584/7) partial parafeather
(PIN 2584/9) six distal parafeathers
Referred- (FG 596/V/1) parafeather (Voigt et al., 2009)
(FG 596/V/2) parafeather (Voigt et al., 2009)
(FG 596/V/3) parafeather (Voigt et al., 2009)
Comments- Longisquama was originally described as a pseudosuchian, which at the time was used for all basal archosauriforms that were not parasuchians. Surprisingly few authors have addressed its relationships since, traditionally assuming it to be an archosaur(-iform) because of the supposed antorbital and mandibular fenestrae. More recently, most reviewers (e.g. Reisz and Sues, 2000) have been even more cautious and refer to it merely as a diapsid. The matter is especially difficult to resolve as the holotype only received a short original description with poor illustrations and has not been osteologically described since, with the exception of Peters' (2000) seemingly overimaginative attempt. Several characters have proven controversial and will be discussed first.
Though originally described as having an antorbital fenestra, Senter (2003) found this was caused by breaks which were not present in the main slab, a conclusion also reached by Prum (2001). Peters (2000), Martin (2004) and James and Pourtless (2009) all disagree however, believing it to have maxillary and (for the former two authors) promaxillary fenestrae as well. The latter authors provide a photograph and interpretive drawing which allow evaluation. Their supposed antorbital fossa could easily be a naris if it is indeed a hole, and if properly identified by them would be unlike other taxa in its location along the dorsal edge of the anterior half of the antorbital/maxillary fenestra. Similarly, the entire ventral edge of the supposed antorbital fenestra looks like a crack to me, including the supposed dorsal process of the maxilla, said to support an interfenestral bar. I don't see any posterior edge of the antorbital fenestra either, so the whole area could be a solid maxilla. The ventral process for the interfenestral bar identified by them is apparent, but if anything seems separated from the nasal by a suture. This would place it on the lacrimal or prefrontal, both unlike any known taxon with a maxillary fenestra. Given the above information, I agree with Prum and Senter that no antorbital fenestra is present.
An external mandibular fenestra was also originally described, though all authors including Peters, Martin and Senter now agree this was due to damage.
The mode of tooth implantation was described as acrodont by Sharov, though Martin believes it is thecodont. He provided no evidence for this in 2004, and even qualified the character with a question mark. In a later 2008 paper, Martin states the mandible had become split between slabs, and what Sharov interpreted as tooth crowns were actually entire teeth with expanded bases. Yet supposing the mandible with preserved teeth is being viewed laterally, the teeth would more probably be pleurodont since no lingual wall is apparent. Certainly the roots would be too short for thecodont teeth in any case, though they might be subthecodont if socketed.
The posterodorsal skull is preserved expanding significantly past the orbit, described as two tubercles by Sharov, and interpreted as a crest by Peters and Senter. Martin (2008) believes it is merely part of the skull roof that has become disarticulated. Either interpretation seems possible.
The clavicles were described by Sharov as fused ("concrescent") though a suture was illustrated. Senter and Martin agree they are fused to form a furcula, Peters thinks they merely overlap each other, and Unwin and Benton (2001) believe they are unfused (incorrectly saying this was Sharov's opinion).
Senter (2003, 2004), Peters (2000) and James and Pourtless have been the only authors to include the taxon in a phylogenetic analysis.
Longisquama an avicephalan? Longisquama has often been placed close to Megalancosaurus, though opinions on where this clade or grade belongs have varied wildly. Olshevsky (1991) had both as basal theropods ("basitheropod theropodomorphs"), Feduccia and Wild (1993) and others against the dinosaurian origin of birds consider both to be "avimorph archosaurs" or "avimorph thecodonts" close to the base of birds, and Peters (2006) has both close to pterosaur origins in Lepidosauromorpha. There is usually little opinion on whether the two taxa are more closely related to each other than to the clade they're viewed as ancestral to, so the discussions under each comment section below also function to examine these authors' evidence for placing Longisquama and Megalancosaurus in similar phylogenetic positions in their classification schemes.
Senter's analyses found Longisquama to emerge either in a polytomy with Coelurosauravus and Megalancosaurus (2003), or as the sister taxon of Coelurosauravus with both forming the sister taxon to simiosaurs (2004). He named this clade Avicephala, and supported it with several characters- premaxilla and dentary sharply pointed in lateral view; craniomandibular joint anteriorly placed below orbit; posterior border of skull strongly inclined posteriorly (correlated with previous character); scapula elongate; width of distal expansion of humerus <1/3 humeral length; length of manus subequal to or less than humeral length (untrue in Longisquama); metacarpal IV shorter than metacarpal III (only present in Hypuronecter among simiosaurs). Two characters (parietals posteriorly upswept, forming posterodorsal crest; parietal rim ornamented) placed Longisquama closer to Coelurosauravus in the published version. The homology of their posterior skull crests is doubtful however, since the elongate, ornamented bone in Coelurosauravus has been more recently identified as the squamosal, which is very short in Longisquama. Also, the parietal of Coelurosauravus is a midline strut, unlike the laterally placed rounded bone in Longisquama. While this is the most extensive analysis (the thesis version moreso than the published version), it still suffers from a lack of taxonomic sampling of non-archosauromorphs and a paucity of characters for such a broad taxonomic scope. Only Paleothyris, Petrolacosaurus, Youngina and Gephyrosaurus were used in the 2004 paper, while the thesis also used Icarosaurus, Askeptosaurus, Palaeopleurosaurus and two choristoderes. Yet many taxa including pterosaurs, most 'protorosaurs', choristoderes, thalattosaurs, and some basal lepidosauromorphs have pointed snouts. Icarosaurus, the basal squamate Tamaulipasaurus, the basal choristodere Lazarussuchus, Cosesaurus and pterosaurs have inclined posterior skulls. Icarosaurus, the basal squamate Huehuecuetzpalli, choristoderes, trilophosaurs, pterosaurs and most archosauriforms have slender scapulae. Araeoscelidians, kuehneosaurids, Lazarussuchus, trilophosaurs, most 'protorosaurs', pterosaurs and many archosauriforms have slender humeri. Huehuecuetzpalli, archosaurs and some 'protorosaurs' have metacarpal IV shorter than III. Renesto and Binelli (2006) added Eudimorphodon to Senter's matrix and updated his simiosaur codings, which resulted in simiosaurs moving to a position sister to archosauriforms, providing further proof the matrix size and taxon sample are both too small to provide robust results. Notably, while Longisquama stayed outside Neodiapsida sister to Coelurosauravus in their tree, they stated that further analyses may indicate the taxon is related to simiosaurians after all. Besides the characters used by Senter, Renesto and Binelli also noted the lack of cervical ribs and the tight interclavicular articulation as potential synapomorphies. Adding a wide variety of over forty diapsids to Senter's published matrix results in Longisquama clading with pterosaurs, Lazarussuchus, a clade of simiosaurs plus Tamaulipasaurus and Trilophosaurus, and Coelurosauravus (with decreasing closeness), suggesting it has too few characters to properly place taxa.
Longisquama not a saurian? Senter (2003, 2004) found Avicephala to emerge outside of Neodiapsida (and thus also Sauria and Archosauromorpha), but closer to those clades than Petrolacosaurus and Araeoscelis. The absent retroarticular process excludes it from Icarosaurus+Neodiapsida in his thesis, but this is also true in basal squamates, Askeptosaurus, Claudiosaurus, Monjurosuchus and Dinocephalosaurus. The deltopectoral crest extending less than halfway down the humerus excludes it from Neodiapsida in his thesis, but a quick survey of potentially related taxa shows this is also true of 'protorosaurs', trilophosaurs, archosaurs and pterosaurs at least. Characters in Senter's analyses which exclude Longisquama from Icarosaurus+Neodiapsida, Sauria and Archosauromorpha are- preorbital skull shorter than postorbital skull + orbit, short posterior jugal process, no ventral squamosal process, and ulnare over twice as large as radiale. However, it has nasals longer than frontals, a poorly developed olecranon, and a short metacarpal I, which are all diagnostic of neodiapsid groups in his trees. Again, note that Cosesaurus also has a short snout, short posterior jugal process, and no ventral squamosal process. Cosesaurus was included in the thesis analysis, but not the published version. While the published version has rather high bootstrap values for Avicephala, Neodispsida and Sauria (87, 97 and 73 respectively), in the thesis these are not supported by the bootstrap analysis at all. In conclusion, the placement of Longisquama sister to Coelurosauravus seems doubtful, while its placement in Avicephala and outside Sauria or Archosauromorpha is only weakly supported.
Longisquama a lepidosauromorph? Unwin et al. (2000) and Unwin and Benton (2001) both stated that the supposedly acrodont teeth and interclavicle were "more typical of lepidosaurs" than archosaurs. Yet interclavicles are present in most non-ornithodiran archosauromorphs (as noted by Jones et al., 2001), while Longisquama's tooth implantation is controversial (see above).
Peters has recently proposed a massive revision of amniote relationships where Longisquama and taxa he places close to it (pterosaurs, tanystropheids and drepanosaurs) are squamates, but his unpublished analysis is so filled with correlated characters, non-homologous states and miscodings based on his discredited photo-analyzing method that it can be ignored. Similarly ignorable is his recent (2006) magazine article on Longisquama where he claims to have found the entire posterior half of the animal (including wings, propatagia and uropatagia), a juvenile preserved next to it (also complete with patagia) and an embryo(!) inside the holotype. Nobody who has examined the actual specimen (e.g. Sharov, Jones, Martin, Prum, Senter, Voigt) has ever seen these things, leading one to conclude they are merely Peters' imagination inspired by noise in his photos and what he thinks should be there.
Longisquama a tanystropheid? Peters (2000) added Longisquama, additional taxa and sometimes additional characters to three previously published matrices (Evans, 1988; Jalil, 1997; Bennett, 1996). Longisquama emerged related to pterosaurs, Sharovipteryx and Cosesaurus in each analysis (a clade he named Fenestrasauria), which was always inside a clade containing Tanystropheus and Langobardisaurus (which he named Characiopoda, though it was previously defined as Tanystropheidae by Dilkes, 1998). However, only a few non-archosauromorphs were included in each analysis, and the accuracy of his codings have been questioned by several experts. In relation to Longisquama, Senter (2003) questioned the antorbital fenestra's presence as noted above, stated the coracoid's outline is uncertain, and that the sternum and interclavicle "are bony wisps that defy interpretation." Of Peters' fenestrasaurian characters, Longisquama lacks antorbital fenestrae, a retroarticular process, and a manual digit IV which is at least three times as long as metacarpal IV. The anterior expansion of the lacrimal could be the prefrontal instead if Coelurosauravus is used as a guide. Pterosaurs, Sharovipteryx and Cosesaurus all lack short cervicals, making this character invalid (but supporting the exclusion of Longisquama from a traditional Prolacertiformes/Protorosauria). Clavicular overlap is uncertain in Longisquama, as noted above. The coracoid, sternum and interclavicle characters (reduced posterior interclavicle process; elongate anterior interclavicle process; sternum fuses with clavicles; elongate coracoid with articulation for sternum) must be considered uncertain given Senter's statements. Of his characiopodan characters, the posterior skull is also ventrally deflected in Coelurosauravus, and posterior teeth "reduced, elongated or multi-cusped" is not a valid character since it describes an array of sometimes exclusive states. Finally, of Peters' characters joining Characiopoda with Macrocnemus, whether Longisquama is "without a primitive quadratojugal" is uncertain as the area is fragmented (though Coelurosauravus lacks the bone regardless), while Coelurosauravus shares the elongate scapula and straight, closely appressed radius and ulna. Thus Peters' characters are highly problematic, and none of those which can be verified are lacking in Coelurosauravus.
Longisquama a non-dinosaurian archosauromorph? Benton (1993) listed Longisquama as an ornithodiran without reason.
While opponents to the dinosaurian ancestry of birds have generally labeled Longisquama an archosaur, they are also generally silent as to where in that clade it belongs. However, Martin (2005) provides a cladogram which places Longisquama (and birds) outside the Crocodilia+Dinosauria clade but closer to it than prolacertids are, which would technically make Longisquama a non-archosaurian archosauromorph. This hypothesis is based only on the absent external mandibular fenestra that would exclude it from Archosauriformes, and the supposed antorbital fenestra (probably absent) that would place it closer to that taxon than Prolacerta. Needless to say, such minimal character evidence is worthless except to suggest exclusion from Archosauriformes, and Martin's other included characters (such as dinosaurs and 'thecodonts' being united by having "superpleurodont" teeth) are equally flawed.
Longisquama a theropod? Olshevsky (1991) believed Longisquama to be a basal theropod (or in his taxonomy, a basitheropod theropodomorph), but of his noted characters for that group, it only has "generally avian appearence of the skull" (vague and unlike basal theropods), carnivorous dentition (plesiomorphic for gnathostomes), furcula (controversial), relatively large forelimbs with pentadactyl manus (plesiomorphic for tetrapods and not found in basal theropods), and "featherlike scales" (which is problematic, as parafeathers do not seem to be scales or necessarily homologous with feathers, and scales are not homologous with feathers in any case).
James and Pourtless (2009) included Longisquama in a cladistic analysis mostly composed of TWG coelurosaur characters, Chiappe's bird characters and characters proposed by people disputing theropod ancestry for birds. In this analysis, Longisquama emerged as the sister group of Ornithes deep within Eumaniraptora, Maniraptora, Coelurosauria, Theropoda and Saurischia. In their trees, it is a theropod due to- straplike scapula (also present in simiosaurs, pterosaurs and to a lesser extent in Coelurosauravus; none of which were included in the matrix); an avepod based on- maxillary fenestra (absent in Longisquama); a coelurosaur based on- anterior postorbital process dorsally curved (untrue even in their own drawing); intramandibular joint poorly developed (true for most tetrapods); a maniraptoriform based on- boomerang-shaped furcula (if Peters' interpretation is correct, the concave edge would face posteroventrally, making the shape topographically unlike theropods'); a maniraptoran based on- T-shaped lacrimal (at least equally likely to be a prefrontal); maxillary and dentary teeth with constricted roots (misleadingly coded as two separate characters; if Martin 2008 is correct that the entire teeth are visible, there is no constriction); weakly developed olecranon (common in tetrapods, e.g. Cosesaurus, Langobardisaurus, Prolacerta; none of which were included in the matrix); a eumaniraptoran based on- serrationless maxillary teeth (true of almost all non-archosauriforms); less than 25 dentary teeth (very common in numerous tetrapods, e.g. simiosaurs, Langobardisaurus, Azendohsaurus; none of which were included in the matrix); interdental plates absent (true of ?all non-archosauriforms); weakly developed acromion (very common in numerous tetrapods, e.g. some simiosaurs, Protorosaurus, pterosaurs; none of which were included in the matrix); furcula (actually an avepod character, and controversial in Longisquama); an avialan based on- inflated frontals (controversial as noted above; also in Megalancosaurus and pterosaurs, which were not included in the matrix); serrationless dentary teeth (true of almost all non-archosauriforms); a metornithine based on- reduced dorsal jugal process (not obvious in Longisquama, where it could merely be broken); parietals unfused (very common in numerous tetrapods, e.g. Coelurosauravus, simiosaurs, pterosaurs; none of which were included in the matrix); all teeth with expanded roots (not actually different in birds compared to other theropods, the roots of Ichthyornis and Hesperornis illustrated by Martin and Stewart being due to cementum concretion); short coracoid (only optimized here due to Caudipteryx, this is the primitive state shared with most tetrapods and unlike birds or most paravians); poorly developed deltopectoral crest (same situation as previous character).
Thus no established characters of Longisquama are shared with theropods or subgroups yet not found in various potentially related non-archosauriforms. At best, it's possible James and Pourtless are correct in identifying the T-shaped lacrimal, low dorsal jugal process, and fused boomerang-shaped furcula if Peters got the orientation wrong. However, the lack of non-archosauriforms is a fatal flaw, as shown by Mortimer's (online, 2015) experiment adding the front half of the basal diapsid Coelurosauravus to their matrix. If this is done, Coelurosauravus and Longisquama both emerge as ornithomimosaurs (as does the pseudosuchian Effigia), proving not only is Longisquama's maniraptoran identification poorly supported, but that the characters and taxa necessary to keep non-archosaurs out of Theropoda are absent. Indeed, no manual characters were coded for any dinosaur. Characters more plesiomorphic than dinosaurs include the absent external mandibular fenestra (though reversed in some coelurosaurs and other obviously unrelated taxa), tooth implantation (whether acrodont, pleurodont or subthecodont), less than nine cervical vertebrae, interclavicle (if correctly identified), short deltopectoral crest, five phalanges on manual digit IV and four phalanges on digit V. It is thus near certainly not a dinosaur, theropod or otherwise.
Longisquama a bird ancestor? Another connection to dinosaurs has been the claim that Longisquama is related to the ancestor of birds. This originated with Sharov (1970), who believed the furcula and elongate forelimb scales were birdlike, but has more recently been popular among those arguing birds are not dinosaurs (beginning with Jones et al., 2000). Besides the furcula and forearm scales (which cannot be homologous with secondary feathers based on developmental data), Martin (2004) listed several other characters as being similar to birds- subdivided antorbital fenestra (absent as noted above), pointed snout (also in Coelurosauravus, simiosaurs, pterosaurs, most coelurosaurs, and numerous other taxa), "expanded cranium" (uncertain even if the supposed crest is really a displaced skull roof, as its three dimensional placement is unknown, as is endocranial size; also in Megalancosaurus, pterosaurs and coelurosaurs), elongate postorbital (vague and actually reduced in most birds, with even basal forms like Archaeopteryx having a shorter ventral ramus that ironically resembles Coelurosauravus more), absent mandibular fenestra (variable in basal birds and true of almost all non-archosauriforms), teeth with expanded roots (unverified as noted above), neck attaches low to skull (only true if the parietal crest is taken as part of the skull, and not actually true in basal birds like Archaeopteryx), strap-like scapula (also present in neotheropods, simiosaurs, pterosaurs and to a lesser extent in Coelurosauravus), elongate manus (also in pterosaurs, coelurosaurs and many other taxa), elongate penultimate manual phalanges (untrue in digit I, and only homologous in Martin's view in digit IV; also present in Coelurosauravus, simiosaurs, pterosaurs and theropods), and feathers.
The supposed feather homologs of Longisquama have generated the most research since Jones et al. first redescribed them, and have since been more accurately described by Voigt et al. (2009). Jones et al. described a number of similarities to feathers, and it seems fitting to use Feduccia's (2002) term 'parafeather' for the structures. Though Haubold and Buffetaut (1987) proposed the parafeathers were paired and could be horizontally extended as gliding surfaces (a claim followed by Martin), there is no evidence of this and Voigt et al. noted any such 'thoracic wing' would be compromised by having the aerodynamic surfaces so distally placed. The cylindrical, tapered base is similar to follicular structures like feathers, though Voigt et al. noted some scales such as those on iguanid dorsal frills have this characteristic as well. However, the supposed transverse partitions homologized to avian pulp caps by Jones et al. (and claimed to be pedal phalanges by Peters, 2006) are actually transverse ridges on both sides of the parafeather's middle lobe. The supposed calamus walls surrounding them are the anterior and posterior lobes, which lack ridges basally (Voigt et al., 2009). While parafeathers look roughly feather-like distally in having a central shaft and surrounding vane, the actual structure is quite different. Instead of a hollow rachis and separate barbs to form the vane, Longisquama has a pair of membranes which join at their edges (Reisz and Sues, 2000) and enclose two longitudinal lobes distally, as the posterior lobe tapers out before the vane-like expansion. The shaft analog is a continuation of the boundary between the anterior and middle lobes (so is not even continuous with the basal 'calamus' as identified by Jones et al.), while the supposed barbs never separate even at the parafeather's tip and often merge (blamed on taphonomy by Jones et al.). Instead, the 'barbs' are transverse ridges in the continuous membrane. There seems to be an outer sheath on the base of each parafeather, which was homologized by Jones et al. with the sheaths on avian feathers. Resemblences to feathers thus seem limited to the cylindrical and tapered base and basal sheath. While parafeathers may be homologous to feathers at the level of the follicle, they are no more similar than the stage 1 feathers of Tianyulong, Kulindadromeus and basal coelurosaurs, the quills of Psittacosaurus or the pycnofibres of pterosaurs.
More recently, James and Pourtless (2009) have recovered Longisquama as the sister to Ornithes in their flawed cladistic analysis described above. This is actually more closely related to birds than hypothesized by Martin and Feduccia in their latest papers, as the latter have oviraptorosaurs and deinonychosaurs closer to Aves albeit outside Dinosauria. The characters listed under "Longisquama a theropod?" above for Maniraptora through Metornithes could thus also be used to defend relationships with birds in Feduccia's or Martin's view. In addition to those, the characters James and Pourtless recovered as shared with Ornithes are- scapula parallel to dorsal column (very untrue in Longisquama); ossified sternum absent (optimized in their topology this way due to alvarezsaurids and Caudipteryx being next most basal, but also true of most theropods, as well as many non-archosaurs such as Coelurosauravus and Protorosaurus).
Longisquama does not share any characters with birds not found in basal coelurosaurs, and can be excluded from Tetanurae based on numerous characters such as maxillary teeth extending posteriorly under the orbit, lacking an enlarged distal carpal I+II, and having phalanges on manual digit IV and having digit V, in addition to the non-dinosaurian characters noted in the previous section.
References- Sharov, 1970. Svoyeobrazaya reptiliya iz nizhnego triasa Fergany. Paleontologicheskii Zhurnal. 1, 127-130.
Sharov, 1970. An unusual reptile from the Lower Triassic of Fergana. Paleontological Journal. 1, 127-130.
Haubold and Buffetaut, 1987. Une novelle interprétation de Longisquama insignis, reptile énigmatique du Trias supérieur d'Asie centrale. Comptes Rendus Académie des Sciences du Paris. 305, 65-70.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Benton, 1993. Reptilia. in Benton (ed). The Fossil Record 2. London. 681-715.
Jones, 2000. New insight into the morphology and physiology of Mesozoic archosaurs. PhD thesis, Oregon State University. 87 pp.
Jones, Ruben, Martin, Kurochkin, Feduccia, Maderson, Hillenius, Geist and Alifanov, 2000. Nonavian feathers in a Late Triassic archosaur. Science. 288(5474), 2202-2205.
Peters, 2000. A reexamination of four prolacertiforms with implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia. 106(3), 293-336.
Reisz and Sues, 2000. The "feathers" of Longisquama. Nature. 408(6811), 428.
Unwin, Alifanov and Benton, 2000. Enigmatic small reptiles from the Middle-Late Triassic of Kirgizstan. In Benton, Shishkin, Unwin and Kurochkin (eds). The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press, Cambridge. 177-186.
Jones, Ruben, Maderson and Martin, 2001. Longisquama fossil and feather morphology. Science. 291(5510), 1901-1902.
Prum, 2001. Longisquama fossil and feather morphology. Science. 291(5510), 1899-1900.
Reisz and Sues, 2001. Longisquama does not have feathers. Journal of Vertebrate Paleontology. 21(3), 92A.
Unwin and Benton, 2001. Longisquama fossil and feather morphology. Science. 291(5510), 1900-1901.
Feduccia, 2002. Birds are dinosaurs: Simple answer to a complex problem. The Auk. 119(4), 1187-1201.
Senter, 2003. Taxonomic sampling artifacts and the phylogenetic position of Aves. Unpublished PhD thesis. Northern Illinois University. 147 pp.
Senter, 2004. Phylogeny of the Drepanosauridae (Reptilia: Diapsida). Journal of Systematic Palaeontology. 2, 257-268.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica. 50(6), 978-990.
Peters, 2006. The other half of Longisquama. Prehistoric Times. 75, 10-11.
Renesto and Binelli, 2006. Vallesaurus cenensis Wild 1991, a drepanosaurid (Reptilia, Diapsida) from the Late Triassic of northern Italy. Rivista Italiana di Paleontologia e Stratigrafia. 112, 77-94.
Martin, 2008. Origins of avian flight- a new perspective. Oryctos. 7, 45-54.
Buchwitz, Voigt and Fischer, 2009. Dorsal appendages of Longisquama reconsidered: Aspects of development and the link to the evolution of filamentous integumentary structures. Journal of Vertebrate Paleontology. 29(3), 72A.
James and Pourtless, 2009. Cladistics and the origin of birds: A review and two new analyses. Ornithological Monographs. 66, 78 pp.
Voigt, Buchwitz, Fischer, Krause and Georgi, 2009. Feather-like development of Triassic diapsid skin appendages. Naturwissenschaften. 96, 81-86.
Mortimer, online 2015. http://theropoddatabase.blogspot.com/2015/01/bandit-cladogram-evaluated-james-and.html

Thecodontia Owen, 1859
Definition- (Protorosaurus speneri + Thecodontosaurus diagnosticus) (modified from Kischlat, 2000)
= Protorosauria sensu Merck, 1997
Definition- (Protorosaurus speneri + Tanystropheus conspicuus) (modified)
= Archosauromorpha sensu Gauthier et al., 2004
Definition- (Rhynchosaurus articeps + Protorosaurus speneri + Caiman crocodilus)

unnamed possible thecodont (Chatterjee, 1987)
Carnian, Late Triassic
Maleri Formation, India
Material
- (ISI R 306) cervical vertebra (~27 mm)
Comments- This material is part of the Alwalkeria holotype as described by Chatterjee (1987), but Rauhut and Remes (2005) found it was similar to Malerisaurus and Pamelaria. Some of the dorsal vertebrae may belong to this taxon also.
References- Chatterjee, 1987. A new theropod dinosaur from India with remarks on the Gondwana-Laurasia connection in the Late Triassic. Geophysical Monograph. 41, 183-189.
Remes and Rauhut, 2005. The oldest Indian dinosaur Alwalkeria maleriensis Chatterjee revised: a chimera including remains of a basal saurischian. in Kellner, Henriques and Rodrigues (eds). II Congresso Latino-Americano de Paleontologia de Vertebrados, Boletim de Resumos.Museu Nacional, Rio de Janeiro. 218.

Protorosauria Huxley, 1871
Definition- (Protorosaurus speneri <- Crocodylus niloticus) (modified from Kischlat, 2000)
Other definitions- (Protorosaurus speneri + Tanystropheus conspicuus) (modified from Merck, 1997)

Parathecodontia Gow, 1975
Definition- (Prolacerta broomi + Tanystropheus conspicuus) (modified from Merck, 1997)
?= Tapinoplatia Peters, 2000
Definition- (Macrocnemus bassanii + Preondactylus buffarinii) (modified from Peters, 2000)
?= Characiopoda Peters, 2000
Definition- (Tanystropheus conspicuus + Preondactylus buffarinii) (modified from Peters, 2000)
?= Fenestrasauria Peters, 2000
Definition- (Cosesaurus aviceps + Preondactylus buffarinii) (modified from Peters, 2000)

Tanystropheidae Gervais, 1858 or 1859
= Tribelesodontidae Nopcsa, 1922
= Tribelesodontoidea Nopcsa, 1922 sensu Nopcsa, 1928
= Tribelesodontia Nopcsa, 1923
= Macroscelosauridae Kuhn, 1935
Definition- (Tanystropheus conspicuus + Macrocnemus bassanii + Langobardisaurus pandolfii) (modified from Dilkes, 1998)
Other definitions- (Tanystropheus conspicuus + Tanytrachelos ahynis) (modified after Merck, 1997)
Comments- Tribelesodontidae was named by Nopcsa (1922) as a basal family of pterosaurs including Tribelesodon and Rhabdopelix.
References- Nopcsa, 1922. Neubeschreibung des Trias-Pterosauriers Tribelesodon. Paläontologische Zeitschrift. 5(2), 161-181.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Nopcsa, 1923. Die familien der Reptilien.
Kuhn, 1935. Rhynchocephalia (Eosuchia). Fossilium Catalogus I: Animalia. 71, 1-38.

Cosesaurus

Gwyneddosaurus Bock, 1945
= Tanytrachelos Olsen, 1979
G. erici Bock, 1945
= Tanytrachelos ahynis Olsen, 1979
Late Carnian, Late Triassic
Gwynedd or North Wales Member of Lockatong Formation, Pennsylvania, US
Holotype
- (ANSP 15072) braincase or palatal element?, axis (8 mm), two anterior cervical neural arches (~8 mm), three posterior cervicals (7.5 mm), cervical vertebra, cervical ribs, dorsal vertebra, at least 11 dorsal ribs, gastralia, distal caudal vertebra (4 mm), three chevrons, heterotopic bone?, neural arch, scapulocoracoids, clavicles, interclavicle, radius (9 mm), proximal ulna (~9 mm), incomplete ilium (10 mm), pubis?, ischium?, femur (23 mm), proximal fibula, three phalanges (2.5, 4 mm), fragments, coelacanth fragments?
Paratype- ?(ANSP coll.) femur (36.5 mm), tibia (30 mm)
Comments- Gwyneddosaurus erici was described by Bock (1945) based on a disarticulated specimen, in which he labeled the bones he identified with numbers 1-51. Bock provisionally placed the reptile in Theropoda and furthermore in Podokesauridae within Coelurosauria. This was based on the supposedly short forelimbs and gracile (perhaps hollow) postcrania, with the Triassic age probably responsible for the familial identification. The procoelous vertebrae, distally expanded presacral neural spines, and differently shaped pectoral girdle were all seen as different from Podokesaurus though. Bock also referred to "several leg bones, similar to those of our specimen" from the same locality though he did not describe these. Huene (1948) examined photographs of the material from before it was prepared by Bock, leading him to reidentify almost every element. The supposed pterygoid (16), and two other cranial elements were reidentified as a possible squamosal, basisphenoid or parasphenoid (15), and cervical vertebra (13) respectively. The supposed dorsal vertebrae (3 and 4) were reidentified as posterior cervicals, the cervical ribs (39) as chevrons, and one of the dorsal ribs (45) as an ulna. The scapula (18) and coracoid (17) were each actually a complete scapulocoracoid, and the bone Bock identified as a fused radius and ulna (22) was not mentioned but is far too short to be such. One of the ilia is actualy a probable ischium (2), and a supposed tarsal (25, perhaps meant as metatarsal) is a radius. Huene also described and figured a femur and tibia larger than the holotype which is seemingly the additional specimen noted by Bock. He felt these were possibly referrable to Gwyneddosaurus, as the femur "may be structurally identical." Huene assigned the taxon to Protorosauria based on cervicals similar to Macrocnemus and Microcnemus, holocephalous dorsal ribs, elongate and only slightly curved gastralia as in Macrocnemus and Tanystropheus, extremely short scapula with one broad and one rounded corner as in Adelosaurus, clavicle similar to Macrocnemus, ilium with small preacetabular and large postacetabular process as in Protorosaurus, and distally narrowing paratype femur as in Macrocnemus. This was followed by Camp et al. (1953) and Romer (1966), who placed it in Protorosauridae and ?Prolacertidae respectively. However, the cervical and clavicle similarities were never specified, the ilial proportions are plesiomorphic for amniotes, Adelosaurus has been more recently recognized as being of Youngina-grade, holocephalous dorsals are found in numerous taxa, gastralial morphology has not been shown to be diagnostic for tanystropheids, and the holotype femur seems not to distally narrow. Huene later (1956) placed Gwyneddosaurus in Askeptosauridae, but Askeptosaurus has shorter amphicoelous presacrals, a shorter scapula, rod-like interclavicle, no preactabular process, a longer postacetabular process, and much more robust limb elements with poorly developed features. Perhaps it was a typo.
Steel (1970) followed Bock in placing Gwyneddosaurus in Theropoda, Romer (1976) listed it as a podokesaurid coelurosaur, and Welles (1984) placed it in Theropoda incertae sedis (though stating "it certainly is not even remotely related to Dilophosaurus"). While Huene never explicitly stated reasons to exclude the taxon from Theropoda, they include- procoelous cervical centra, no cervical pleurocoels, holocephalous dorsal ribs, short scapular blade, unfused clavicles, ossified interclavicle, short preacetabular process, femoral head not inturned, internal trochanter and intertrochanteric fossa present. Furthermore, the supposedly short forelimb elements were incorrectly identified, and numerous reptiles have gracile postcrania which can appear hollowed.
Olsen first (1979) stated that Gwyneddosaurus had an emargnate scapulocoracoid (though this is untrue), so could be a lizard, but was not closely related to his new tanystropheid Tanytrachelos. He did state it was nearly identical to some elements of Rhabdopelix though, and would be redescribed in the future. Olsen later (1980) thought it and Rhabdopelix could be senior synonyms of Tanytrachelos, as some elements could not be distinguished from it. As other elements differ between the taxa, Olsen thought Gwyneddosaurus should be a nomen dubium. Yet chimaerical status does not make a taxon a nomen dubium. Olsen and Baird (1986) considered Gwyneddosaurus to be a chimaera of coelacanth and possible Tanytrachelos bones. Similarly, Olsen and Flynn (1989) thought it to be a chimaera of Tanytrachelos and possible coelacanth fragments. Specifically, they stated "the large, quadranglular and keeled interclavicle characteristic of Tanytrachelos is present as are procoelous vertebrae." Instead of sinking Tanytrachelos into Gwyneddosaurus, Olsen and Flynn incorrectly consider the latter a nomen dubium because "there is significant room for doubt- certainly at the species level it is indeterminate." This statement is meaningless though as Tanytrachelos only has one recognized species, and the historical recognization of genus- and species-level characters for monospecific taxa is outdated. Indeed, when "at least part of Gwyneddosaurus more assuredly is Tanytrachelos", the former cannot be a nomen dubium. An ICZN petition would be necessary to sink Gwyneddosaurus, and since this has not been made yet it is more proper to make Gwyneddosaurus a senior synonym of Tanytrachelos. The identity of Rhabdopelix deserves further scrutiny, though it does include procoelous vertebrae. Whether the referred hindlimb is correctly referred is uncertain, as there are no proposed hindlimb apomorphies of Gwyneddosaurus, but is does seem to be tanystropheid based on the distal narrowing and is similar in size to large Tanytrachelos specimens. Spamer et al. (1995) placed Gwyneddosaurus in Tanystropheidae.
Unfortunately, comparison between the two taxa is difficult, as the pre-preparation photos of Gwyneddosaurus remain unpublished (though they are archived at the ANSP), while Tanytrachelos has only received a preliminary description. Bone 2 is not mentioned by either Bock or Huene but may be another posterior cervical. I think Huene switched bones 5 and 6 in his description, with 5 being the axis and 6 being a neural arch in dorsal view. Element 14 was believed to be an atlas or cranial bone by Bock, but is certainly too large for the former. Olsen and Baird's description of the interclavicle and comparison with Tanytrachelos suggests it is element 15, identified as a cranial bone by Bock and a para- or basisphenoid by Huene. Element 16 was identified as a possible squamosal by Huene, but does not match that element in Tanytrachelos and may be from the braincase or palate instead. Bock's supposed fused radius and ulna (22), may be an anterior dorsal rib if element 22 refers to the elongate curved bone and not the posterior cervical(?) the distal end lies over. Elements 23 and 24, identified as ischia by Bock, seem to be posterior dorsal ribs. Based on the identification as congeneric with Tanytrachelos, the two metatarsals or phalanges identified by Huene (31 and another unlabeled) are definitely the latter. The radius (25) and perhaps the ulna (45) as identified by Huene may actually be metatarsals then. A supposed (meta?)tarsal identified by Bock (27) seems more likely to be a posterior dorsal rib. Interestingly, curved triangular element 29 (called a rib by Bock), may be a heterotopic bone as found in some Tanytrachelos individuals. All of the "foot bones (32-38) which probably are toes" are anterior cervical ribs or gastralia as is at least one of the supposed manus bones (41). The other proposed manual elements (40 and 42) are very small and could be distal gastralia or fish bones. Notably, despite Olsen's statements Gwyneddosaurus contains coelacanth material (presumably of the common and contemporaneous Osteopleurus newarki), which material this refers to has never been specified nor has the referral been justified.
References- Bock, 1945. A new small reptile from the Triassic of Pennsylvania. Notulae Naturae of the Academy of Natural Sciences of Philadelphia. 154, 1-8.
Huene, 1948. Notes on Gwyneddosaurus. American Journal of Science. 246, 208-213.
Camp, Welles and Green, 1953. Bibliography of Fossil Vertebrates 1934-1938. The Geological Society of America, GSA Special Paper 42.
Huene, 1956. Palaeontologie und Phylogenie der Niederen Tetrapoden. Jena. 716 pp.
Romer, 1966. Vertebrate Paleontology. Chicago University Press. 468 pp.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie. Gustav Fischer Verlag, Stuttgart. 1-87.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Olsen, 1979. New aquatic eosuchian from the Newark Supergroup (Late Triassic-Early Jurassic) of North Carolina and Virginia. Postilla. 176, 14 pp.
Olsen, 1980. Comparison of the vertebrate assemblages from the Newark and Hartford basins (Early Mesozoic, Newark Supergroup) of eastern North America. in Jacobs (ed.). Aspects of Vertebrate History. Flagstaff, Museum of Northern Arizona Press. 35-53.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica Abteilung A. 185, 85-180.
Olsen and Baird, 1986. The ichnogenus Atreipus and its significance for Triassic biostratigraphy. in Padian (ed.). In the Beginning of the Age of Dinosaurs: Faunal Change Across the Triassic-Jurassic Boundary. Cambridge: Cambridge University Press. 61-87.
Olsen and Flynn, 1989. Field guide to the vertebrate paleontology of Late Triassic rocks in the southwestern Newark Basin (Newark Supergroup, New Jersey and Pennsylvania). The Mosasaur. 4, 1-35.
Spamer, Daeschler and Vostreys-Shapiro, 1995. A study of fossil vertebrate types in the Academy of Natural Sciences of Philadelphia: Taxonomic, systematic and historical perspectives. The Academy of Natural Sciences of Philadelphia. Special Publication 16, 435 pp.

Tanystropheus Meyer, 1852
= "Macroscelosaurus" Muenster vide Meyer, 1852
= Tribelesodon Bassani, 1886
= Chelyzoon Huene, 1902
= Pectenosaurus Huene, 1902
= Procerosaurus Huene, 1902
= Thecodontosaurus "latespinatus" Huene, 1905
= Thecodontosaurus latespinatus Huene, 1908
Comments- The misspelling Tanystrophaeus was very common in early works, perhaps originating with Owen (1860) and being used consistantly by Cope and Huene.
Tanystropheus conspicuus was named for cervical vertebrae originally identified as caudals. Owen (1860) thought they might belong to the tail of a contemporaneous sauropterygian such as Nothosaurus, Simosaurus or Placodus. Cope (1887b) referred the genus to Theropoda (his Goniopoda) based on coelophysid remains he had previously described as new Coelurus species, but which he moved to Tanystropheus (T. bauri, T. longicollis and the new T. willistoni) that year. This was followed by Lydekker (1889), Zittel (1890) and Haeckel (1895) who both placed it in Coeluridae. Even after Cope's species were made the types of his new genus Coelophysis in 1889, Tanystropheus remained in Coeluridae (e.g. Huene 1902, 1908). Huene later (1914, 1926) moved it to Podokesauridae once he separated the Triassic coelurosaurs into that family. Nopcsa (1928) listed it as a podokesaurine podokesaurid. The comparisons to Coelurus and coelophysid distal caudal vertebrae were due to all three being elongate, amphicoelous, thin-walled, with low neural spines and no transverse processes. Of course articulated Tanystropheus material now shows these are cervicals, and Huene (1908) even identified parapophyses but thought the majority of evidence indicated they were caudals. Unlike theropods, they lack pleurocoels, angled articular facets, and laminae. Once Tanystropheus was reinterpreted by Peyer (1930, 1931) based on more complete material of T. longobardicus, Huene (1931) agreed in removing it from Dinosauria.
Not Tanystropheus- Cope (1887a) described Coelurus bauri and C. longicollis from the Petrified Forest Formation of New Mexico, and later that year (1887b) moved them to Tanystropheus along with his new species T. willistoni from the same locality. This was due to the elongate amphicoelous caudal centra, as Tanystropheus conspicuus cervicals were thought to be caudals at the time. Cope later (1889) erected the genus Coelophysis for the three species, because he incorrectly thought Tanystropheus' vertebrae lacked neural canals. Coelophysis bauri has since received a more complete neotype and is a coelophysid theropod, but Cope's material and other species have proven harder to diagnose within Coelophysidae. Huene (1908) named Tanystropheus posthumus, which was later renamed Tanystrosuchus and seems to be an archosaur caudal vertebra.
References- Meyer, 1852. Die saurier des Muschelkalkes mit ruecksicht auf die saurier aus Buntem Sanstein und Keuper. in Zur fauna der Vorwelt, zweite Abteilung 42. 167 pp.
Owen, 1860. Palaeontology, or a Systematic Summary of Extinct Animals and their Geological Relations. Second Edition. Adam and Charles Black, Edinburgh. 463 pp.
Bassani, 1886. Sui Fossili e sull’ età degli schisti bituminosi triasici di Besano in Lombardia. Atti della Società Italiana di Scienze Naturali. 19, 15-72.
Cope, 1887a. The dinosaurian genus Coelurus. American Naturalist. 21, 367-369.
Cope, 1887b. A contribution to the history of the Vertebrata of the Trias of North America. Proceedings of the American Philosophical Society. 24(126), 209-228.
Cope, 1889. On a new genus of Triassic Dinosauria. American Naturalist. 23, 626.
Lydekker, 1889. On a coelurid dinosaur from the Wealden. Geological Magazine. 6, 119-121.
Zittel, 1890. Handbuch der Palaeontologie. Volume III. Vertebrata (Pisces, Amphibia, Reptilia, Aves). 900 pp.
Haeckel, 1895. Systematische Phylogenie der Wirbelthiere: (Vertebrata). 660 pp.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen. 6, 1-84.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs of the European Triassic formations with consideration of occurrences outside Europe]. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Huene, 1914a. Das natürliche System der Saurischia [The systematics of the Saurischia]. Centralblatt für Mineralogie, Geologie und Paläontologie. 1914, 154-158.
Edinger, 1923. Eine Bemerkung über Tanystropheus. Senckenbergiana. 5, 143-144.
Edinger, 1924. Rückenmark im Wirbelkörper. Anatomischer Anzeiger. 57, 515-519.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Peyer, 1930. Tanystropheus longobardicus Bass. sp. Centralblatt für Mineralogie Abteilung B. 1930, 336-337.
Edinger, 1931. Tanystropheus aufgeklärt. Naturwissenschaften, 19, 846-847.
Wild, 1971. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). PhD thesis. Universitat Zurich.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). Schweizerische Palaontologische Abhandlungen. 95, 1-162.
Wild, 1975. Der Giraffenhals-Saurier. Naturwissenschaften. 62, 149-153.
Wild, 1980. Die Triasfauna der Tessiner Kalkalpen. XXIV. Neue Funde von Tanystropheus (Reptilia, Squamata). Abhandlungen der schweizerischen paläontologischen Gesellschaft. 102, 1-31.
Wild, 1985. Tanystropheus and its importance for stratigraphy. Mémoires de la Société géologique de France. 139, 201-206.
Wild, 1987. An example of biological reasons for exctinction: Tanystropheus (Reptilia, Squamata). Mémoires de la Société géologique de France. 150, 37-44.
Vickers-Rich, Rich, Rieppel, Thulborn and McClure, 1999. A Middle Triassic vertebrate fauna from the Jilh Formation, Saudi Arabia. Neues Jahrbuch für Geologie und Paläontologie. 213(2), 201-232.
Dalla Vecchia, 2000. Tanystropheus (Archosauromorpha, Prolacertiformes) remains from the Triassic of Northern Friuli (NE Italy). Revista Italiana di Paleontologia e Stratigrafia. 106(2), 135-140.
Li, 2007. A juvenile Tanystropheus sp. (Protorosauria, Tanystropheidae) from the Middle Triassic of Guizhou, China. Vertebrata PalAsiatica. 45, 37-42.
T. conspicuus Meyer, 1852
= "Macroscelosaurus" Muenster vide Meyer, 1852
= Chelyzoon blezinger Huene, 1902
= Chelyzoon latum Juene, 1902
= Pectenosaurus strunzi Huene, 1902
= Procerosaurus cruralis Huene, 1902
= "Macroscelosaurus" conspicuus (Meyer, 1852) Kuhn, 1937
Anisian, Middle Triassic
Upper Muschelkalk, Germany

Syntypes- (material of "Macroscelosaurus") cervical vertebrae
Referred- (Blezinger coll.; holotype of Procerosaurus cruralis) incomplete femur (~415 mm) (Huene, 1902)
(Blezinger coll.) cervical vertebrae (Huene, 1908)
(GPIT coll.) cervical vertebrae (Huene, 1908)
(SMNS coll.) cervical vertebrae (Huene, 1908)
(Strasbourg University coll.) cervical vertebra (Huene, 1908)
(Strunz coll.) cervical vertebrae (Huene, 1908)
Comments- Huene (1908) thought Zanclodon schuetzii (based on a tooth) might belong to T. conspicuus, but Tanystropheus differs in having conical, unserrated teeth with longitudinal crests.
"Macroscelosaurus"- In the original desciption of Tanystropheus, Meyer (1852; sometimes cited as 1847 to/or 1855) stated that Muenster thought the type cervical vertebrae were reptilian limb bones and had named them Macroscelosaurus. No species name was given, and references to "Macroscelosaurus janseni" are misspellings of the therapsid Macroscelesaurus janseni (inappropriately renamed Haughtoniscus by Kuhn, who thought the name was too similar to "Macroscelosaurus"). Kuhn (1934) used "Macroscelosaurus" as a senior synonym of Tanystropheus, citing a "Muenster, 1834" reference which subsequent authors have not been able to verify (Muenster's 1834 "Vorläufige Nachricht über einige neue Reptilien im Muschelkalke von Baiern" being about Nothosaurus mirabilis instead). 1830 is another year that has often been cited, though Wild (1976) stated that a search of all known works of Muenster had proved fruitless. Wild (1975, 1976) noted that since "Macroscelosaurus" was first published as a junior synonym, it is invalid and petitioned the ICZN to have it officially suppressed. This was approved by the ICZN in 1981 (Melville, 1981). Huene (e.g. 1902, 1908) and Nopcsa (1901) listed "Macroscelosaurus" as a junior synonym of Tanystropheus in Coeluridae. Piveteau (1955) used "Macroscelosaurus" as a valid genus of tanystropheid non-synonymous with Tanystropheus, which is not possible as they are based on the same material. Similarly, Kuhn's (1935) Macroscelosauridae is an objective junior synonym of Tanystropheidae, even if "Macroscelosaurus" were the valid name.
Procerosaurus- Procerosaurus cruralis was described by Huene (1902) as a eusuchian most similar to gavials, though in a rather confusing paragraph which also states the lack of a fourth trochanter seems to prove it is dinosaurian. He ironically stated some similarity could exist with Tanystropheus longicollis (now recognized as coelophysid material, in this case AMNH 2704) based on Cope's description, but examination of the specimen itself showed they weren't similar after all. In 1910, Huene stated it was similar to Saltopus and that he never regarded it as crocodilian (though he did not synonymize it with Tanystropheus yet, contra Olshevsky, 1991). Huene later (1914a) placed Procerosaurus in parentheses under Tanystropheus, perhaps indicating he had synonymized them by this time, and included both as a central coelurosaur lineage within Podokesauridae. In the same year (1914b), he described additional supposed coelurosaur femora from the slightly younger Lower Muschelkalk as being most similar to Procerosaurus. However, Huene still had Procerosaurus listed as a valid genus of podokesaurid in 1920, as did Nopcsa in 1928 (in the subfamily Podokesaurinae). Kuhn (1939) listed it as an indeterminate coelurosaur, and stated it was a possible synapsid ("theromorphe"). General reference works such as Zittel (1927), Romer (1956) and Kuhn (1969) used it as a synonym of Tanystropheus, which was confirmed by Wild's (1973) osteology of the latter genus. It was synonymized with T. conspicuus by Wild, probably due to provenence. Notably, Huene's favorable comparisons to theropods used taxa which are no longer thought to be dinosaurian (Saltopus, the Lower Muschelkalk femora), while his comparison to Tanystropheus was based on a theropod femur as no appendicular elements were yet referred to that genus. The femur of Procerosaurus differs from Triassic theropods in lacking a fourth trochanter and an ectocondylar tuber. While Olshevsky stated it has sometimes been classified as an ornithischian, this seems to only be true of Procerosaurus Fritsch, 1905. This was a new genus for Iguanodon exogirarum, a Cretaceous taxon named by Fritsch in 1878 for a possible tibia. Because Fritsch's genus was preoccupied by Huene's, Olshevsky renamed it Ponerosteus in 2000.
References- Meyer, 1852. Die saurier des Muschelkalkes mit ruecksicht auf die saurier aus Buntem Sanstein und Keuper. In Zur fauna der Vorwelt, zweite Abteilung 42. 167 pp.
Fritsch, 1878. Die Reptilien und Fische der böhmischen Kreideformation. Prague. 46 pp.
Nopcsa, 1901. Synopsis und Abstammung der Dinosaurier. Földtani Közlöny. 31, 247-288.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen. 6, 1-84.
Fritsch, 1905. Synopsis der Saurier der böhm. Kreideformation [Synopsis of the saurians of the Bohemian Cretaceous formation]. Sitzungsberichte der königlich-böhmischen Gesellschaft der Wissenschaften, II Classe. 1905(8), 1-7.
Huene, 1910. Ein primitiver Dinosaurier aus der mittleren Trias von Elgin. Geologische und Paläontologische Abhandlungen, 8, 315-322.
Huene, 1914a. Das natürliche System der Saurischia [The systematics of the Saurischia]. Centralblatt für Mineralogie, Geologie und Paläontologie. 1914, 154-158.
Huene, 1914b. Coelurosaurier Reste aus dem Unteren Muschelkalk. Centralblatt für Mineralogie, Geologie und Palaontologie. 1914, 670-672.
Broili, 1915. Beobachtungen über Tanystropheus conspicuus. Neues Jahrbuch für Minerologie, Geologie und Paläontologie. 2, 51-62.
Huene, 1920. Stammesgeschichtliche Ergebnisse einiger Untersuchungen an Trias-Reptilien [Phylogenetic results of some investigations of Triassic reptiles]. Zeitschrift für Induktive Abstammungsund Vererbungslehre. 24, 159-163. .
Zittel, 1927. Textbook of Paleontology. Macmillan Co. Limited. Vol. I, 839 pp.
Nopcsa, 1928. The genera of reptiles. Palaeobiologica. 1, 163-188.
Peyer, 1931. Tanystropheus longobardicus Bass sp. Die Triasfauna der Tessiner Kalkalpen. Abhandlungen Schweizerische Paläontologie Gesellschaft. 50, 5-110.
Huene, 1931. Über Tanystropheus und verwandte Formen. Neues Jahrbuch für Geologie und Paläontologie, Beilageband 67 Abteilung B. 65-86.
Kuhn, 1934. Sauropterygia. Fossilium Catalogus I: Animalia. 69, 127 pp.
Kuhn, 1935. Rhynchocephalia (Eosuchia). Fossilium Catalogus I: Animalia. 71, 38 pp.
Kuhn, 1937. Die fossilen Reptilien. Berlin: Gebrueder Borntrager. 121 pp.
Kuhn, 1939. Saurischia. In Fossilium Catalogus I. Animalia. 87. 124 pp.
Adam, 1953. Ein Tanystropheus-Fund aus dem Hauptmuschelkalk bei Schlo Stetten (Kreis Kunzelsau). Neues Jahrbuch fur Geologie und Palaeontologie. 1, 40-43.
Piveteau, 1955. Traite de paleontologie. V. Amphibiens, reptiles, oiseaux. Masson, Paris. 1113 pp.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Kuhn, 1969. Proganosauria, Bolosauria, Placodontia, Araeoscelidia, Trilophosauria, Weigeltisauria, Millerosauria, Rhynchocephalia, Protorosauria. Handbuch der Palahoerpetologie. 9, 74 pp.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). Schweizerische Palaontologische Abhandlungen. 95, 1-162.
Wild, 1975. Tanystropheus H. v. Meyer, 1855 (Reptilia): Request for conservation under the plenary powers. Z.N.(S.) 2084. Bulletin of Zoological Nomenclature.32, 124-126.
Wild, 1976. Tanystropheus H. von Meyer, [1852] (Reptilia): Revised request for conservation under the plenary powers. Z.N.(S.) 2084. Bulletin of Zoological Nomenclature. 33, 124-126.
Melville, 1981. Opinion 1186. Tanystropheus H. von Meyer, [1852] (Reptilia) conserved. Bulletin of Zoological Nomenclature. 38(3), 188-190.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Olshevsky, 2000. An Annotated Checklist of Dinosaur Species by Continent. Mesozoic Meanderings. 3, 157 pp.
T. antiquus Huene, 1908
= Tanystropheus "antiquus" Huene, 1905
= Thecodontosaurus "primus" Huene, 1905
= Thecodontosaurus primus Huene, 1908
= Tanystropheus primus (Huene, 1908) Huene, 1932
= "Macroscelosaurus" antiquus (Huene, 1908) Kuhn, 1937
Early Anisian, Middle Triassic
Lower Gogolin Formation of the Lower Muschelkalk, Poland
Lectotyp
e- (SMNS 10110; Huene's No. 7) cervical vertebra (63.1 mm)
Paralectotypes- (Vienna Natural History Museum coll.; Huene's No. 10) cervical vertebra (47 mm)
(SMNS coll.; Huene's No. 1; lost) cervical vertebra (93 mm)
(SMNS coll.; Huene's No. 2; lost) cervical vertebra (83 mm)
(SMNS coll.; Huene's No. 3; lost) cervical vertebra (74 mm)
(SMNS coll.; Huene's No. 4; lost) cervical vertebra (76 mm)
(SMNS coll.; Huene's No. 5; lost) cervical vertebra (~75 mm)
(SMNS coll.; Huene's No. 6; lost) cervical vertebra (37 mm)
(SMNS coll.; Huene's No. 8; lost) cervical vertebra (62 mm)
(SMNS coll.; Huene's No. 9; lost) cervical vertebra (61 mm)
(SMNS coll.; Huene's No. 11; lost) cervical vertebra (40 mm)
(SMNS coll.; Huene's No. 12; lost) cervical vertebra (42 mm)
Referred- (Breslau University coll.; syntypes of Thecodontosaurus primus)
Early Anisian, Middle Triassic
Lower Gogolin Formation of the Lower Muschelkalk, Netherlands

Early Anisian, Middle Triassic
Lower Gogolin Formation of the Lower Muschelkalk, Germany

Early Anisian, Middle Triassic
Lower Gogolin Formation of the Lower Muschelkalk, Europe

Comments- Wild's (1973) identification of Huene's No. 7 as SMNS 16687 is incorrect, and the former is the lectotype, not the latter (Fraser and Rieppel, 2006).
Huene (1905) first announced Tanystropheus antiquus without a diagnosis, making it a nomen nudum until the official description in 1908. It was described as a coelurid, and like T. conspicuus was thought to be represented by caudal vertebrae which were in actuality cervical vertebrae. Ortlam (1967) described material from the Upper Buntsandstein of Germany as T. longobardicus and Macrocnemus, but this was reinterpreted as T. antiquus by Wild (1980) who redescribed it. Wild (1987) claimed T. antiquus was no longer considered to belong to Tanystropheus based on these specimens, but the material was later described as Amotosaurus rotfeldensis by Fraser and Rieppel (2006). Evans (1988) stated T. antiquus was less closely related to T. longobardicus than Tanytrachelos based on having less cervical vertebrae and recommended either sinking Tanytrachelos or creating a new genus for antiquus. However, this was also based on the Amotosaurus material. Fraser and Rieppel state T. antiquus vertebrae can only be distinguished from T. conspicuus/longobardicus by their elongation and stratigraphy. While some new specimens have been described as T. antiquus (e.g. Huene, 1931; Wild and Oosterink, 1984), the species remains poorly known and needs redescription.
References- Huene, 1905. Uber die Trias-Dinosaurier Europas. Zeitschrift der Deutschen Geologischen Gesellschaft. 57, 345-349.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs of the European Triassic formations with consideration of occurrences outside Europe]. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Huene, 1914. Das natürliche System der Saurischia [The systematics of the Saurischia]. Centralblatt für Mineralogie, Geologie und Paläontologie. 1914, 154-158.
Huene, 1931. Über Tanystropheus und verwandte Formen. Neues Jahrbuch für Geologie und Paläontologie. 67, 65-86.
Peyer, 1931. Tanystropheus longobardicus Bass sp. Die Triasfauna der Tessiner Kalkalpen. Abhandlungen Schweizerische Paläontologie Gesellschaft. 50, 5-110.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Kuhn, 1937. Die fossilen Reptilien. Berlin: Gebrueder Borntrager. 121 pp.
Ortlam, 1967. Fossile Böden als Leithorizonte für die Gliederung des Höheren Buntsandsteins im nördlichen Scwarzwald und südlichen Odenwald. Geologisches Jahrbuch. 84, 485-590.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). Schweizerische Palaontologische Abhandlungen. 95, 1-162.
Wild, 1980. Tanystropheus (Reptilia: Squamata) and its importance for stratigraphy.Mémoires de la Société Géologique de France. 139, 201-206.
Wild and Oosterink, 1984. Tanystropheus (Reptilia, Squamata) aus dem Unteren Muschelkalk von Winterswijk, Holland. Grondboor en Hamer. 5(1984), 142-148.
Wild, 1987. An example of biological reasons for extinction: Tanystropheus (Reptilia, Squamata). Mémoires de la Société géologique de France. 150, 37-44.
Evans, 1988. The early history and relationships of the Diapsida. in Benton (ed.). The Phylogeny and Classification of the Tetrapods, Vol. 1: Amphibians, Reptiles, Birds. Clarendon Press, Oxford. 221-260.
Fraser and Rieppel, 2006. A new protorosaur from the Upper Buntsandstein of the Black Forest, Germany. Journal of Vertebrate Paleontology. 26(4), 866-871.
T. longobardicus (Bassani, 1886) Peyer, 1930
= Tribelesodon longobardicus Bassani, 1886
= "Macroscelosaurus" longobardicus (Bassani, 1886) Kuhn, 1937
= Tanystropheus meridensis Wild, 1980
Late Anisian-Early Ladinian, Middle Triassic
Grenzbitumenzone, Italy
Holotype-
(Milan Museum coll.; holotype of Tribelesodon longobardicus) skull, mandible, cervical vertebrae,
Late Anisian-Early Ladinian, Middle Triassic
Grenzbitumenzone, Switzerland

Ladinian, Middle Triassic
Meride Limestone, Switzerland

Late Ladinian, Middle Triassic
lowermost Keuper, SW Germany

Late Ladinian-Early Carnian, Middle-Late Triassic
Zhuganpo Member of the Falang Formation, Guizhou, China

Comments- Tribelesodon longobardicus was described by Bessani (1886) as a possible pterosaur, the doubt being caused by the tricusped posterior teeth which were seen as more similar to some ornithischians, mammals and lizards. Subsequent authors followed this assignment or felt Tribelesodon's placement was too uncertain owing to the short original description, until Nopcsa (1922, 1923) redescribed it in detail and assigned it to his new basal pterosaurian family Tribelesodontidae. While Olshevsky (2000) claims Tribelesodon has sometimes been called a theropod, I'm unaware of any such instances in the literature. Peyer discovered a complete skeleton in 1929, which was announced in 1930 and described in detail in 1931. He referred this specimen to longobardicus but recognized it was also Tanystropheus, with the supposed elongate manual digit IV phalanges of Tribelesodon being cervical vertebrae instead. Thus Peyer synonymized the genera and placed Tanystropheus in Protorosauridae. Ortlam (1967) described material from the Upper Buntsandstein of Germany as T. longobardicus and Macrocnemus, but this was later redescribed as Amotosaurus rotfeldensis by Fraser and Rieppel (2006). Numerous additional specimens have been described as Tanystropheus longobardicus by Wild (1970, 1973), Renesto (2005), Nosotti (2007) and Rieppel et al. (2010). Wild (1987) suggested T. longobardicus was probably identical with T. conspicuus, but no study of this issue has yet appeared. Fraser et al. (2004) find that T. meridensis' holotype can not be distinguished from small specimens of T. longobardicus and synonymized the species. They state that two species may be present though, one including all the largest specimens, having a short posterolateral premaxilla process, and unicuspate to very weakly tricuspate marginal teeth. As some small specimns also show this morphology, it may not be ontogenetic.
References- Bassani, 1886. Sui Fossili e sull’ età degli schisti bituminosi triasici di Besano in Lombardia. Atti della Società Italiana di Scienze Naturali. 19, 15-72.
Nopcsa, 1922. Neubeschreibung des Trias-Pterosauriers Tribelesodon. Paläontologische Zeitschrift. 5(2), 161-181.
Nopcsa, 1923. Neubeschreibung des Trias-Pterosauriers Tribelesodon. Paläontologische Zeitschrift. 5(3), 161-181.
Peyer, 1930. Tanystropheus longobardicus Bass. sp. Centralblatt für Mineralogie Abteilung B. 1930, 336-337.
Peyer, 1931. Tanystropheus longobardicus Bass sp. Die Triasfauna der Tessiner Kalkalpen. Abhandlungen Schweizerische Paläontologie Gesellschaft. 50, 5-110.
Huene, 1931. Über Tanystropheus und verwandte Formen. Neues Jahrbuch für Geologie und Paläontologie. 67, 65-86.
Peyer, 1939. Über die Rekonstruktion des Skelettes von Tanystropheus. Eclogae geologicae Helvetiae. 32(2), 203-209.
Kuhn, 1948. Der Schaedel von Tanystropheus. Eclogae geologicae Helvetiae. 40, 390.
Ortlam, 1967. Fossile Böden als Leithorizonte für die Gliederung des Höheren Buntsandsteins im nördlichen Scwarzwald und südlichen Odenwald. Geologisches Jahrbuch. 84, 485-590.
Wild, 1971. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). PhD thesis. Universitat Zurich.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). Schweizerische Palaontologische Abhandlungen. 95, 1-162.
Kuhn-Schnyder, 1974. Die Triasfauna der Tessiner Kalkalpen. Vierteljahresschrift der naturforschenden Gesellschaft in Zürich, Neujahrsblatt. 118, 1-119.
Wild, 1980. Die Triasfauna der Tessiner Kalkalpen. XXIV. Neue Funde von Tanystropheus (Reptilia, Squamata). Abhandlungen der schweizerischen paläontologischen Gesellschaft. 102, 1-31.
Wild, 1987. An example of biological reasons for extinction: Tanystropheus (Reptilia, Squamata). Mémoires de la Société géologique de France. 150, 37-44.
Tschanz, 1985. Tanystropheus- An unusual reptilian construction. Konstruktion lebender und ausgestorbener Reptilien. Konzepte SFB 230. 4(1985), 169-178.
Tschanz, 1986. Funktionelle Anatomie der Halswirbelsäule von Tanystropheus longobardicus (Bassani) aus der Trias (Anis/Ladin) des Monte San Giorgio (Tessin) auf der Basis vergleichend morphologischer Untersuchungen an der Halsmuskulatur rezenter Echsen. Unpublished Thesis. University of Zurich.
Tschanz, 1988. Allometry and heterochrony in the growth of the neck of Triassic prolacertiform reptiles. Palaeontology. 31(4), 997-1011.
Fraser, Nosotti and Rieppel, 2004. A re-evaluation of two species of Tanystropheus (Diapsida, Protorosauria), from Monte San Giorgio, Southern Alps. Journal of Vertebrate Paleontology. 24(3), 278.
Dalla Vecchia, 2005. Resti di Tanystropheus, saurotterigi e "rauisuchi" (Reptilia) nel Triassico Medio della Val Aupa (Moggio Udinese, Udine) [Remains of Tanystropheus, sauropterygians and "rauisuchians" (Reptilia) in the Middle Triassic of Aupa Valley (Udine, Friuli Venezia Giulia, NE Italy]. Gortania- Atti Museo Friulano di Storia Naturale. 27, 25-48.
Renesto, 2005. A new specimen of Tanystropheus (Reptilia Protorosauria) from the Middle Triassic of Switzerland and the ecology of the genus. Revista Italiana di Paleontologia e Stratigrafia. 11(3), 377-394.
Fraser and Rieppel, 2006. A new protorosaur from the Upper Buntsandstein of the Black Forest, Germany. Journal of Vertebrate Paleontology. 26(4), 866-871.
Nosotti, 2007. Tanystropheus longobardicus (Reptilia, Protorosauria): Re-interpretations of the anatomy based on new specimens from the Middle Triassic of Besano (Lombardy, northern Italy). Memorie della Societ`a Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano. 35, 1-88.
Rieppel, Jiang, Fraser, Hao, Motani, Sun and Sun, 2010. Tanystropheus cf. T. longobardicus from the early Late Triassic of Guizhou Province, southwestern China. Journal of Vertebrate Paleontology. 30(4), 1082-1089.
T. haasi Rieppel, 2001
Late Anisian, Middle Triassic
Gevanim Formation, Isreal
References
- Peyer, 1955. Demonstration von Trias-Vertebraten aus Palaestina. Eclogae Geologicae Helvetiae. 48, 486-490.
Haas, 1970. Eine bemerkenswerte Interclavicula von (?) Tanystropheus aus dem Muschelkalk des Wadi Ramon, Israel. Paläontologische Zeitschrift. 34(3-4), 207-214.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). Schweizerische Palaontologische Abhandlungen. 95, 1-162.
Rieppel, 2001. A new species of Tanystropheus (Reptilia: Protorosauria) from the Middle Triassic of Makhtesh Ramon, Israel. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 221(2), 271-287.
T. biharicus Jurcsak, 1975
Anisian, Middle Triassic
Alesd, Bihor, Romania
Reference
- Jurcsak, 1975. Tanystropheus biharicus n.sp. (Reptilia, Squamata), o nouva specie pentru fauna Triasica a României. Nymphaea. 3, 45-52.
T? fossai Wild, 1980
Late Norian, Late Triassic
Argillite di Riva di Solto, Italy
References
- Wild, 1980. Die Triasfauna der Tessiner Kalkalpen. XXIV. Neue Funde von Tanystropheus (Reptilia, Squamata). Abhandlungen der schweizerischen paläontologischen Gesellschaft. 102, 1-31.
Renesto, 2005. A new specimen of Tanystropheus (Reptilia Protorosauria) from the Middle Triassic of Switzerland and the ecology of the genus. Revista Italiana di Paleontologia e Stratigrafia. 11(3), 377-394.

Rhynchosauria Parker, 1879
Definition- (Mesosuchus browni + Rhynchosaurus articeps) (suggested)
Other definitions- (Mesosuchus browni + Hyperodapedon gordoni) (modified after Merck, 1997)
(Mesosuchus browni + Howesia browni) (modified after Dilkes, 1998)
Comments- The authorship is often given as "Osborn, 1903 (Gervais, 1859)", but Gervais used "Rhynchosaures" which is merely French for "rhynchosaurs." Rhynchosauria goes back at least to Parker (1879), though Osborn (1903) did establish it more officially.

Rhynchosauroidea Huxley, 1887 vide Nopcsa, 1928
Other definitions- (Howesia browni + Hyperodapedon gordoni) (modified after Merck, 1997)
Comments- According to the ICZN any superfamily must include its eponymous family, in this case Rhynchosauridae. If the latter were defined to exclude Hyperodapedon, Rhynchosauroidea could exist to include both Rhynchosauridae and Hyperodapedontidae.

Rhynchosauridae Huxley, 1887
Other definitions- (Rhynchosaurus articeps + Scaphonyx fischeri + Stenaulorhynchus stockleyi + Hyperodapedon gordoni) (modified after Dilkes, 1998)
(Stenaulorhynchus stockleyi + Hyperodapedon gordoni) (modified after Merck, 1997)
Comments- While Rhynchosauridae was defined prior to Hyperodapedontidae, according to the ICZN a family including both Rhynchosaurus and Hyperodapedon must be named after the latter since its family level taxon was named two years earlier. Rhynchosauridae remains available for a sister taxon to Hyperodapedontidae.

Hyperodapedontidae Lydekker, 1885
Definition- (Hyperodapedon gordoni <- Rhynchosaurus articeps) (modified after Langer and Schultz, 2000)
= Scaphonychidae Huene, 1928

Hyperodapedontinae Lydekker, 1885 vide Chatterjee, 1969
Definition- (Hyperodapedon gordoni <- Fodonyx spenceri) (Langer and Schultz, 2000)

Hyperodapedon Huxley vide Murchison, 1858
= Stenometopon Boulenger, 1904
= Scaphonyx Woodward, 1907
= Cephalastron Huene, 1926
= Cephalonia Huene, 1926
= Cephalostronius Huene, 1926
= Scaphonychimus Huene, 1926
= Macrocephalosaurus Tupi-Caldas, 1933
= Paradapedon Huene, 1938
= Supradapedon Chatterjee, 1980
Definition- (Hyperodapedon gordoni <- Teyumbaita sulcognathus) (Langer and Schultz, 2000)
Comments- Huene's (1926) new taxa (Cephalastron brasiliense, Cephalastron gondwanicum, Cephalonia lotziana, Cephalostronius angustispinatus, Scaphonyx australis and Scaphonychimus eurychors) are all near certainly referrable to Hyperodapedon (H. sanjuanensis, H. huenei or H. mariensis), but currently viewed as indeterminate (Langer, 1996). These species are not given their own entries here, as I lack both Langer's thesis and Huene's original description and none have been formally made new combinations with Hyperodapedon.
References- Murchison, 1858. On the sandstones of Morayshire (Elgin, &c.) containing reptilian remains; and on their relations to the Old Red Sandstone of that country. Quarterly Journal of the Geological Society of London. 15, 419-439.
Boulenger, 1904. On reptilian remains from the Triass of Elgin. Philosophical Transactions of the Royal Society of London B. 196, 175-189.
Woodward, 1907. On some fossil reptilian bones from the state of Rio Grande do Sul. Revista do Museu Paulista. 7, 46-57.
Huene, 1926. Gondwana-Reptilien in Südamerika. Palaeontologia Hungarica. 2, 1-108.
Tupi-Caldas, 1933. Contribuição ao estudo do fossil da Alemoa, Município de Santa Maria, Rio Grande do Sul. In Tupi-Caldas (ed.). Curso Geral de Mineralogia e Geologia, aplicada ao Brasil. Edições da Livraria do Globo. 333-339.
Huene, 1938. Stenaulorhynchus, ein Rhynchosauridae der ostafrikanischen Obertrias. Nova Acta Leopoldina. 1938, 83-121.
Chatterjee, 1980. The evolution of rhynchosaurs. Memoires de la Societe Geologique de France, Nouvelle Serie. 139, 57-65.
Langer, 1996. Rincossauros sul-brasileiros: Historico e filogenia. Masters thesis, Universidade Federal do Rio Grande do Sul. 361 pp.
H. fischeri (Woodward, 1907) Whatley, 2005
= "Scaphonyx fischeri" White, 1906
= Scaphonyx fischeri Woodward, 1907
Carnian, Late Triassic
Alemoa Member of the Santa Maria Formation, Brazil

Holotype- (BM R-5033) two cervical centra, dorsal centrum, central fragment, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, pedal ungual I
Comments- The holotype was discovered in 1902, and initially announced by Woodward in 1903 before being described and named by him in 1907. White (1906) first published the name in a note in Science, but did not provide a description or definition (ICZN Article 12.1), making the name a nomen nudum. Woodward (1907) identified Scaphonyx as a Euskelosaurus-like dinosaur based on several characters. First, the dorsal centrum lacks a parapophysis, supposedly unlike 'anomodonts' (under which he included pareiasaurs, procolophonids and therapsids), but rhynchosaurs (which Woodward classified as rhynchocephalians) possess the same state as Scaphonyx. Second, the cervical supposedly resembled Euskelosaurus, but this was based on a specimen (BMNH R2791) now referred to Erythrosuchus (as foreseen in Woodward's postscript). The large pedal ungual I with obliquely curved unguals was compared favorably to sauropods, but is also present in derived hyperodapedontines. Finally, a pedal digit with four phalanges was considered similar to dinosaurs and unlike 'anomodonts', but rhynchosaurs have three pedal digits with this many phalanges as well, and the digit closely matches manual digit III of Alemoa Hyperodapedon. Woodward's 1907 paper was actually written in 1904, and when reprinted in 1908 he included a postscript which recognized BMNH R2791 as non-dinosaurian. As he compared it favorably to Erythrosuchus (considered by Woodward to resemble both 'anomodonts' and 'belodonts'- the latter containing parasuchians and aetosaurs), Woodward now considered Scaphonyx an 'anomodont'.
Huene (1908) noted Scaphonyx was unlike dinosaurs in the presence of postaxial intercentra, cervical diapophyses and parapophyses which are placed high on the vertebra, and dissimilar unguals. He suggested it might be a therapsid or parasuchian. In 1911, Huene proposed Scaphonyx and Erythrosuchus were members of his new 'thecodont' group Pelycosimia, which continued through 1926 when he gave Scaphonyx its own family. In 1929, Huene finally recognized the similarity between Scaphonyx and rhynchosaurs, assigning the genus to the group. While long considered a valid genus of rhynchosaur, Langer (1996; published in Langer and Schultz, 2000a) proposed Scaphonyx fischeri's holotype is indeterminate, as multiple species are known from the Alemoa member (mariensis, sanjuanensis, and what would be named huenei) which have only been distinguished using cranial characters. Although huenei is not known from postcrania (so can't be compared to fischeri), mariensis and sanjuanensis have not had their vertebral or pedal anatomy compared in detail in the published literature. Indeed, Alemoa rhynchosaurs have not had their postcrania well described in over seventy years. Given these facts and that I lack access to both Langer's thesis and mariensis' original and only published description, I only consider it provisionally indeterminate here. Additional specimens assigned to S. fischeri by Huene (1926, 1942) have been considered indeterminate or referrable to S. sanjuanensis (Langer and Schultz, 2000b; Montefeltro, 2008; Langer, pers. comm. 2015).
References- Woodward, 1903. On some dinosaurian bones from south Brazil. Geological Magazine. 10(11), 512.
White, 1906. Geology of south Brazil. Science. 24(612), 377-379.
Woodward, 1907. On some fossil reptilian bones from the state of Rio Grande do Sul. Revista do Museu Paulista. 7, 46-57.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Woodward, 1908. On some fossil reptilian bones from the state of Rio Grande do Sul. Geological Magazine. 5(6), 251-255.
Huene, 1911. Über Erythrosuchus, Vertreter der neuen Reptil-Ordnung Pelycosimia. Geologische und Paläontologische Abhandlungen. 10(1), 1-60.
Huene, 1926. Gondwana-Reptilien in Südamerika. Palaeontologia Hungarica. 2, 1-108.
Huene, 1929. Über Rhynchosaurier und andere Reptilien aus den Gondwana-Ablagerungen Südamerikas. Geologie und Palaeontogie Abhandlungen. 17, 1-61.
Huene, 1942. Die fossilen Reptilien des sudamerikanischen Gondwanalandes. C. H. Beck, Munich. 342 pp.
Langer, 1996. Rincossauros sul-brasileiros: Historico e filogenia. Masters thesis, Universidade Federal do Rio Grande do Sul. 361 pp.
Langer and Schultz, 2000a. Rincossauros-herbivoros cosmopolitas do Triassico. In Holz and de Ros (eds.). Paleontologia do Rio Grande do Sul. Porto Alegre. Ediitora da Universidade, CIGO/UFRGS, Brazil. 246-272.
Langer and Schultz, 2000b. A new species of the Late Triassic rhynchosaur Hyperodapedon from the Santa Maria Formation of south Brazil. Palaeontology. 43, 633-652.
Whatley, 2005. Phylogenetic relationships of Isalorhynchus genovefae, the rhynchosaur (Reptilia, Archosauromorpha) from Madagascar. PhD thesis, University of California. 276 pp.
Montefeltro, 2008. Inter-relações filogenéticas dos rincossauros (Diapsida, Archosauromorpha). Masters thesis, Universidade de Sao Paulo. 203 pp.
H. huenei Langer and Schultz, 2000
Carnian, Late Triassic
Alemoa Member of the Santa Maria Formation, Brazil
Reference
- Langer and Schultz, 2000. A new species of the Late Triassic rhynchosaur Hyperodapedon from the Santa Maria Formation of south Brazil. Palaeontology. 43, 633-652.
H. mariensis (Tupi-Caldas, 1933) Langer, Boniface, Cuny and Barbieri, 2000
= Macrocephalosaurus mariensis Tupi-Caldas, 1933
Carnian, Late Triassic
Alemoa Member of the Santa Maria Formation, Brazil

References- Tupi-Caldas, 1933. Contribuição ao estudo do fossil da Alemoa, Município de Santa Maria, Rio Grande do Sul. In Tupi-Caldas (ed.). Curso Geral de Mineralogia e Geologia, aplicada ao Brasil. Edições da Livraria do Globo. 333-339.
Langer, Boniface, Cuny and Barbieri, 2000. The phylogenetic position of Isalorhynchus genovefae, a Late Triassic rhynchosaur from Madagascar. Annales de Paléontologie. 86(2), 101-127.
H. sanjuanensis (Sill, 1970) Langer, Boniface, Cuny and Barbieri, 2000
= Scaphonyx sanjuanensis Sill, 1970
Late Carnian, Late Triassic
La Pena and Cancha de Bochas Members of the Ischigualasto Formation, San Juan, Argentina

Carnian, Late Triassic
Alemoa Member of the Santa Maria Formation, Brazil

References- Sill, 1970. Scaphonyx sanjuanensis, nuevo rincosaurio (Reptilia) de la formacion Ischigualasto, Triasico de San Juan, Argentina. Ameghiniana. 7, 341-354.
Langer, Boniface, Cuny and Barbieri, 2000. The phylogenetic position of Isalorhynchus genovefae, a Late Triassic rhynchosaur from Madagascar. Annales de Paléontologie. 86(2), 101-127.
H. huxleyi Lydekker, 1881
= Paradapedon huxleyi (Lydekker, 1881) Huene, 1938
Carnian, Late Triassic
Lower Maleri Formation, India
References
- Lydekker, 1881. Note on some Gondwana vertebrates. Records of the Geological Survey of India. 15, 174-178.
Huene, 1938. Stenaulorhynchus, ein Rhynchosauridae der ostafrikanischen Obertrias. Nova Acta Leopoldina. 1938, 83-121.
H. gordoni Huxley vide Murchison, 1858
?= Hyperodapedon minor Burckhardt, 1900b
= Stenometopon taylori Boulenger, 1904
Early Norian, Late Triassic
Lossiemouth Sandstone Formation, Scotland
Comments
- The citation for Murchison (1858) is often listed incorrectly, misspelled 'Murchinson', cited as 1859, with an erroneous title, and pagination from Huxley's 1869 paper.
Hyperodapedon minor- While Hyperodapedon minor has never been associated with saurischians, its details are not widely known so are discussed here. This species was established by Burckhardt (1900b; page 492) for two small maxillae and a mandible from Warwickshire which were mentioned in a footnote by Huxley (1869) as H. gordoni. Only a few statements were made about H. minor in Burckhardt's work, with the only proposed distinguishing character being a more posteriorly extensive dentary tooth row than H. gordoni. The taxon has been virtually ignored in the literature since, though Huene (1942) did say its distinctiveness from H. gordoni is unfounded and that it should probably be rejected. Discussion with Benton (pers. comm 2015) indicates Burckhardt only visited the BMNH, though no specimens there were indicated as belonging to this species. Based on Benton's unpublished thesis notes, I believe BMNH R3150 (listed as "Partial skull 18 pieces, some fitting: palate views of mx, pal etc - small animal") is the best possibility for being H. minor's holotype, though it's possible the holotype has remained unnoticed or become lost. Regardless, the fact all diagnostic Lossiemouth Sandstone rhynchosaurs have been referred to H. gordoni suggests H. minor is similarly referrable.
References- Murchison, 1858. On the sandstones of Morayshire (Elgin, &c.) containing reptilian remains; and on their relations to the Old Red Sandstone of that country. Quarterly Journal of the Geological Society of London. 15, 419-439.
Huxley, 1869. On Hyperodapedon. Quarterly Journal of the Geological Society of London. 25, 138-152.
Burckhardt, 1900a. On Hyperodapedon gordoni. Geological Magazine. 7(12), 529-535.
Burckhardt, 1900b. On Hyperodapedon gordoni. Geological Magazine. 7(37), 486-492.
Boulenger, 1904. On reptilian remains from the Triass of Elgin. Philosophical Transactions of the Royal Society of London B. 196, 175-189.
Huene, 1942. Die fossilen Reptilien des sudamerikanischen Gondwanalandes. C. H. Beck, Munich. 342 pp.
Benton, 1981. The Triassic reptile Hyperodapedon from Elgin, functional morphology and relationships. PhD thesis, University of Newcastle upon Tyne. [? pp].
H. stockleyi (Boonstra, 1953) Mukherjee and Ray, 2014
= Scaphonyx stockleyi Boonstra, 1953
= Supradapedon stockleyi (Boonstra, 1953) Chatterjee, 1980
Carnian?, Late Triassic
Tunduru District, Tanzania
Comments
- While the combination Hyperodapedon stockleyi is usually credited to Langer et al. (2000), they merely listed Scaphonyx stockleyi under "Described rhynchosaurs attributable to Hyperodapedon" without claiming the species was valid or using the new combination.
References- Boonstra, 1953. A note on some rhynchosaurian remains from Tanganyika territory. Annals of the South African Museum. 42, 1-4.
Chatterjee, 1980. The evolution of rhynchosaurs. Memoires de la Societe Geologique de France, Nouvelle Serie. 139, 57-65.
Langer, Boniface, Cuny and Barbieri, 2000. The phylogenetic position of Isalorhynchus genovefae, a Late Triassic rhynchosaur from Madagascar. Annales de Paléontologie. 86(2), 101-127.
Mukherjee and Ray, 2014. A new Hyperodapedon (Archosauromorpha, Rhynchosauria) from the Upper Triassic of India: Implications for rhynchosaur phylogeny. Palaeontology. 57(6), 1241-1276.
H. tikiensis Mukherjee and Ray, 2014
Carnian, Late Triassic
Tiki Formation, India
Reference
- Mukherjee and Ray, 2014. A new Hyperodapedon (Archosauromorpha, Rhynchosauria) from the Upper Triassic of India: Implications for rhynchosaur phylogeny. Palaeontology. 57(6), 1241-1276.

Allokotosauria Nesbitt, Flynn, Pritchard, Parrish, Ranivoharimanana and Wyss, 2015
Definition- (Azendohsaurus madagaskarensis, Trilophosaurus buettneri <- Tanystropheus longobardicus, Proterosuchus fergusi, Protorosaurus speneri, Rhynchosaurus articeps) (Nesbitt et al., 2015)
Comments- The type species (Azendohsaurus laaroussii, Tanystropheus conspicuus) should have been used in the definition.

Azendohsauridae Nesbitt, Flynn, Pritchard, Parrish, Ranivoharimanana and Wyss, 2015
Definition- (Azendohsaurus madagaskarensis <- Trilophosaurus buettneri, Tanystropheus longobardicus, Proterosuchus fergusi, Protorosaurus speneri, Rhynchosaurus articeps, Passer domesticus) (Nesbitt et al., 2015)
Comments- The type species (Azendohsaurus laaroussii, Tanystropheus conspicuus) should have been used in the definition.

Azendohsaurus

Trilophosauridae Gregory, 1945
= Trilophosauria Romer, 1956

Spinosuchus Huene, 1932
= Chinleogomphius Sues and Olsen, 1993
S. caseanus Huene, 1932
= Trilophosaurus jacobsi Murry, 1987
= Chinleogomphius jacobsi Sues and Olsen, 1993
= Trilophosaurus dornorum Mueller and Parker, 2006
Late Carnian, Late Triassic
Tecovas Formation of the Dockum Group, Texas, US
Holotype
- (UMMP 7507) presacral vertebral column
Comments- Discovered in 1921 and originally referred to Coelophysis by Case (1922, 1927), this was separated as a new genus of coelophysoid by Huene (1932). Viewed as non-dinosaurian by Padian (1986) and Murry and Long (1989), it was referred to Theropoda by Hunt et al. (1998) because of the supposedly hollow centra. The latter are found in some pseudosuchians too however. Richards (1999) referred Spinosuchus to Trilophosauridae in his unpublished thesis, but Nesbitt et al. (2007) stated the characters he used are found in other archosaurmorphs, though they confusingly listed it as Archosauriformes indet. instead of Archosauromorpha indet.. Spielmann et al. (2007; 2009) have more recently demonstrated its trilophosaurid identity convincingly and redescribed the taxon.
References- Case, 1922. New reptiles and stegocephalians from the Upper Triassic of Western Texas. Carnegie Institution of Washington Publication. 321, 1-84.
Case, 1927. The vertebral column of Coelophysis Cope. Contributions from the Museum of Geology, University of Michigan. 10, 209-222.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Padian, 1986. On the type material of Coelophysis Cope (Saurischia: Theropoda) and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation). in Padian (ed.). The Beginning of the Age of Dinosaurs. Cambridge University Press, Cambridge. 45-60.
Murry and Long, 1989. Geology and paleontology of the Chinle Formation, Petrified Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils of the southwestern Upper Triassic. in Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. 29-64.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the western United States. Geobios. 31(4), 511-531.
Richards, 1999. Osteology and relationships of Spinosuchus caseanus HUENE, 1932 from Texas (Dockum Group, Upper Triassic): A new interpretation. Masters thesis, Fort Hays State University. 157 pp.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.
Spielmann, Lucas, Hunt and Heckert, 2007. A revision of the Trilophosauridae (Archosauromorpha) from the Late Triassic of western North America. Journal of Vertebrate Paleontology. 27(3), 151A.
Spielmann, Lucas, Heckert, Rinehart and Richards, 2009. Redescription of Spinosuchus caseanus (Archosauromorpha: Trilophosauridae) from the Upper Triassic of North America. Palaeodiversity. 2, 283-313.

Prolacertiformes Camp, 1945
Definition- (Prolacerta broomi <- Crocodylus miloticus) (modified from Senter, 2004)

Prolacerta

Archosauriformes Gauthier et al., 1988
Definition- (Proterosuchus fergusi + Crocodylus niloticus) (Nesbitt, 2011; modified from Kischlat, 2000)
Other definitions- (Proterosuchus fergusi + Caiman crocodilus + Vultur gryphus) (Gauthier et al., 2004)

Ankistrodon Huxley, 1865
= Epicampodon Lydekker, 1885
A. indicus Huxley, 1865
= Epicampodon indicus (Huxley, 1865) Lydekker, 1885
= Thecodontosaurus indicus (Huxley, 1865) Huene, 1908
= Chasmatosaurus indicus (Huxley, 1865) Huene, 1942
Early Scythian, Early Triassic
Panchet Formation, India
Holotype
- (GSI coll.) dentary fragment, two teeth
Referred- ?(GSI coll.) cervical vertebrae, anterior dorsal vertebrae, sacral vertebrae, proximal caudal vertebrae (Huene, 1942)
Comments- Huxley named and described the holotype, believing it to be a thecodont most similar to the parasuchian Clepsysaurus based on the presence of only distal serrations. He later (1870) referred Thecodontia to Dinosauria, leading him to refer Ankistrodon to the latter clade. Seeley (1880) mentioned the genus as a dinosaur related to Megalosaurus. He later (1885) created the genus Epicampodon for the taxon, since he incorrectly thought Ankistrodon was preoccupied by the recent viperid genus Agkistrodon Palisot de Beauvois, 1799 (or its own unjustified emmendation Ancistrodon Wagler, 1830; or the pycnodontiform fish Ancistrodon Roemer, 1849, which was renamed Ankistrodus then Grypodon). Seeley (1888) illustrated Epicampodon as an anchisaurid theropod, Nicholson and Lydekker (1889) included it in Megalosauridae, while Nopcsa (1901) listed it as an anchisaurine megalosaurid. It was listed as a zanclodontid theropod by Zittel (1890) and Huene (1902), who incorrectly believed it to be from the Late Triassic Maleri beds. Huene (1906) stated that the species was probably referrable to Thecodontosaurus, though he did not explicitly list the new combination until 1908. In that work he called it Thecodontosaurus(?) indicus, believing it to be most similar to Paleosaurus (his Thecodontosaurus cylindrodon). Das-Gupta (1931) thought the species was similar to his new theropod Orthogoniosaurus in having only distal serrations and having a straight distal edge (actually caused by apical breakage in indicus), placing both in Anchisauridae within Theropoda. Huene still referred Epicampodon to Saurischia in 1940, but in 1942 recognized it was a more basal archosauriform and made it a species of Chasmatosaurus. This was followed by Tatarinov (1961) and Hughes (1963; who nonetheless considered the holotype generically indeterminate), but Charig et al. (1976) correctly noted Ankistrodon has priority over Chasmatosaurus (and its senior synonym Proterosuchus). Charig and Reig (1970) and Reig (1970) considered it an indeterminate proterosuchian. Romer (1972) considered the material to be Proterosuchus without explicitly naming the new combination, though he later (1976) questionably synonymized it with Chasmatosaurus. Most recently, Charig et al. (1976) considered Ankistrodon a seemingly valid genus of proterosuchid, which was followed by Olshevsky (1991). While the presence of serrations does indicate an archosauriform, the Early Triassic age excludes parasuchians and theropods, and the recurved crowns exclude a relationship to sauropodomorphs like Thecodontosaurus or Anchisaurus, evidence supporting a close relationship with Proterosuchus was perhaps only given by Huene (1942), prior to the discovery of a large amount of 'proterosuchian' diversity. Recent analyses suggest that the traditional Proterosuchidae is a paraphyletic grade of basal archosauriforms, so Ankistrodon is here placed in Archosauriformes until further studies on mandibular and dental variation are performed.
Huxley (1865) also described vertebrae as belonging to Dicynodon orientalis (now Lystrosaurus murrayi), which were recognized by Huene (1942) as being archosauriform and referred to his Chasmatosaurus indicus. As the vertebrae cannot be compared to the type jaw fragment, Charig and Reig (1970) merely called them cf. Chasmatosaurus sp..
References- Huxley, 1865. On a collection of vertebrate fossils from the Panchet Rocks, Ranigunu, Bengal. Memoirs of the Geological Survey of India; Paleontologia Indica, Series IV. Indian Pretertiary Vertebrata. i, 3-24.
Huxley, 1870. Triassic Dinosauria. Nature. 1, 23-24.
Lydekker, 1880. A sketch of the history of the fossil Vertebrata of India. Journal and Proceedings of the Asiatic Society of Bengal. 69(2), 8-40.
Lydekker, 1885. The Reptilia and Amphibia of the Maleria and Denwa Groups. Memoirs of the Geological Survey of India. Palaeontologia Indica, Series IV. Indian Pretertiary Vertebrata. 1(5), 1-38.
Seeley, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum of Natural History, London. 309 pp.
Nicholson and Lydekker, 1889. A manual of palaeontology for the use of students: with a general introduction on the principles of palæontology, Volume 2. 1624 pp.
Zittel, 1890. Handbuch der Palaeontologie. Volume III. Vertebrata (Pisces, Amphibia, Reptilia, Aves). 900 pp.
Nopcsa, 1901. A dinosaurusok atnezete es szarmazasa. Földtani Közlöny. 31, 193-224.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen (Neue Serie). Gustav Fischer Verlag, Jena. 6, 1-84.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs of the European Triassic formations with consideration of occurrences outside Europe]. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Das-Gupta, 1931. On a new theropod dinosaur (Orthogoniosaurus matleyi, n. gen. et n. sp.) from the Lameta beds of Jubbulpore. Journal and Proceedings of the Asiatic Society of Bengal. 26, 367-369.
Huene, 1940. Die Saurier der Karroo-, Gondwana-, und verwandten Ablagerungen in faunistischer, biologischer und phylogenetischer Hinsicht [Saurians of the Karroo, Gondwana, and other deposits in faunistic, biological, and phylogenetic regard]. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie. 83, 246-347.
Huene, 1942. Die Fauna der Panchet-Schichten in Bengalen [The fauna of the Panchet beds in Bengal]. Zentralblatt für Mineralogie, Geologie und Paläontologie, Abteilung B: Geologie und Paläontologie. 1941(11), 354-360.
Tatarinov, 1961. Pseudosuchians of the USSR. Paleontologicheskii Zhurnal. 1961(1), 117-132.
Hughes, 1963. The earliest archosaurian reptiles. South African Journal of Science. 59, 221-240.
Charig and Reig, 1970. The classification of the Proterosuchia. Biological Journal of the Linnean Society. 2, 125-171.
Reig, 1970. The Proterosuchia and the early evolution of the archosaurs; an essay about the origin of a major taxon. Bulletin of The Museum of Comparative Zoology. 139, 229-292.
Romer, 1972. The Chaneres (Argentina) Triassic reptile fauna. XVI. Thecodont classification. Breviora. 395, 24 pp.
Charig, Krebs, Sues and Westphal, 1976. Thecodontia. Encyclopedia of Paleoherpetology. 13, 137 pp.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.

Cladeiodon Owen, 1841
C. lloydi Owen, 1841
= Teratosaurus lloydii (Owen, 1841) Huene, 1908
Anisian, Middle Triassic
Bromsgrove Sandstone Formation, England
Holotype
- (lost) tooth
Referred- (BMNH R2645) tooth (Huene, 1908)
(BMNH R4847) tooth (Benton and Gower, 1997)
(BMNH R4848) tooth (Benton and Gower, 1997)
(CAMSM G352) tooth (Huene, 1908)
(WARMS G.7) tooth (Murchison and Strickland, 1840)
(WARMS G.8) tooth (Murchison and Strickland, 1840)
(WARMS G.954) tooth (18 x 10 x ? mm)
(WARMS G.956.1) tooth (Walker, 1969)
(WARMS G.956.2) tooth (47 mm)
(WARMS G.957) tooth (Walker, 1969)
(WARMS G.969) tooth (18 mm) (Walker, 1969)
Comments- The entry for this taxon is in progress, though the opinions of several authors are liasted below.
Owen, 1841- 'thecodont' intermediate between Thecodontosaurus and "Paleosaurus" platyodon in Lacertilia.
Owen, 1842b- same as above, states "one of the teeth of Cladyodon" is WARMS G.7.
Pictet, 1845- Lacertiformes.
Geinitz, 1846- Lacertia.
Owen, 1860- Thecodontia related to Paleosaurus, not distinguishable from Belodon, states original specimen is WARMS G.7.
Owen, 1868- Crocodilia.
Huxley, 1870- dinosaur, not similar to Thecodontosaurus or "Paleosaurus" platyodon, two teeth referrable to Paleosaurus cylindrodon, another possibly Teratosaurus (lost; contra Benton and Gower, 1997 this is not WARMS G.956.2, as it is less elongate).
Phillips, 1871- dinosaur.
Nicholson and Lydekker, 1889- megalosaurid, possibly synonym of Paleosaurus.
Woodward and Sherborn, 1890- =Belodon?, type in the Geol. Soc. London.
Zittel, 1902- megalosaurid close to Zanclodon.
Huene, 1908- in Teratosaurus.
Colbert and Chaffee, 1941- theropod.
Walker, 1969- belongs to poposaurid (later named Bromsgroveia).
Kuhn, 1970- ornithosuchid.
Charig et al., 1976- saurischian dinosaur.
Romer, 1976- teratosaurid carnosaur.
Welles, 1984- probably theropod, indeterminate, nomen nudum.
Chure and McIntosh, 1989- teratosaurid theropod.
Olshevsky, 1991- synonym of Teratosaurus suevicus.
Benton et al., 1994- Archosauria indet., maybe Bromsgroveia or "large thecodont" (= Bromsgroveia- Benton and Gower, 1997).
Benton and Gower, 1997- provisional synonym of Bromsgroveia, within range of shape of Postosuchus but higher serration density, WARMS G.954 may be the holotype (though this is not true, as it is too short and lacks recurvature).
Often misspelled as- Cladyodon Owen, 1842b; Kladyodon and Kladeisteriodon Plieninger, 1846;Claderodon Agassiz, 1846; Claydyodon Mantell, 1848.
References- Murchison and Strickland, 1840. On the upper formations of the New Red Sandstone
Owen, 1841. Odontography; or a treatise on the comparative anatomy of the teeth, I Part 11. Dental system of reptiles. Hippolyte Bailliere, London. 179-295.
Owen, 1842a. Second rapport sur le reptiles fossiles de le Grande Bretagne. L'Institut. 11-14.
Owen, 1842b. Report on British fossil reptiles. Part II. Report of the Eleventh Meeting of the British Association for the Advancement of Science. 60-204.
Pictet, 1845. Traité élémentaire de paléontologie: ou, Histoire naturelle des animaux fossiles considérés dans leurs rapports zoologiques et géologiques, Volume 2. 407 pp.
Geinitz, 1846. Grundriss der Versteinerungskunde. 813 pp.
Plieninger, 1846a.
Mantell, 1848. The wonders of geology: or A familiar exposition of geological phenomena. Volume 2. 938 pp.
Owen, 1860. Palaeontology, or a systematic summary of extinct animals and their geological relations. 420 pp.
Owen, 1868. On the Anatomy of Vertebrates. Volume III. Mammals. 915 pp.
Huxley, 1870. On the classification of the Dinosauria, with observations on the Dinosauria of the Trias. Quarterly Journal of the Geological Society. 26, 32-51.
Phillips, 1871. Geology of Oxford and the Valley of the Thames: Oxford at the Clarendon Press. 523pp.
Nicholson and Lydekker, 1889. A manual of palaeontology for the use of students: with a general introduction on the principles of palæontology, Volume 2. 1624 pp.
Woodward and Sherborn, 1890. A catalogue of British fossil Vertebrata. 396 pp.
Zittel, 1902. Text-book of palaeontology, Volume 2. 283 pp.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs of the European Triassic formations with consideration of occurrences outside Europe]. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Colbert and Chaffee, 1941. The type of Clepsysaurus pennsylvanicus
Walker, 1969. The reptile fauna of the "Lower Keuper" Sandstone. Geological Magazine. 106, 470-476.
Kuhn, 1970.
Charig, Krebs, Sues and Westphal, 1976. Thecodontia. Encyclopedia of Paleoherpetology. 13, 137 pp.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda): Osteology and comparisons. Palaeontographica Abteilung A. 185, 85-180.
Benton,Warrington, Newell and Spencer, 1994. A review of the British Middle Triassic tetrapod assemblages. in Fraser and Sues (eds). In the shadow of the dinosaurs: Early Mesozoic tetrapods. 131-160.
Galton and Walker, 1996a. Bromsgroveia from the Middle Triassic of England
Galton and Walker, 1996b. Supposed prosauropod dinosaurs from Middle Triassic
Benton and Gower, 1997. Richard Owen's giant Triassic frogs:

"Teleocrater rhadinus"

Teratosaurus? bengalensis Das-Gupta, 1929
= Teratosaurus "bengalensis" Das-Gupta, 1927
Early Scythian, Early Triassic
Panchet Formation, India

Holotype- tooth
Comments- This species was first reported by Das-Gupta in 1927 without a description, making that mention a nomen nudum. In 1927 it was explicitly called a theropod, while in 1929 it was referred to as a carnivorous dinosaur. The description does not include any explicit rationale for referring bengalensis to Teratosaurus, which was done because it was similar to teeth illustrated by Huene. Chure and McIntosh (1989) listed it as incorrectly referred to Teratosaurus, while Olshevsky (1991) retains it questionably in that genus. It is probably indeterminate based on the description and is here referred to Archosaauriformes due to its serrations.
References- Das-Gupta, 1927. Batrachian and reptilian remains found in the Panchet Beds at Deoli, Bengal. Proceedings of the Indian Science Congress. 14, 240.
Das-Gupta, 1929. Batrachian and reptilian remains found in the Panchet Beds at Deoli, Bengal. Journal and Proceedings of the Asiatic Society of Bengal. 24(4), 473-479.
Rozhdestvensky, 1977. The study of dinosaurs in Asia. Journal of the Palaeontological Society of India. 20, 102-119.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.

Zanclodon Plieninger, 1846
Z. laevis
Middle Triassic
Erfurt Formation, Germany
Lectotype-
(SMNS 6045) partial maxilla
Z? crenatus

Z? schuetzii Fraas, 1900 emmend. Olshevsky, 2000
= Zanclodon schützii Fraas, 1900
= Teratosaurus schützii (Fraas, 1900) Huene, 1932
Middle Triassic
Muschelkalk, Germany

Holotype- (SMNS coll.) tooth
Comments- Huene (1908) thought this tooth might be referrable to Tanystropheus conspicuus, which he placed in Coeluridae.
References- Fraas, 1900. Zanclodon schützii n. sp. aus dem Trigonodusdolomit von Hall [Zanclodon schützii n. sp. from the Trigonodus-dolomite of Halle]. Jahreshefte des Vereins für Vaterländische Naturkunde in Württemberg. 56, 510-513.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs of the European Triassic formations with consideration of occurrences outside Europe]. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings. 3, 1-157.
Z? silesiacus

Proterosuchia Broom, 1906
Definition- (Proterosuchus fergusi <- Crocodylus niloticus) (modified from Kischlat, 2000)

Erythrosuchia Watson, 1957
Definition- (Erythrosuchus africanus <- Crocodylus niloticus) (modified from Kischlat, 2000)

Euparkeriidae

unnamed clade (Proterochampsa barrionuevoi + Crocodylus niloticus)

Avalonianus Kuhn, 1961
= Avalonia Seeley, 1898 (preoccupied Walcott, 1890)
A. sanfordi (Seeley, 1898) Kuhn, 1961
= Avalonia sanfordi Seeley, 1898
Rhaetian, Late Triassic
Westbury Formation, England
Holotype
- (BMNH R2869) tooth (33 x 15 x 7 mm)
Comments- The holotype was found in 1894 along with Picrodon herveyi's holotype and various postcrania (BMNH R2870-2874, 2676-2878), all described by Sanford (1894) as a dinosaur similar to Megalosaurus. Seeley (1898) later more fully described and named the remains, associating the tooth BMNH R2869 with BMNH R2870-2873 (two posterior dorsal vertebrae, femur, proximal tibia, two pedal phalanges, pedal ungual) to form Avalonia sanfordi. Seeley referred most of the other remains to Picrodon and referred to both as zanclodont dinosaurs. Newton (1899) believed Avalonia was not sufficiently distinguished from Zanclodon and that Picrodon and its associated postcrania probably belonged to the same individual. He thought the Avalonia holotype closely resembled his new taxon "Zanclodon" cambrensis but differed in having more strongly curved teeth. Seeley (in Newton, 1899) also thought cambrensis was most similar to Avalonia, but felt the latter was generically diagnostic compared to Zanclodon. Nopcsa (1901) listed Avalonia as an anchisaurine megalosaurid, with Picrodon listed as Avalonia? herveyi. Huene (1902) listed both Avalonia and Picrodon as synonyms of Gressylosaurus, which he believed was a zanclodontid theropod. He later (1905) listed them as synonyms of Gresslyosaurus cf. ingens in the precursor to his 1908 study which more firmly accepted their synonymy with G. ingens, which he placed as a plateosaurid theropod. In that paper, Huene stated the synonymy was based on the postcrania and believed it and the two teeth came from one individual. The slight differences in serration morphology and density, tooth size and serration extent were seen as due to different positions in the jaw, with Avalonia being more anterior than Picrodon. Huene (1932) continued to list Avalonia and Picrodon as synonyms of Gresslyosaurus ingens, which he had moved to Teratosauridae within Carnosauria. In 1956, Huene still followed that synonymy. Romer (1976) and Chure and McIntosh (1989) both listed Avalonia as a teratosaurid theropod as well, with the former believing it to be a junior synonym of Teratosaurus within Carnosauria. Kuhn (1961) noted that Avalonia was preoccupied by an ?atopid trilobite genus named in 1890, so proposed the replacement name Avalonianus. Charig et al. (1965) referred the Avalonianus postcrania to Melanorosauridae, which was followed by Steel (1970), White (1973) and Cooper (1980, 1981). While Cooper seemed to accept the referral of carnivorous teeth to Melanorosauridae, Charig et al. correctly noted such teeth did not belong to sauropodomorphs, so presumably thought the same of the Avalonianus holotype. Van Heerden (1979) followed Charig et al.'s thought on this matter, making Avalonianus an indeterminate archosaur(-iform) but noting the associated femur resembles Riojasaurus. The issue was finally resolved in 1985, when Galton made the postcrania the type material of a new melanorosaurid sauropodomorph- Camelotia borealis, though he did not comment further on the identity of Avalonianus. Further references to Camelotia are not given here, but it is now placed as a basal sauropod or very closely related to that group, more derived than Gresslyosaurus but perhaps a melanorosaurid. Storrs (1993) also declared Avalonianus to be a nomen dubium, Glut (1997) merely called it a "thecodontian", and Naish and Martill (2007) referred it to Archosauria indet.. Olshevsky (1991) listed it as an ornithosuchid, with Picrodon herveyi as a junior synonym of the species. The most recent extensive review has been Galton's (1998) paper, where he agreed with past authors that the teeth are not sauropodomorph, are nomina dubia, and are similar enough to each other to belong to the same taxon. Galton also agreed with Newton and Seeley that the teeth were similar to cambrensis, so that he provisionally referred them to "?Megalosaurus cambrensis (Newton) of the suborder Theropoda (and possibly Infraorder Carnosauria)."
When determining the affinities of Avalonianus sanfordi, we can exclude a close relationship to Gresslyosaurus, melanorosaurids or other sauropodomorphs, as these have lanceolate teeth with enlarged apically oriented serrations. However, despite an apparent consensus the tooth is conspecific with Picrodon herveyi and both are most similar to those of "Newtonsaurus" cambrensis (which is neither Zanclodon nor Megalosaurus), this has never been supported by explicit characters. Indeed, the teeth of "Newtonsaurus" have not been shown to be diagnostic compared to the large variety of Late Triassic and Early Jurassic theropods now known. If Avalonianus did happen to be synonymous with "Newtonsaurus" (and thus its correct genus name), sanfordi has priority over cambrensis by one year, so the former cannot be a junior synonym of the latter (contra Galton). An additional issue is that while Avalonia has priority over Picrodon (thanks to Nopcsa, 1901 being the first revisor), Avalonianus does not. Thus if the genera were synonymous, Picrodon would be the correct name (contra Nopcsa and Olshevsky). Confounding matters is that sanfordi has priority over herveyi, making the correct name the unpublished new combination Picrodon sanfordi. None of this matters though, as Avalonianus and Picrodon have not been shown to be more similar to each other than they are to other archosauriforms, and indeed both have been called indeterminate by several authors. If this is true, they cannot be synonyms of each other or any other taxon (contra Galton). Thus Avalonianus, Picrodon and "Newtonsaurus" have no established relationship to each other, and placing the former two more precisely within Archosauriformes will require further study. The presence of serrations does suggest they are archosauriform, and the Late Triassic age suggests they are at least as derived as proterochampsids. It can also be noted that the presence of serrations is different from Zanclodon, suggesting no close relationship with that genus.
References- Walcott, 1890. Descriptive notes of new genera and species from the Lower Cambrian or Olenellus zone of North America. Proceedings of the United States National Museum, 12(763), 33-46.
Sanford, 1894. On the bones of an animal resembling the megalosaur, found in the Rhaetic Formation at Wedmore. Proceedings of the Somerset Archaeological Society. 40, 227-235.
Seeley, 1898. On large terrestrial saurians from the Rhaetic beds of Wedmore Hill, described as Avalonia sanfordi and Picrodon herveyi. Geological Magazine. 5(1), 1-6.
Newton, 1899. On a megalosauroid jaw from Rhaetic beds near Bridgend (Glamorganshire). Quarterly Journal of the Geological Society. 55, 89-96.
Nopcsa, 1901. A dinosaurusok atnezete es szarmazasa. Földtani Közlöny. 31, 193-224.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen. 6, 1-84.
Huene, 1905. Uber die Trias-Dinosaurier Europas. Zeitschrift der Deutschen Geologischen Gesellschaft. 57, 345-349.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [The dinosaurs of the European Triassic formations with consideration of occurrences outside Europe]. Geologische und Palaeontologische Abhandlungen. Supplement 1(1), 1-419.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Huene, 1956. Palaontologie und Phylogenie der Niederen Tetrapoden. 716 pp.
Kuhn, 1961. Fossilium Catalogus I: Animalia, Reptila. supplementum 1(2), 1-163.
Charig, Attridge and Crompton, 1965. On the origin of the sauropods and the classification of the Saurischia. Proceedings of the Linnean Society of London. 176, 197-221.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp.
White, 1973. Catalogue of the genera of dinosaurs. Annals of Carnegie Museum. 44(9), 117-155.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.
van Heerden, 1979. The morphology and taxonomy of Euskelosaurus (Reptilia: Saurischia; Late Triassic) from South Africa. Navorsinge van die Nasionale Museum, Bloemfontein. 4, 21-84.
Cooper, 1980. The first record of the prosauropod dinosaur Euskelosaurus from Zimbabwe. Arnoldia. 9(3), 1-17.
Cooper, 1981. The prosauropod dinosaur Massospondylus carinatus Owen from Zimbabwe: Its biology, mode of life and phylogenetic significance. Occasional Papers of the National Museums and Monuments of Rhodesia (series B, Natural Sciences). 6, 689-840.
Galton, 1985. Notes on the Melanorosauridae, a family of large prosauropod dinosaurs (Saurischia: Sauropodomorpha). Géobios. 18(5), 671-676.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Storrs, 1993. Terrestrial components of the Rhaetian (uppermost Triassic) Westbury Formation of southwestern Britain. In Lucas and Morales (eds.). The Nonmarine Triassic. New Mexico Museum of Natural History and Science Bulletin. 3, 447-451.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076 pp.
Galton, 1998. Saurischian dinosaurs from the Upper Triassic of England: Camelotia (Prosauropoda, Melanorosauridae) and Avalonianus (Theropoda, ?Carnosauria). Palaeontographica Abteilung A. 250, 155-172.
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, London. 164, 493-510.

Basutodon Huene, 1932
B. ferox
Huene, 1932
Norian, Late Triassic
Lower Elliot Formation, Lesotho

Holotype- (NMB R.610) tooth
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Kuhn, 1939. Saurischia. In Fossilium Catalogus I. Animalia. 87. 124 pp.
Charig, Attridge and Crompton, 1965. On the origin of the sauropods and the classification of the Saurischia. Proceedings of the Linnean Society of London. 176, 197-221.
Kuhn, 1970.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.
van Heerden, 1979. The morphology and taxonomy of Euskelosaurus (Reptilia: Saurischia; Late Triassic) from South Africa. Navorsinge van die Nasionale Museum, Bloemfontein. 4, 21-84.
Cooper, 1980. The first record of the prosauropod dinosaur Euskelosaurus from Zimbabwe. Arnoldia Zimbabwe. 9, 1-17.
Cooper, 1981. The prosauropod dinosaur Massospondylus carinatus Owen from Zimbabwe: Its biology, mode of life and phylogenetic significance. Occasional Papers of the National Museums and Monuments of Rhodesia (series B, Natural Sciences). 6, 689-840.
Hopson, 1984. Late Triassic traversodont cynodonts from Nova Scotia and Southern Africa. Palaeontologia Africana. 25, 181-201.
Kitching and Raath, 1984. Fossils from the Elliot and Clarens Formations (Karoo Sequence) of the Northeastern Cape, Orange Free State and Lesotho, and a suggested biozonation based on tetrapods. Palaeontologia Africana. 25, 111-125.
Anderson, Anderson and Cruikshank, 1998. Late Triassic ecosystems of the Molteno/Lower Elliot biome of southern Africa. Palaeontology. 41(3), 387-421.
Olsen and Galton, 1984. A review of the reptile and amphibian assemblages from the Stormberg of Southern Africa with special emphasis on the footprints and the age of the Stormberg. Palaeontologia Africana. 25, 87-110.
Galton and van Heerden, 1998. Anatomy of the prosauropod dinosaur Blikanasaurus cromptoni (Upper Triassic, South Africa) with notes on the other tetrapods from the Lower Elliot Formation. Palaontologische Zeitschrift. 72, 163-177.
Ray and Chinsamy, 2002. A theropod tooth from the Late Triassic of southern Africa. Journal of Biosciences. 27(3), 295-298.
Knoll, 2004. Review of the tetrapod fauna of the "Lower Stormberg Group" of the main Karoo Basin (southern Africa): Implication for the age of the Lower Elliot Formation. Bulletin de la Societe Geologique de France. 175(1), 73-83.

"Cinizasaurus" Heckert, 1997 vide Nesbitt, Irmis and Parker, 2007
"C. hunti" Heckert, 1997 vide Nesbitt, Irmis and Parker, 2007
Late Carnian, Late Triassic
Mesa Redondo Member of the Chinle Formation, New Mexico, US
Material
- (NMMNH P-18400) vertebrae, tibia, fragments
Comments- Originally described as a new genus of theropod in Heckert's (1997) unpublished thesis, the specimen was later claimed to be indeterminate when published without a name by Heckert et al. (2000). Nesbitt et al. (2007) noted the vertebrae could not be distinguished from non-theropods, while the tibia lacks a cnemial crest and lateral and medial condyles, unlike dinosauriforms and theropods respectively. They assigned it to Archosauriformes indet..
Although the name "Cinizasaurus hunti" was originally used in thesis, and thus not available for use in this website, it was later published by Nesbitt et al. (2007).
References- Heckert, 1997. Litho- and biostratigraphy of the Lower Chinle Group, east-central Arizona and west-central New Mexico, with a description of a new theropod (Dinosauria: Theropoda) from the Bluewater Creek Formation. Unpublished MSc Thesis, University of New Mexico. 278 pp.
Heckert, Lucas and Sullivan, 2000. Triassic dinosaurs of New Mexico. New Mexico Museum of Natural History & Science Bulletin. 17, 17-26.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.

"Likhoelesaurus" Ellenberger, 1970
"L. ingens" Ellenberger, 1970
= "Likhoelesaurus ferox" Ellenberger, 1972
Norian, Late Triassic
Lower Elliot Formation, Lesotho

Material- (?LES coll.) five teeth (70 mm)
Comments- Ellenberger and Ginsberg (1966) mentioned carnivorous dinosaur teeth from the Lower Elliot Formation, which were later called "Likhoelesaurus ingens" by Ellenberger (1970) and referred to Ornithosuchidae. These were not described properly though, making the name a nomen nudum. Ellenberger (1972) referred to "Likhoelesaurus ferox" as a "giant carnosaur" from zone A/5 of his lower red beds, illustrating five associated recurved teeth in a plate. This was another nomen nudum, with no explanation of the different species name (it may involve Basutodon ferox, but even if "Likhoelesaurus" were officially named, Basutodon would have priority). Kitching and Raath (1984) suggested it may be a junior synonym of Basutodon ferox. It was listed as a teratosaurid theropod by Chure and McIntosh (1989) and a melanorosaurid by Olshevsky (1991). Glut (1997) listed it as ?Theropoda incertae sedis, listed "?bones" among the remains and stated the teeth were 70 mm long. Knoll (2004) discusses the material under Rauisuchia, but notes it could be theropod as well. If the material is not lost, it is presumably held with the rest of Ellenberger's material in the "Collection Paul Ellenberger - Pistes du Stormberg", stored in the Laboratoire de Paleontologie, Institut de Sciences de l’Evolution, Universite Montpellier II.
With no published description, and only one undetailed photograph, the phylogenetic position of "Likhoelesaurus" remains uncertain. While their recurved morphology excludes referral to Melanorosauridae or any other sauropodomorph clade, differences between the teeth of carnivorous dinosaurs, crutotarsans (including ornithosuchids and teratosaurs) and other carnivorous archosauriforms have yet to be studied. Indeed, they are only assumed to be archosauriform here due to past identifications, since the presence of serrations has not been confirmed. Synonymy with Basutodon is possible, but no shared derived characters have been suggested, Basutodon itself is probably undiagnostic and multiple archosauriform taxa are known from other Late Triassic localities.
References- Ellenberger and Ginsberg, 1966. Le gisement de Dinosauriens triasiques de Maphutseng (Basutoland) et l'origine des Sauropodes [The Triassic dinosaur locality of Maphutseng (Basutoland) and the origin of sauropods]. Comptes Rendus de l'Académie des Sciences à Paris, Série D. 262, 444-447.
Ellenberger, 1970. Les niveaux paléontologiques de première apparition des mammifères primoridaux en Afrique du Sud et leur ichnologie. Establissement de zones stratigraphiques detaillees dans le Stormberg du Lesotho (Afrique du Sud) (Trias Supérieur à Jurassique) [The paleontological levels of the first appearance of primordial mammals in southern Africa and their ichnology. Establishment of detailed stratigraphic zones in the Stormberg of Lesotho (southern Africa) (Upper Triassic to Jurassic). in Haughton (ed.). Second Symposium on Gondwana Stratigraphy and Paleontology, International Union of Geological Sciences. Council for Scientific and Industrial Research, Pretoria. 343-370.
Ellenberger, 1972. Contribution à la classification des Pistes de Vertébrés du Trias: Les types du Stormberg d'Afrique du Sud (I). Palaeovertebrata. 104, 152 pp.
Kitching and Raath, 1984. Fossils from the Elliot and Clarens Formations (Karoo Sequence) of the Northeastern Cape, Orange Free State and Lesotho, and a suggested biozonation based on tetrapods. Palaeontologia Africana. 25, 111-125.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076 pp.
Knoll, 2004. Review of the tetrapod fauna of the "Lower Stormberg Group" of the main Karoo Basin (southern Africa): Implication for the age of the Lower Elliot Formation. Bulletin de la Societe Geologique de France. 175(1), 73-83.

"Megalosaurus" cloacinus Quenstedt, 1858
= Plateosaurus cloacinus (Quenstedt, 1858) Huene, 1905
= Gresslyosaurus cloacinus (Quenstedt, 1858) Huene, 1932
Rhaetian, Late Triassic
Rhatsandstein, Germany

Holotype- (SMNH 52457) tooth
Reference- Quenstedt, 1858. Der Jura. H. Laupp'schen, Tübingen. 842 pp.

"Megalosaurus" obtusus

Palaeosauria Romer, 1956
Palaeosauridae Huene, 1926
Paleosaurus Riley and Stutchbury, 1840
= "Palaeosaurus" Riley and Stutchbury, 1836 (preoccupied Geoffroy Saint-Hilaire, 1833)
= "Paleosaurus" Riley and Stutchbury, 1837
= Palaeosauriscus Kuhn, 1959
Comments- This genus has an extremely complicated history due to several nomina nuda and homonyms. Geoffroy Sant-Hillaire named Palaeosaurus in 1836, which is currently a junior synonym of the teleosaurid Steneosaurus priscus. Riley and Stutchbury (1836) used the names "Palaeosaurus cylindricum" and "P. platyodon" in an abstract, but these are nomina nuda. Those same authors wrote another short paper in 1837 in which the genus for those speciesw was spelled both "Palaeosaurus" and "Paleosaurus", though these were also nomina nuda. The taxa were finally validly published in 1840 under the genus Paleosaurus, though this was misspelled Palaeosaurus by many later authors. Another Palaeosaurus was named by Sternberg (1940), though this was renamed Sphenosaurus by Meyer (1847) and is currently classified as a procolophonid. Kuhn noticed Geoffroy Saint-Hillaire's Palaeosaurus and thought this caused Riley and Stutchbury's genus to be preoccupied, so proposed the replacement name Palaeosauriscus. Yet as Paleosaurus is spelled differently than Palaeosaurus and is not the valid name for any other genus, it does not need to be replaced. Though originally based on archosaurian teeth, the genus and its spelling variants were widely used until the 1970s for basal sauropodomorphs based on the specimen later named Efraasia.
References- Riley and Stutchbury, 1836. A description of various fossil remains of three distinct saurian animals discovered in the authumn of 1834, in the Magnesian conglomerates of Diirdham Down, near Bristol. Proc. Geol. Soc. London 2, 397-399.
Riley and Stutchbury, 1837. On an additional species of the newly-discovered saurian animals in the Magnesian Conglomerate of Durdham Down, near Bristol. Annual Report of the British Association for the Advancement of Science, Transactions of the Sections. 1836, 90-94.
Fitzinger, 1840. Ãber Palaeosaurus sternbergii, eine neue Gattung vorweltlicher Reptilien und die Stellung dieser Thiere im Systeme Ãberhaupt. Wiener Mus. Annalen II, 175-187.
Riley and Stutchbury, 1840. A description of various fossil remains of three distinct saurian animals discovered in the authumn of 1834, in the Magnesian conglomerates of Diirdham Down, near Bristol. Trans. Geol. Soc. London 5(2), 349-357.
Kuhn, 1959. Ein neuer Microsaurier aus dem deutschen Rotliegenden. N. Jb. Geol. Palaeontol. Mh. 1959: 424-426.
P. cylindrodon Riley and Stutchbury, 1840
= "Palaeosaurus cylindricum" Riley and Stutchbury, 1836
= "Palaeosaurus cylindrodon" Riley and Stutchbury, 1837
= "Paleosaurus cylindrodon" Riley and Stutchbury, 1837
= Thecodontosaurus cylindrodon (Riley and Stutchbury, 1840) Huene, 1908
= Thecodontosaurus "cylindricum" (Riley and Stutchbury, 1836) Huene, 1914
= Palaeosauriscus cylindrodon (Riley and Stutchbury, 1840) Kuhn, 1959
Early Norian, Late Triassic
Durdham Down, England

Holotype- (destroyed) tooth
References- Riley and Stutchbury, 1836. A description of various fossil remains of three distinct saurian animals discovered in the authumn of 1834, in the Magnesian conglomerates of Diirdham Down, near Bristol. Proc. Geol. Soc. London 2, 397-399.
Riley and Stutchbury, 1837. On an additional species of the newly-discovered saurian animals in the Magnesian Conglomerate of Durdham Down, near Bristol. Annual Report of the British Association for the Advancement of Science, Transactions of the Sections. 1836, 90-94.
Riley and Stutchbury, 1840. A description of various fossil remains of three distinct saurian animals discovered in the authumn of 1834, in the Magnesian conglomerates of Diirdham Down, near Bristol. Trans. Geol. Soc. London 5(2), 349-357.
Huene, 1908. Die Dinosaurier der Europaiaschen Triasformation mit Berucksichtiging der aussereuropaischen Vorkommnisse. Geol. Paleont. Abhandl. Suppl. 1, 1-419.
Huene, 1914. Nachtrige zu meinen fruheren Beschreibungen triassischer Saurischia. Geol. Palaeontol. Abhandl. 13: 69-82.
Huene, 1926. Vollstandige Osteologie eines Plateosauriden aus dem schwabischen Trias. Geol. Palaeontol. Abhandl. 15:129-179.
Romer, 1956. Osteology of the Reptiles. University of Chicago Press, Chicago.
Kuhn, 1959. Ein neuer Microsaurier aus dem deutschen Rotliegenden. N. Jb. Geol. Palaeontol. Mh. 1959: 424-426.
Benton, Juul, Storrs and Galton, 2000. Anatomy and systematics of the prosauropod dinosaur Thecodontosaurus antiquus from the Upper Triassic of southern England. J. Vertebr. Palaeontol. 20, 77-108.

Paleosaurus? subcylindrodon (Huene, 1908) Huene, 1932
= Thecodontosaurus "subcylindrodon" Huene, 1905
= Thecodontosaurus subcylindrodon Huene, 1908
= Palaeosauriscus subcylindrodon (Huene, 1908) Kuhn, 1965
Early Carnian, Late Triassic
Schilfsandstein, Germany

Holotype- (SMNS 52456) tooth
Comments- Olshevsky said Galton (1985) considered this a herrerasaurid, but he only wrote it was from a carnivorous archosaur.
References- Huene, 1905. Uber die Trias-Dinosaurier Europas. Z. Dtsch. Geol. Ges. Mb. 57: 345-349.
Huene, 1908. Die Dinosaurier der Europaiaschen Triasformation mit Berucksichtiging der aussereuropaischen Vorkommnisse. Geol. Paleont. Abhandl. Suppl. 1, 1-419.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Kuhn, 1965.
Galton, 1985. The poposaurid thecodontian Teratosaurus suevicus v. Meyer, plus reffered specimens mostly based on prosauropod dinosaurs, from the Middle Stubensandstein (Upper Triassic) of Nordwurttemberg. Stuttgarter Breitage Naturkunde Ser. B. 116, 1-29.

Picrodon Seeley, 1898
P. herveyi Seeley, 1898
= Avalonia herveyi (Seeley, 1898) Nopcsa, 1901
Rhaetian, Late Triassic
Westbury Formation, England
Holotype
- (BMNH R2875) fragmented dentaries , tooth (23 x 9.5 x ? mm)
Comments- This taxon shares much of its history with Avalonianus sanfordi. Kuhn (1970) listed it as an ornithosuchid
References- Nopcsa, 1901. A dinosaurusok atnezete es szarmazasa. Földtani Közlöny. 31, 193-224.
Kuhn, 1970.
Romer, 1976. Osteology of the Reptiles. University of Chicago Press. 772 pp.

"Thecodontosaurus" elizae

Zanclodon? bavaricus

Zatomus

unnamed archosauriform (Jenkins, Shubin, Amaral, Gatesy, Schaff, Downs, Clemmenssen, Bonde, Davidson and Osbaeck, 1993)
Late Norian-Early Rhaetian, Late Triassic
Orsted Dal Member of the Fleming Fjord Formation, Greenland, Denmark
Material
- (MCZ coll.) vertebrae including dorsal centra, ribs, partial pelvis, partial hindlimb including femur (330 mm), phalanges and pedal unguals
Comments- Jenkins et al. (1993) reported theropods from this formation, but the eventual more detailed version of Jenkins et al. (1994) reported only a single specimen. This was referred to Theropoda based on- constricted dorsal vertebral centra; slender and hollow limb elements; mediolaterally compressed and curved pedal unguals. Clammensen et al. (2015) reported "classification of the remains attributed to theropods is not supported by any clear diagnostic theropodan feature and might alternatively be phytosaurian in origin."
References- Jenkins, Shubin, Amaral, Gatesy, Schaff, Downs, Clemmenssen, Bonde, Davidson and Osbaeck, 1993. A Late Triassic continental vertebrate fauna from the Fleming Fjord Formation, Jameson Land, east Greenland. New Mexico Museum of Natural History and Science Bulletin. 3, 74.
Jenkins, Shubin, Amaral, Gatesy, Schaff, Clemmensen, Rowns, Davidson, Bonde and Osbaeck, 1994. Late Triassic continental vertebrates and depositional environments of the Fleming Fjord Formation, Jameson Land, east Greenland. Meddelelser om Grønland, Geoscience. 32, 1-25.
Clemmensen, Milàn, Adolfssen, Estrup, Frobøse, Klein, Mateusand Wings, 2015. The vertebrate-bearing Late Triassic Fleming Fjord Formation of central east Greenland revisited: Stratigraphy, palaeoclimate and new palaeontological data. Geological Society Special Publications. 434. doi: 10.1144/SP434.3

unnamed archosauriform (Heckert et al., 2000)
Late Carnian, Late Triassic
Mesa Redondo Member of the Chinle Formation, New Mexico, US
Material
- (NMMNH P-18401) dorsal vertebrae
Comments- Heckert et al. (2000) referred these vertebrae to an unnamed new theropod taxon with a ventral keel, though Nesbitt et al. (2007) noted some pseudosuchians had ventral keels and the vertebrae shared no unique characters with those of theropods. They assigned them to Archosauriformes indet..
References- Heckert, Lucas and Sullivan, 2000. Triassic dinosaurs of New Mexico. New Mexico Museum of Natural History & Science Bulletin. 17, 17-26.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.

Proterochampsia Bonaparte, 1971
Definition- (Proterochampsa barrionuevoi <- Crocodylus niloticus) (modified from Kischlat, 2000)
Other definitions- (Proterochampsa barrionuevoi <- Euparkeria capensis, Erythrosuchus africanus, Passer domesticus, Crocodylus niloticus) (Nesbitt, 2011)

Proterochampsidae Sill, 1967
= Rhadinosuchidae Hofstetter, 1955

Rhadinosuchinae Hofstetter, 1955 vide Ezcurra, DeSojo and Rauhut, 2015
Definition- (Rhadinosuchus gracilis, Chanaresuchus bonapartei <- Cerritosaurus binsfeldi, Tropidosuchus romeri, Doswellia kaltenbachi) (Ezcurra, DeSojo and Rauhut, 2015)

Crurotarsi Sereno and Arcucci, 1990
Definition- (Parasuchus hislopi + Ornithosuchus longidens + Prestosuchus chiniquensis + Crocodylus niloticus) (modified from Sereno, 1991)
Other definitions- (Belodon plieningeri + Ornithosuchus longidens + Crocodylus niloticus) (modified from Kischlat, 2000)
(Phytosaurus cylindricodon + Ornithosuchus longidens + Prestosuchus chiniquensis+ Crocodylus niloticus) (modified from Benton, 2004)
(fully rotary, hemicylindrical, fibulocalcaneal crurotarsal articulation as in Caiman crocodilus) (Gauthier et al., 2004)
(Rutiodon carolinensis + Crocodylus niloticus) (Nesbitt, 2011)

Parasuchia Huxley, 1875
Definition- (Parasuchus hislopi <- Aetosaurus ferratus, Crocodylus niloticus) (suggested)
= Belodontia Marsh, 1895
Definition- (Belodon plieningeri <- Aetosaurus ferratus, Crocodylus niloticus) (modified from Kischlat, 2000)
= Phytosauria Baur, 1895
Definition- (Paleorhinus bransoni + Phytosaurus cylindricodon) (modified from Doyle and Sues, 1995)
Other definitions- (Rutiodon carolinensis <- Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, Ornithosuchus longidens, Crocodylus niloticus) (Nesbitt, 2011)
= Phytosauriformes Hay, 1830
= Phytosauromorphi Hay, 1930

"Zanclodon" arenaceus

Phytosauridae Jaeger, 1828
Definition- (Phytosaurus cylindricodon + Angistorhinus grandis + Leptosuchus studeri + Mystriosuchus westphali) (suggested)
Other definitions- (Angistorhinus grandis + Rutiodon carolinensis + Mystriosuchus planirostris + Nicrosaurus kapffi + Pseudopalatus pristinus) (modified from Doyle and Sues, 1995)
(Angistorhinus grandis + Leptosuchus studeri + Mystriosuchus westphali) (modified from Stocker, 2010)
= Parasuchidae Lydekker, 1885
Definition- (Parasuchus hislopi + Wannia scurriensis + Mystriosuchus planirostris) (Kammerer, Butler, Bandyopadhyay and Stocker, 2015)
= Belodontidae Cope, 1871
Comments- Kammerer et al. (2015) argued neither Doyle and Sues' (1995) or Stocker's (2010) definitions of Phytosauridae were valid, because Phytosaurus could belong outside either of these apparently equivalent parasuchian clades. Furthermore, I note they are invalid because the eponym Phytosaurus cylindricodon is not used as an internal specifier. Kammerer et al. say "the argument could be made that regardless of the status of Phytosaurus as a nomen dubium, if it exhibits the diagnostic features of the larger clade currently called Phytosauridae, then the latter taxon could be retained", but go on to define the later-named family Parasuchidae in a way that would include Phytosaurus. Since there's no reason to keep the first definitions of Phytosauridae, as they don't use Phytosaurus, I suggest the family be redefined to include Phytosaurus cylindricodon but encompass all known parasuchians.

Belodon

Centemodon

Clepsysaurus
C. pennsylvanicus
Clepsysaurus? fraserianus (Cope, 1878a) Hay, 1930
= Paleosaurus fraserianus Cope, 1878a (as Palaeosaurus fraserianus)
= Thecodontosaurus fraserianus (Cope, 1878a) Hay, 1902
= Palaeosauriscus fraserianus (Cope, 1878a) Kuhn, 1965
Norian, Late Triassic
New Oxford Formation, Pennsylvania, US
Holotype
- (AMNH 1861) tooth (20x6.5x? mm)
Comments- Cope (1878a) described this tooth (which was first presented the year prior) as a new species of Palaeosaurus, at the time a common misspelling of Paleosaurus. Olshevsky (2000) was the first to correct the genus' spelling in this binomial. Note there is a valid genus Palaeosaurus (Geoffroy Sant-Hillaire, 1836; which is currently a junior synonym of the teleosaurid Steneosaurus) however, which caused Kuhn (1965) to incorrectly think Paleosaurus was preoccupied since its spelling is so similar. Thus he referred all Paleosaurus species to his new genus Palaeosauriscus, but this is unnecessary according to the ICZN. Paleosaurus itself is based on P. cylindrodon, an archosauriform tooth of uncertain affinities from the Norian of England which differs from fraserianus in having elongate and oblique serrations, being less recurved, and having a more tapered distal edge in section.
fraserianus a dinosaur? Nopsca (1901) was the first to assign the species explicitly to Dinosauria or Theropoda, assigning it to a subfamily Anchisauridae [sic] within Megalosauridae, but his anchisaurids consisted largely of basal sauropodomorphs and Triassic carnivorous archosauriform teeth. Hay (1902) had a similar concept for Anchisauridae within his Theropoda, similarly placing fraserianus there though assigning it to Thecodontosaurus, as he synonymized the genus with Paleosaurus. Note Colbert and Chaffee (1941) wrongly cited Cope (1878b) as using the combination Thecodontosaurus fraserianus, but that work only uses Thecodontosaurus for T. gibbidens. Hay (1930) retained fraserianus in Anchisauridae and Theropoda, but now placed it in the genus Clepsysaurus (a parasuchian), perhaps based on the similarity noted by Huene (see below). Steel (1970) referred it to his theropodan Ornithosuchidae, which besides Ornithosuchus contained Teratosaurus, Triassic carnivorous archosauriform teeth, and basal sauropodomorph remains incorrectly associated with the latter. The most confusing generic assignment has been that of Olshevsky (1991, 2000), who made fraserianus a junior synonym of Anchisaurus polyzelus, which is from the much later (Pliensbachian) Portland Formation of Connecticut. fraserianus is quite unlike sauropodomorph teeth (including Thecodontosaurus and Anchisaurus) in being highly recurved, with an unconstricted base, little labiolingual compression, and small serrations which are perpendicular to the tooth axis. The connection was maintained through history largely via ignorance of fraserianus' actual morphology in addition to continued confusion of Paleosaurus with Thecodontosaurus and Efraasia.
fraserianus a parasuchian? Lesley (1889) may be the first author to suggest fraserianus is parasuchian, albeit without evidence. Huene (1921) has been the only author to illustrate fraserianus, and briefly described the specimen as well. Huene convincingly illustrated the similarity with other New Oxford parasuchian teeth. He suggested fraserianus was synonymous with Clepsysaurus? veatleianus and/or Rutiodon carolinensis from the same formation. Colbert and Chaffee (1941) made fraserianus a junior synonym of Clepsysaurus pennsylvanicus, based on geography. Of Norian archosauriforms which have recurved teeth with small serrations, only some proterochampsids and phytosaurid parasuchians are reported to have reduced labiolingual compression as in fraserianus (80% of FABL). While proterochampsid teeth remain largely undescribed, they are exclusively South American, so are an unlikely identification for fraserianus. Indeed, phytosaurid material is common in the New Oxford Formation, with Rutiodon carolinensis the only currently recognized valid taxon. It's therefore possible Huene was correct and that fraserianus is synonymous with Rutiodon, but the most recent review also suggested the presence of a larger, poorly characterized form (e.g. SMP VP-36; YPM-PU 11544). The tooth of fraserianus is much smaller than these latter elements, but could be ontogenetically young as well. Unfortunately, heterodonty is so great among phytosaurid dentitions, few of which have been described in detail, that it is not currently possible to assign isolated teeth to particular genera or species. Thus fraserianus remains Phytosauridae indet., and is here placed questionably in Clepsysaurus as that is the only phytosaurid genus prior authors have referred the species to.
References- Cope, 1878a. On some saurians found in the Triassic of Pennsylvania, by C. M. Wheatley. Proceedings of the American Philosophical Society. 17(100), 231-232.
Cope, 1878b. Triassic saurians from Pennsylvania. The American Naturalist. 12, 58.
Lesley, 1889. A Dictionary of the Fossils of Pennsylvania and Neighboring States Named in the Reports and Catalogues of the Survey. Volume 2. The Board of Commissioners for the Geological Survey, Harrisburg. 914 pp.
Nopcsa, 1901. A dinosaurusok atnezete es szarmazasa. Földtani Közlöny. 31, 193-224.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America. United States Geological Survey Bulletin. 179, 868 pp.
Huene, 1921. Reptilian and stegocephalian remains from the Triassic of Pennsylvania in the Cope collection. Bulletin American Museum of Natural History. 44(19), 561-574.
Hay, 1930. Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution of Washington. 390(II), 1-1074.
Colbert and Chaffee, 1941. The type of Clepsysaurus pennsylvanicus and its bearing upon the genus Rutiodon.
Kuhn, 1965. Saurischia (Supplementum 1). In Fossilium Catalogus 1. Animalia. 109, 94 pp.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings. 3, 1-157.
C? veatleianus

"Megalosaurus" terquemi

Palaeoctanus
P. appalachianus
P? aulacodus
P? dumblianus
P? orthodon

"Paleosaurus" platyodon

"Paleosaurus" stricklandi

Rileyasuchus
R. bristolensis
R? stutchburyi

Rutiodon? leaii

Suchoprion
S. cephodon
S? sulcidens

Archosauria Cope, 1869
Definition- (Crocodylus niloticus + Megalosaurus bucklandii) (modified from Kischlat, 2000)
Other definitions- crown(Caiman crocodilus + Compsognathus longipes + Vultur gryphus) (Gauthier et al., 2004)
(Crocodylus niloticus + Passer domesticus) (Sereno, 2004; modified from Gauthier and Padian, 1985)
= Hyperosauria Jaekel, 1911
= Proquadrata Walker, 1977
= Nidosuchia Whetstone and Whybrow, 1983
= Neoarchosauria Benton, 1985
= Avesuchia Benton, 1985
Definition- (Crurotarsi + Avemetatarsalia) (Benton, 1985)
References- Gauthier, de Queiroz, Joyce, Parham, Rowe and Clarke, 2004. A phylogenetic nomenclature for the major clades of Amniota Haeckel 1866, with emphasis on non-avian Reptilia Laurentus 1768. First International Phylogenetic Nomenclature Meeting, abstracts. 24.

Archosauria indet. (Seeley, 1887)
Late Albian, Early Cretaceous
Gault Clay (reworked into Cambridge Greensand), England
Lectotype
- (SMC B58402) incomplete sacral vertebra (11 mm)
Diagnosis- (after Barrett and Evans, 2002) femoral head strongly curved anteriorly; very strong trochanteric crest and intertrochanteric fossa.
Comments- Patricosaurus merocratus was based on two unassociated specimens, a sacral vertebra found before 1859 and a proximal femur (SMC B58401) discovered in the 1880's. Seeley (1887) believed both belonged to the same taxon since he thought "there was little chance of any remains of two lizards occurring" in the Cambridge Greensand, but stated the femur could remain as the type if the vertebra was found to not belong to the same taxon. Barrett and Evans (2002) redescribed the specimens and removed the vertebra from Patricosaurus, believing it to be either a crocodylian or an ornithischian. They thus referred it to Archosauria indet..
References- Seeley, 1887. On Patricosaurus merocratus, Seeley, a lizard from the Cambridge Greensand, preserved in the Woodwardian Museum of the University of Cambridge. Quarterly Journal of the Geological Society of London. 43, 216-220.
Barrett and Evans, 2002. A reassessment of the Early Cretaceous reptile 'Patricosaurus merocratus' Seeley from the Cambridge Greensand, Cambridgeshire, UK. Cretaceous Research. 23, 231-240.

unnamed archosaur (Lapparent, Claracq and Nougarède, 1958)
Late Triassic
Lower Sandstone Member of Lower Zarzaitine Formation, Algeria
Material
- (MNNHM coll.) tooth (43x16x? mm)
(MNNHM coll.) tooth (22x9x? mm)
Comments- These ware described by Lapparent et al. (1958) as a teratosaurid theropod, and later (1960) as Teratosaurus sp.. Both are recurved and serrated on both carinae, typical of basal pseudosuchians like Teratosaurus as well as basal theropods. They are of similar shape and from the same locality, but have no described diagnostic characters and may belong to different taxa.
References- Lapparent, Claracq and Nougarède, 1958. Nouvelles découvertes de Vertébrés dans les séries continentales au Nord d'Edjeleh (Sahara central) [New discoveries of vertebrates in the continental series north of Edjeleh (central Sahara)]. Comptes Rendus de l'Académie des Sciences à Paris. 247, 2399-2402.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.

Archosauria indet. (Kessler and Jurcsak, 1984)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania

Material- (MTCO 14422; = MTCO-P 1503) incomplete long bone
(MTCO 17956; = MTCO-P 6966; paratype of Limnornis corneti) long bone shaft
Comments- MTCO 14422 was discovered in 1978 and described by Kessler and Jurcsak (1984 and subsequent publications) as a humerus of 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. Dyke et al. (2011) redescribed it, noting it cannot even be definitively identified as a humerus and could only refer it to Archosauria indet. as a long bone.
MTCO 17956 was originally (Kessler and Jurcsak, 1984) decribed as an ulnar shaft and paratype of Limnornis corneti (later renamed Palaeocursornis corneti), and later (Kessler and Jurcsak, 1986) a paratype of Eurolimnornis corneti. Hope (2002) noted it couldn't be referred definitively to either species, and Dyke et al. (2011) couldn't identify it past Archosauria indet., as the shaft of a long bone. The supposed nodes for secondary feathers are largely taphonomic.
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.
Hope, 2002. The Mesozoic radiation of Neornithes. In Chiappe and Witmer (eds). Mesozoic birds: Above the heads of dinosaurs. Berkeley: University of California Press. 339-388.
Dyke, Benton, Posmosanu and Naish, 2011. Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology. 54(1), 79-95.

Archosauria indet. (Long and Murray, 1995)
Late Carnian, Late Triassic
Mesa Redondo Member of the Chinle Formation, Arizona, US

Material- (MNA V3091; = V2777 in Long and Murray, 1995) dorsal centrum
(UCMP 138591) partial sacrum
(UCMP 177317) dorsal centrum
(UCMP 178047) incomplete sacrum
(UCMP 178048) incomplete sacrum
(UCMP 178049) incomplete sacrum
Comments- Originally identified as Ceratosauria indet. by Long and Murray (1995), then made paratypes of Camposaurus by Hunt et al. (1998), they were relegated to Archosauria indet. by Nesbitt et al. (2007). They cannot be distinguished from shuvosaurids and other dinosauriforms.
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the western United States. Geobios. 31(4), 511-531.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

Archosauria indet. (Heckert, Lucas and Sullivan, 2000)
Late Carnian, Late Triassic
Santa Rosa Formation of the Dockum Group, New Mexico, US
Material
- (NMMNH P-13006) two fused sacral centra
(NMMNH P-25749) pubis, fragmentary femur(?)
(NMMNH P-25750) metatarsals
Comments- Heckert et al. (2000) considered these specimens to be theropod and probably coelophysoid, in part based on the hollow sacral centra. Nesbitt et al. (2007) notes that the latter are found in shuvosaurids too, and that the pubis lacks an acetabular rim, so ma be from a more basal archosaur. They assign them to Archosauria indet..
References- Heckert, Lucas and Sullivan, 2000. Triassic dinosaurs of New Mexico. New Mexico Museum of Natural History & Science Bulletin. 17, 17–26.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

Archosauria indet. (Heckert, Zeigler, Lucas, Rinehart and Harris, 2000)
Norian, Late Triassic
Petrified Forest Member of Chinle Formation, New Mexico
Material
- (NMMNH P-29996) distal caudal vertebra
(NMMNH P-33691) dorsal vertebra
Comments- Heckert et al. (2000) assigned the dorsal vertebra to Eucoelophysis sp., but it is not identifiable past Archosauria (Nesbitt et al., 2007). The distal caudal vertebra is indistinguishable from Coelophysis and catalogued as Theropoda, but is also not identifiable past Archosauria (Nesbitt et al., 2007).
References- Heckert, Zeigler, Lucas, Rinehart and Harris, 2000. Preliminary description of coelophysoids (Dinosauria: Theropoda) from the Upper Triassic (Revueltian: early-mid Norian) Snyder quarry, north-central New Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 27–32.
Heckert, Zeigler, Lucas and Rinehart, 2003. Coelophysids (Dinosauria: Theropoda) from the Upper Triassic (Revueltian) Snyder quarry. New Mexico Museum of Natural History and Science Bulletin. 24, 127–132.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

undescribed archosaur (Nesbitt, 2001)
Anisian, Middle Triassic
Holbrook Member of the Moenkopi Formation, Arizona, US

Material- distal pubis
Comments- Nesbitt mentions this as a possible herrerasaurid, which would be notable as dinosaurs are otherwise unknown from Middle Triassic strata. Nesbitt (pers. comm, 2010) now believes the morphology is equivocal between poposauroids and basal theropods, suggesting it should be referred to Archosauria incertae sedis.
Reference- Nesbitt, 2001. New fossil vertebrate material from the Holbrook Member, Moenkopi Formation (Middle Triassic) from Northern Arizona. Journal of Vertebrate Paleontology. 21(3), 83A.

Archosauria indet. (Hunt, 1994)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US
Material
- (NMMNH P4126) proximal femur (Heckert et al., 2000)
(NMMNH P4375) vertebral fragment (Hunt, 1994)
(NMMNH P4380) vertebral fragment (Hunt, 1994)
(NMMNH P4415; paratype of Chindesaurus bryansmalli) proximal femur (Hunt, 1994)
(NMMNH P4440) central fragment (Hunt, 1994)
(NMMNH P4547) phalanx (Hunt, 1994)
(NMMNH P4666) two pubic fragments (Hunt, 1994)
(NMMNH P16607) tooth fragments (Hunt, 1994)
(NMMNH P16656; paratype of Chindesaurus bryansmalli) dorsal centra, caudal centra (Hunt, 1994)
(NMMNH P16946) two centrum fragments (Hunt, 1994)
(NMMNH P17134) dorsal vertebrae, caudal vertebrae, pelvic fragments (Hunt, 1994)
(NMMNH P17154) three centra (Hunt, 1994)
(NMMNH P17258) vertebrae, scapula, fragments (Hunt, 1994)
(NMMNH P17325; paratype of Chindesaurus bryansmalli) dorsal centrum (Hunt, 1994)
(UCM 47221; in part; holotype of Gojirasaurus quayi) tooth, proximal cervical rib, mid dorsal rib, mid dorsal rib shaft, proximal posterior dorsal rib, two partial gastralia, proximal chevron, scapula (314 mm), partial metatarsal (Hunt, 1994)
(UMMP 7274) two dorsal centra (Hunt, 1994)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, Texas, US

(TTUP 11045; = TTUP 9281 in part; paratype of Shuvosaurus inexpectatus) proximal scapula (Chatterjee, 1993)
Comments- Chatterjee (1993) referred a scapula (then part of TTUP 9281; since separated as TTUP 11045) to his new supposed ornithomimosaur Shuvosaurus, but the latter ended up being a pseudosuchian. The element doesn't resemble shuvosaurid scapulae (Long and Murry, 1995), and is too small to belong to any Shuvosaurus individuals preserved in the quarry (Lehane, 2005). Lehane ascribed it to Theropoda indet., and indeed there is a small coelophysoid femur in the quarry, but it also resembles other small taxa from the quarry such as silesaurids. While Martz et al. (2013) retained its referral to Shuvosaurus, they didn't comment further and this seems unlikely based on the slender shaft which is more like avemetatarsalians.
NMMNH P4415, P4666, P16607, P16946, P17134, P17258, UCM 47221 and UMMP 7274 were originally referred to Hunt's (1994) new taxon of herrerasaurid "Revueltoraptor lucasi", called herrerasaurid A by Hunt et al. (1998). UCM 47221 (which also includes dorsal material, a pubis and a tibia) was made the holotype of the coelophysoid Gojirasaurus quayi by Carpenter (1997) after being described by Parrish and Carpenter (1986). NMMNH P4415 and P16656 were referred to Chindesaurus by Long and Murry (1995). Nesbitt et al. (2007) found that the referred specimens noted by Hunt could not be assigned to a particular taxon, nor could the Gojirasaurus material except for the dorsal vertebrae (Shuvosaurus) and the pubis and tibia (a possibly indeterminate coelophysoid). The remaining parts of the Gojirasaurus/'Revueltoraptor" holotype and the referred "Revueltoraptor" material are possibly shuvosaurid or theropod, and are listed here.
NMMNH P4375, P4380, P4440 and P4547 were originally referred to Hunt's (1994) new taxon of herrerasaurid "Comanchesaurus kuesi", called herrerasaurid B by Hunt et al. (1998). Nesbitt et al. (2007) found they could not be assigned to a particular archosaurian taxon. Heckert et al. (2000) described and illustrated NMMNH P4415 and P4126 as hererrasaurid B, noted to be similar to Chindesaurus in their medially angled and squared-off femoral heads, but these characters are also seen in shuvosaurids and other basal dinosaurs.
NMMNH P16656 consists of dorsal and caudal centra which were referred to Chindesaurus by Murry and Long (1989) and Long and Murry (1995) and made the basis of the unnamed herrerasaurid C by Hunt (1994) and Hunt et al. (1998), but were found to be unidentifiable past Archosauria indet. by Nesbitt et al. (2007). Heckert et al. (2000) stated it "appears dinosaurian" but also that it "documented the presence of herrerasaurs", suggesting confusion in the precision of identification. They agreed it was generically indeterminate however.
NMMNH P17325 was considered a possible theropod by Hunt (1994) and Hunt et al. (1998), and was referred to Chindesaurus by Long and Murry (1995).
References- Parrish and Carpenter, 1986. A new vertebrate fauna from the Dockum Formation (Late Triassic) of eastern New Mexico. In: Padian, K. (ed.). The Beginning of the Age of Dinosaurs. Cambridge Univ. Press, NewYork. Pp.151-160.
Murry and Long, 1989. Geology and paleontology of the Chinle Formation, Petrified Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils of the southwestern Upper Triassic. in Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. 29-64.
Chatterjee, 1993. Shuvosaurus, a new theropod. National Geographic Research and Exploration. 9(3), 274-285.
Hunt, 1994. Vertebrate paleontology and biostratigraphy of the Bull Canyon Formation (Chinle Group: Norian), east-central New Mexico with revisions of the families Metoposauridae (Amphibia: Temnospondyli) and Parasuchidae (Reptilia: Archosauria). Unpublished PhD Dissertation. Albuquerque, Univerrsity of New Mexico. 403 pp.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Carpenter, 1997. A Giant Coelophysoid (Ceratosauria) Theropod from the Upper Triassic of New Mexico, USA. Neues Jahrbuch fuer Geologie und Palaeontologie, Abhandlungen 205(2): 189-208.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.
Heckert, Lucas and Sullivan, 2000. Triassic dinosaurs in New Mexico. in Lucas and Heckert (eds). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 17-26.
Lehane, 2005. Anatomy and relationships of Shuvosaurus, a basal theropod from the Triassic of Texas. Masters thesis, Texas Tech University. 92 pp.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.
Martz, Mueller, Nesbitt, Stocker, Parker, Atanassov, Fraser, Weinbaum and Lehane, 2013. A taxonomic and biostratigraphic re-evaluation of the Post Quarry vertebrate assemblage from the Cooper Canyon Formation (Dockum Group, Upper Triassic) of southern Garza County, western Texas. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103, 1-26.

"Cryptoraptor" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
"C. lockleyi" Hunt, 1994 vide Nesbitt, Irmis and Parker, 2007
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US
Material
- (NMMNH P17375) (small) vertebral centra, pubis, proximal femora
Comments- Originally named by Hunt (1994) in his unpublished thesis, it was identified as a theropod there, by Hunt et al. (1998) and by Hunt (2001). Nesbitt et al. (2007) later found the femora are too fragmentary for identification, the vertebrae are identical to Shuvosaurus, and that the pubis resembles Shuvosaurus more than dinosaurs. Specifically the acetabulum is closed as in non-dinosaurian taxa and the symphysis extends far proximally as in shuvosaurids. They assigned it to Archosauria indet., but it seems possible this is a shuvosaurid or even a specimen of Shuvosaurus itself.
Although the name "Cryptoraptor lockleyi" was originally used in thesis, and thus not available for use in this website, it was later published by Nesbitt et al. (2007).
References- Hunt, 1994. Vertebrate paleontology and biostratigraphy of the Bull Canyon Formation (Chinle Group: Norian), east-central New Mexico with revisions of the families Metoposauridae (Amphibia: Temnospondyli) and Parasuchidae (Reptilia: Archosauria). Unpublished PhD Dissertation. Albuquerque, Univerrsity of New Mexico. 403 pp.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.
Hunt, 2001. The vertebrate fauna, biostratigraphy and biochronology of the type Revueltian land-vertebrate faunachron, Bull Canyon Formation (Upper Triassic), east-central New Mexico. New Mexico Geological Society Guidebook. 52, 123-151.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.

Lukousaurus

"Plateosaurus" ornatus

Sinosaurus? "shawanensis" Young vide Anonymous, 1979
Early Jurassic
Lufeng Formation, Yunnan, China

Comments- This only appears in a faunal list, and may be a misspelling of Sinosaurus triassicus.
Reference- Anonymous, 1979. Stratigraphy of China, Jurassic System, Summary. Chinese Academy of Geological Sciences. May 1979.

Tanystrosuchus Kuhn, 1963
T. posthumus (Huene, 1908) Kuhn, 1963
= Tanystropheus posthumus Huene, 1908
= Thecodontosaurus posthumus (Huene, 1908) Fraas, 1913
= Halticosaurus posthumus (Huene, 1908) Huene, 1932
= Coelophysis posthumus (Huene, 1908) Olshevsky, 1991
Norian, Late Triassic
Middle Stubensandstein, Germany

Holotype- (SMNS 4385) distal caudal vertebra (39 mm)
Diagnosis- (after Huene, 1908) no notch between distal caudal prezygapophysis and centrum.
Other diagnoses- Huene (1908) also distinguished posthumus from Tanystropheus conspicuus and T. antiquus based on its supposedly enlarged and elongated postzygapophyses, while the prezygapophyses were said to be rudimentary. Yet he had the vertebra oriented backwards, as it is actually the prezygapophyses which are elongate (65% of central length). Comparably elongate prezygapophyses are known in other taxa such as Effigia, herrerasaurids and many neotheropods. Additionally, the neural spine was said to be more reduced than other Tanystropheus species, and the ventral surface has a median groove instead of being flat and/or keeled. The former is typical of theropod distal caudals while the latter is also found in Effigia and most avepods.
Huene (1932) states a unique character is the lack of elongation, but this varies continuously in most archosaur caudal series.
Comments- Collected in the 1860's, this was first described and figured by Meyer (1865) as the caudal vertebra of an unknown reptile. Huene (1908) named it Tanystropheus (consistantly misspelled Tanystrophaeus) posthumus, as the elongate cervicals of Tanystropheus were thought to be theropod caudals at the time. SMNS 4385 was noted to be similar to Tanystropheus cervicals in being elongate, lacking transverse processes and having a reduced neural spine. These characters led Huene to refer Tanystropheus to Coeluridae, though they are now recognized as typical features for theropod caudal vertebrae. Today, coelurids and other related basal coelurosaurs are known to be restricted to the Jurassic and Cretaceous and have shorter prezygapophyses than Tanystrosuchus. Both Meyer and Huene had the vertebra backwards, interpreting its elongate prezygapophyses as postzygapophyses. Fraas (1913) thought much of the Pfaffenhofen quarry material was probably referrable to a small thecodontosaur, which he provisionally used the combination Thecodontosaurus (misspelled Thekodontosaurus) posthumus for. However, posthumus is from the Heslach quarry, not Pfaffenhofen. Fraas did not provide evidence for his reassignment, and Tanystrosuchus differs supposed Thecodontosaurus distal caudals (e.g. YPM 56736) in having a ventral groove, smaller more medially placed postzygapophyses, and larger, longer prezygapophyses. By 1932, Huene had oriented the vertebra correctly and removed posthumus from Tanystropheus, since new remains of the latter had shown it was a nondinosaurian reptile whose elongated vertebrae were cervicals. He instead called it "Tanystropheus" (gen. indet.) posthumus and assigned it to Coelurosauria, although in one table it is listed as Halticosaurus posthumus. This may have been mistakenly retained from an earlier version of the paper, since he does say posthumus could belong to Halticosaurus or Dolichosuchus. Steel (1970) also suggested posthumus might be referrable to Halticosaurus, while Olshevsky (1991) listed it as possibly being Halticosaurus or Liliensternus (which was placed in Halticosaurus until 1984). While Halticosaurus and Dolichosuchus are known from the same formation (but different quarries), they do not preserve distal caudal vertebrae so cannot be compared. Liliensternus loses its transverse processes on caudal 22 and its neural spines at about caudal 30. This fits with Tanystrosuchus' holotype being most similar in proportions to caudal 32 of those illustrated for Liliensternus. It differs from Liliensternus in having longer prezygapophyses (65% of central length compared to 17-22%), less dorsally projected zygapophyses, and lacking a notch between the prezygapophysis and centrum. These seem to be true in all illustrated caudals similar in position to Tanystrosuchus' (20, 24, 32, 38), so probably indicate it is not synonymous. Kuhn (1963) created the new genus Tanystrosuchus for posthumus, as he also decided it was not referrable to Tanystropheus. Wild (1973) also rejected the synonymy of posthumus with Tanystropheus conspicuus. Tanystrosuchus differs from Tanystropheus in having longer prezygapophyses and lacks Tanystropheus' tall bladelike neural spine. In 1965, Kuhn stated Tanystrosuchus might be protorosaurid instead of dinosaurian. Yet Protorosaurus distal caudals differ in having short prezygapophyses, longer postzygapophyses, knob-like transverse processes and a tall bifurcated neural spine. Norman (1990) noted the long prezygapophyses were suggestive of theropod relationship, but considered it a nomen dubium. Olshevsky (1991) placed Tanystrosuchus in Halticosauridae and listed the combination Coelophysis posthumus as a prior synonym, though I have yet to locate this in an earlier published work. Glut (1997) merely listed the genus as a nondinosaurian reptile. Most recently, Rauhut and Hungerbuhler (2000) briefly redescribed and illustrated the material, listing it as Tanystrophaeus posthumus. Their conclusions match Norman's- that it is an indeterminate theropod based on the prezygapophyseal length.
Tanystrosuchus is similar to coelophysoids in general form- elongate amphicoelous centrum (3.25 times longer than tall) with median ventral groove, neural spine reduced to a slight ridge over the postzygapophyses, transverse process reduced to a longitudinal ridge, elongate prezygopophysis and short postzygapophysis. However, the prezygapophysis is actually longer than in coelophysids, 65% of central length compared to 35% in Coelophysis rhodesiensis, 25% in Coelophysis bauri, 22% in Liliensternus and 37% in Dilophosaurus. In this respect Tanystrosuchus is more similar to herrerasaurids and neotheropods (e.g. Elaphrosaurus). It differs from herrerasaurids in having the ventral groove and from known neotheropods in being Triassic in age, though if coelophysoids are monophyletic we would expect Norian neotheropods. Another perhaps more plausible identification is as a shuvosaurid, which are common in the Late Triassic. Effigia has distal caudals which exhibit the same characters noted above for coelophysoids (including the ventral groove), except its prezygapophyses are longer (at least 58% of central length). The preserved caudals have neural spines, albeit low ones, but these are all proximal to the thirtieth caudal and it is probable that more distal ones lacked neural spines as in theropods and Tanystrosuchus. It differs from all examined theropods and Effigia in lacking a notch between the prezygapophysis and centrum. While the anatomy favors a shuvosaurid or neotheropod identification, stratigraphy favors the former only to the extent that Triassic shuvosaurids are certainly known while Triassic neotheropods are unknown but possible depending on theropod topology. Also a factor is that variation along the tail is poorly described for most taxa discussed here, so while the published evidence suggests coelophysoid prezygapophyses never exceed half of centrum length and herrerasaurids lack ventral grooves, this is not as well established as it could be. For now, I recommend Tanystrosuchus posthumus be referred to Archosauria incertae sedis.
References- Meyer, 1865. Reptilien aus dem Stubensandstein des oberen Keupers (Dritte Folge). Palaeontographica. 14, 99-124.
Huene, 1908. Die Dinosaurier der europäischen Triasformation mit Berücksichtiging der aussereuropäischen Vorkommnisse [The dinosaurs of the European Triassic Formation, with consideration of non-European occurrences]. Geologische und Paläontologische Abhandlungen Supplement-Band. 1, 419 pp.
Fraas, 1913. Die neuesten Dinosaurierfunde in der schwäbischen Trias [The newest dinosaur finds in the Swabian Trias]. Naturwissenschaften. 1(45), 1097-1100.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Kuhn, 1963. Sauria (Supplementum I). In Fossilium Catalogus I. Animalia. 104. 87 pp.
Kuhn, 1965. Saurischia (Supplementum 1). In Fossilium Catalogus 1. Animalia. 109, 94 pp.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie. Gustav Fischer Verlag, Stuttgart. 1-87.
Wild, 1973. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus (Bassani) (Neue Ergebnisse). Schweizerische Palaontologische Abhandlungen. 95, 1-162.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press: Berkeley. 280-305.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope, 1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076 pp.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia. 15, 75-88.

Trialestes Bonaparte, 1982
= Triassolestes Reig, 1963 (preoccupied Tillyard, 1918)
T. romeri (Reig, 1963) Bonaparte, 1982
= Triassolestes romeri Reig, 1963
Late Carnian-Early Norian, Late Triassic
Cancha de Bochas Member of the Ischigualasto Formation, San Juan, Argentina

Holotype- (PVL 2561) partial skull, mandibles, four cervical vertebrae, cervical centrum, twelve caudal vertebrae, scapula, humerus, radius, ulnae, radiale, ulnare
Paratype- (PVL 2559) cervicals (lost), sacrals (lost), pubis (lost), astragalus (lost), partial metatarsal I, partial metatarsal II, metatarsal III, partial metatarsal IV, phalanges
Referred- (PVL 3889) vertebrae, forelimb without carpus, pelvis, hindlimb (Bonaparte, 1978)
Comments- Originally described as a coelophysid preserved with Proterochampsa forelimb elements (Reig, 1963), it was later thought to be crocodylomorph based on the elongate radiale and ulnare found in the holotype. PVL 3889 however, has several characters seen in dinosaurs but not crocodylomorphs- laterally excavated vertebral centra; perforated acetabulum; well-developed supracetabular crest; distinct inturned femoral head; mesotarsal ankle; functionally tridactyl pes. Clark et al. (2000) find most of the characters uniting PVL 2561 and PVL 3889 are symplesiomorphic, but that they both have very elongate forearms (1.12 times humeral length). This means either PVL 2561 is crocodylomorph and PVL 3889 is dinosaurian, but they both developed elongate forearms in parallel; or that Trialestes is dinosaurian (based on parsimony) and has a crocodylomorph-like carpus. PVL 2559 was referred to Herrerasauridae indet. by Novas (1989) and Dinosauria indet. by Novas (1993).
Reference- Reig, 1963. La presencia de dinosaurios saurisquios en los "Estrados de Ischigualasto" (Mesotriasico superior) de las provincias de San Juan y La Rioja (Republica Argentina). Ameghiniana 3, 3-20.
Bonaparte, 1982. Classification of the Thecodontia. Geobios Mem. Spec. 6, 99-112.
Novas, 1989. The tibia and tarsus in Herrerasauridae (Dinosauria, incertae sedis) and the origin and evolution of the dinosaurian tarsus. Journal of Paleontology 63 p. 677-690.
Novas, 1993. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina. Journal of Vertebrate Paleontology. 13 p. 400-423.
Clark, Sues and Berman, 2000. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology. 20(4), 683-704.

Pseudosuchia Zittel, 1887
Definition- (Crocodylus niloticus <- Passer domesticus) (Nesbitt, 2011; modified from Gauthier and Padian, 1985)
= Crocodylotarsi Benton and Clark, 1988
Definition- (Crocodylus niloticus <- Vultur gryphus) (modified from Kischlat, 2000)

unnamed pseudosuchian (Chatterjee, 1987)
Carnian, Late Triassic
Maleri Formation, India
Material
- (ISI R 306) (juvenile) partial premaxilla, maxilla, partial dentary, teeth (to 5 mm), dorsal vertebrae?
Comments- This material is part of the Alwalkeria holotype as described by Chatterjee (1987), but Rauhut and Remes (2005) found it belonged to a pseudosuchian, perhaps an ornithosuchid. This agrees with Rauhut's (2003) comments that the long mandibular symphysis is present in members of Crurotarsi. Some of the dorsal vertebrae may belong to this taxon also.
References- Chatterjee, 1987. A new theropod dinosaur from India with remarks on the Gondwana-Laurasia connection in the Late Triassic. Geophysical Monograph. 41, 183-189.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Remes and Rauhut, 2005. The oldest Indian dinosaur Alwalkeria maleriensis Chatterjee revised: a chimera including remains of a basal saurischian. in Kellner, Henriques and Rodrigues (eds). II Congresso Latino-Americano de Paleontologia de Vertebrados, Boletim de Resumos.Museu Nacional, Rio de Janeiro. 218.

Dromaeosuchia Juul, 1994
Definition- (Ornithosuchus longidens + Crocodylus niloticus) (modified from Kischlat, 2000; modified from Juul, 1994)

Ornithosuchia Huene, 1908
Definition- (Ornithosuchus longidens <- Crocodylus niloticus, Vultur gryphus) (modified from Kischlat, 2000)
Other definitions- (Vultur gryphus <- Caiman crocodilus) (modified from Gauthier and Padian, 1985)

Ornithosuchidae Huene, 1908
Definition- (Ornithosuchus longidens + Riojasuchus tenuisceps + Venaticosuchus rusconii) (modified from Sereno, 1991)
Other definitions- (Ornithosuchus longidens <- Rutiodon carolinensis, Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, Crocodylus niloticus, Passer domesticus) (Nesbitt, 2011)

Ornithosuchus

Riojasuchus

Venaticosuchus

Suchia Krebs, 1974
Definition- (Aetosaurus ferratus + Poposaurus gracilis + Rauisuchus tiradentes + Gracilisuchus stipanicicorum + Crocodylus niloticus) (modified from Sereno, 1991)
Other definitions- (Aetosaurus ferratus + Rauisuchus tiradentes + Prestosuchus chiniquensis + Crocodylus niloticus) (Nesbitt, 2011)
= Rauisuchiformes Parrish, 1993
Definition- (Aetosaurus ferratus + Rauisuchus tiradentes) (modified from Parrish, 1993)

Spondylosoma

Gracilisuchus

Aetosauria Marsh, 1884
Definition- (Aetosaurus ferratus <- Belodon plieningeri, Crocodylus niloticus) (modified from Kischlat, 2000)
Other definitions- (Aetosaurus ferratus, Desmatosuchus haplocerus <- Leptosuchus crosbiensis, Postosuchus kirkpatricki, Prestosuchus chiniquensis, Poposaurus gracilis, Sphenosuchus acutus, Alligator mississippiensis, Gracilisuchus stipanicicorum, Revueltosaurus callenderi) (modified from Parker, 2007)
(Aetosaurus ferratus + Desmatosuchus haplocerus <- Rutiodon carolinensis, Postosuchus kirkpatricki, Prestosuchus chiniquensis, Poposaurus gracilis, Crocodylus niloticus, Gracilisuchus stipanicicorum, Revueltosaurus callenderi) (Nesbitt, 2011)

Stegonolepididae Lydekker, 1887
Other definitions- (Aetosaurus ferratus + Desmatosuchus spurensis) (modified from Heckert and Lucas, 2000)
(Aetosaurus ferratus + Stegonolepis robertsoni <- Mystriosuchus planirostris, Ornithosuchus longidens, Crocoylus niloticus) (modified from Benton, 2004)

Ticinosuchidae Sennikov, 1995

Paracrocodylomorpha Parrish, 1993
Definition- (Poposaurus gracilis + Crocodylus niloticus) (Nesbitt, 2011; modified from Parrish, 1993)

Poposauroidea Nopcsa, 1928
Definition- (Poposaurus gracilis <- Postosuchus kirkpatricki, Crocodylus niloticus, Ornithosuchus longidens, Aetosaurus ferratus) (Nesbitt, 2011)
= Poposauria Olshevsky, 1991
= Ctenosauriscidae Kuhn, 1964

Arctosaurus Adams, 1875
A. osborni Adams, 1875
Late Triassic
Heiberg Formation, Nunavut, Canada
Holotype- (NMI 62 1971) incomplete third cervical vertebra (33 mm)
Diagnosis- (suggested) differs from Ctenosauriscus in low cervical neural spine; differs from Lotosaurus in offset centrum faces.
Comments- This was discovered in 1853 and though originally identified as a Teleosaurus vertebra, was described by Adams (1875) as a new reptilian genus more similar to lizards than crocodiles. Non-dinosaurian identifications include Huene's (1902) suggestion it is a testudine. This was based on comparison to his new genus Chelyzoon, which he believed to be a turtle but has since been synonymized with the archosauromorph Tanystrophaeus. Thus a relationship with turtles can be discounted. Baird (in Russell, 1984) suggested it was a trilophosaurid. This needs to be seriously considered, as trilophosaurs have epipophyses and spinopostzygapophyseal laminae. However, they differ in having platycoelous to procoelous centra (in Spinosuchus and Trilophosaurus) which lack a posterior centrodiapophyseal lamina and have larger epipophyses. Furthermore, Arctosaurus has closely situated postzygapophyses which lack the broad horizontal transpostzygapophyseal laminae characteristic of Trilophosaurus and Spinosuchus. The rhynchosaur Mesosuchus also has epipophyses, but similarly differs in lacking posterior centrodiapophyseal laminae as well as having prominent posterodorsally placed diapophyses.
Arctosaurus a dinosaur? Lydekker (1890) placed it in Theropoda fam. indet., stating it was probably related to Anchisaurus (which he viewed as a theropod). Indeed, authors such as Lydekker (1889) and Marsh (1895) placed it in Anchisauridae. A relationship to basal sauropodomorphs remained the consensus until Galton and Cluver (1976) placed it in Theropoda instead due to the short and high proportions and supposed pleurocoel. Additional authors who have placed Arctosaurus in Theropoda include Steel (1970), Welles (1984; as a nomen dubium) and Weishampel (1990). The elongate centrum with offset articular surfaces is like dinosauriforms, and the presence of epipophyses is like dinosaurs. Within Dinosauria, the lack of a pleurocoel excludes it from Avepoda, the relatively short centrum excludes it from Sauropodomorpha (even compared to Panphagia and Pantydraco), and the shortness of the epipophyses exclude it from Theropoda. It is unlike basal ornithischians like Eocursor and Heterodontosaurus in the anteroventral placement of the diapophysis and thus probably parapophysis, and in the well developed posterior centrodiapophyseal lamina which marks a lateral fossa (pleurocoel of Galton and Cluver).
Arctosaurus an archosauriform? The specimen was most recently reviewed by Nesbitt et al. (2007), who placed it as Archosauriformes indet.. Ignoring dinosaurs, epipophyses are only known in suchians (e.g. Revueltosaurus, Xilousuchus, Lotosaurus, Rauisuchus), while the elongate centrum with offset sides resembles those of some poposauroids and crocodylomorphs. Thus Poposauroidea is the most likely candidate. It is more similar to derived poposauroids than basal taxa (Yarasuchus, Qianosuchus) in the well developed posterior centrodiapophyseal lamina. Within poposauroids, Arctosaurus is more similar to Lotosaurus, Ctenosauriscus and Poposaurus than Arizonasaurus, Sillosuchus, Effigia and Shuvosaurus in having a shorter centrum. As in Arizonasaurus but unlike Sillosuchus, Effigia and Shuvosaurus, the centrum is transversely compressed. It is further similar to Arizonasaurus and Lotosaurus, but unlike Poposaurus, Sillosuchus, Effigia and Shuvosaurus in lacking a pleurocoel. Like Lotosaurus and apparently Ctenosauriscus, but unlike Arizonasaurus, Poposaurus, Effigia and Shuvosaurus, Arctosaurus has postaxial epipophyses. Thus it seems most similar to Lotosaurus and Ctenosauriscus. As it differs from both taxa, it is not indeterminate, contra Welles and Nesbitt et al..
References- Adams, 1875. On a fossil saurian vertebra, Arctosaurus osborni, from the Arctic regions. Proceedings of the Royal Ireland Academy. 2, 177-179.
Lydekker, 1889. Notes on new and other dinosaurian remains. Geological Magazine, decade 3. 6(8), 352-356.
Lydekker, 1890. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part IV. Containing the orders Anomodontia, Ecaudata, Caudata and Labyrinthodontia; and supplement. British Museum of Natural History, London. 295 pp.
Marsh, 1895. On the affinities and classification of the dinosaurian reptiles. American Journal of Science. 50(300), 483-498.
Huene, 1902. Übersicht über die Reptilien der Trias [Review of the Reptilia of the Triassic]. Geologische und Paläontologische Abhandlungen (Neue Serie). Gustav Fischer Verlag, Jena. 6, 1-84.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 1-87.
Galton and Cluver, 1976. Anchisaurus capensis (Broom) and a revision of the Anchisauridae. Annals of the South American Museum. 69, 121-159.
Russell, 1984. A check list of the families and genera of North American dinosaurs. Syllogeus Series No. 53. National Museums of Canada.
Welles, 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda), osteology and comparisons. Palaeontographica. Beiträge zur Naturgeschichte der Vorzeit. Abteilung A: Paläozoologie, Stratigraphie. 185, 85-180.
Weishampel, 1990. Dinosaurian distribution. in Weishampel, Dodson and Osmolska (eds.). The Dinosauria. University of California Press. 63-139.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.

Hypselorhachis

Arizonasaurus

Poposauridae Nopcsa, 1928
Poposaurus

Shuvosauridae Chatterjee, 1993
Definition- (Shuvosaurus inexpectatus + Sillosuchus longicervix) (Nesbitt, 2011)
= Shuvosaurinae Chatterjee, 1993
= Chatterjeeidae Long and Murry, 1995

Shuvosaurus Chatterjee, 1993
= "Chatterjeea" Murry and Long, 1989
= Chatterjeea Long and Murry, 1995
S. inexpectatus Chatterjee, 1993
= "Chatterjeea elegans" Murry and Long, 1989
= Chatterjeea elegans Long and Murry, 1995
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, Texas, US

Holotype- (TTUP 9280) incomplete skull (~189 mm), partial mandible
Paratypes- (TTUP 9281) maxillary fragment, frontal, squamosal (may belong to 9282), quadrate, anterior dentaries, dorsal vertebra
(TTUP 9282) premaxillae, frontals, quadrate, braincase, atlantal intercentrum, atlantal neural arch
Referred- ?(DMNH 9887) several cervical vertebral fragments, vertebral fragments (Long and Murry, 1995)
?(DMNH 9888) partial posterior cervical vertebra, two fragmentary mid dorsal vertebrae, partial mid caudal vertebra (Long and Murry, 1995)
?(DMNH 9901) four fragmentary dorsal centra (Long and Murry, 1995)
?(DMNH 9905) two distal caudal vertebrae, incomplete distal caudal vertebra (Long and Murry, 1995)
?(DMNH 9907) partial dorsal centrum (Long and Murry, 1995)
?(DMNH 9924) caudal centrum fragment (Long and Murry, 1995)
?(DMNH 9925) three partial cervical centra (Long and Murry, 1995)
?(DMNH 9926) partial cervical centrum (Long and Murry, 1995)
(TTUP 9001; holotype of Chatterjeea elegans; paratype of Postosuchus kirkpatricki) three cervical vertebrae, partial dorsal vertebra, twenty presacral centra, four fused incomplete sacral centra, partial proximal caudal vertebra, two proximal caudal centra, two incomplete distal caudal vertebrae, scapulae (one incomplete; one partial), coracoids, humerus (152 mm), incomplete ilium, partial pubis (200 mm), incomplete ischium (150 mm), femur (231 mm), tibia (198 mm), fibula (201 mm), astragalus (33 mm wide), calcaneum, metatarsal I (66 mm), phalanx I-1, pedal ungual I, metatarsal II (83 mm), phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (85 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (61 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V (41 mm) (Chatterjee, 1985)
(TTUP 9003-9011; paratypes of Postosuchus kirkpatricki) nine incomplete postcranial skeletons (Chatterjee, 1985)
(TTUP 9605) (skull ~257 mm) maxilla (Lehane, 2005)
(TTUP 10969) quadrate (Lehane, 2005)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US

?(NMMNH P4160) partial cervical centrum (Long and Murry, 1995)
?(NMMNH P4380) partial dorsal centrum (Long and Murry, 1995)
?(NMMNH P4440) partial dorsal centrum (Long and Murry, 1995)
?(NMMNH P4601) ischial fragment (Long and Murry, 1995)
?(NMMNH P4687) proximal femur (Long and Murry, 1995)
?(NMMNH P4693) astragalus (Long and Murry, 1995)
?(NMMNH P4695) femur (Long and Murry, 1995)
?(NMMNH P4856) partial dorsal centrum, central fragments (Long and Murry, 1995)
?(NMMNH P4859) ischia (Long and Murry, 1995)
?(NMMNH P4891) partial cervical vertebra (Long and Murry, 1995)
?(NMMNH P4893) cervical vertebral fragments (Long and Murry, 1995)
?(NMMNH P4913) eight fragmentary cervical centra (Long and Murry, 1995)
?(NMMNH P4920) dorsal centrum (Long and Murry, 1995)
?(NMMNH P4927) (several individuals) five dorsal centra, sacral centrum (Long and Murry, 1995)
?(NMMNH P4948) partial centrum (Long and Murry, 1995)
?(NMMNH P16643) partial dorsal centrum (Long and Murry, 1995)
?(NMMNH P16654) partial cervical centrum (Long and Murry, 1995)
?(NMMNH P17134) dorsal centrum (Long and Murry, 1995)
?(NMMNH P17247) cervical vertebra, vertebral fragments (Long and Murry, 1995)
?(NMMNH P17262) cervical centrum, several incomplete presacral centrum (Long and Murry, 1995)
?(NMMNH P17469) six cervical centra (Long and Murry, 1995)
?(NMMNH P17470) nine dorsal central fragments (Long and Murry, 1995)
?(NMMNH P17271) two dorsal central fragments (Long and Murry, 1995)
?(NMMNH P17281) dorsal central fragment (Long and Murry, 1995)
(UCM 47221; in part; holotype of Gojirasaurus quayi) (immature) four mid or posterior dorsal centra (66, 68, 74 mm), partial mid dorsal neural arch (Parrish and Carpenter, 1986)
?...(UCM 52080) partial ilium, pubic fragment (Parrish and Carpenter, 1986)
?...(UCM 52081) premaxilla (Parrish and Carpenter, 1986)
?(UMMP 7438) three fused sacral centra (Long and Murry, 1995)
Other diagnoses- Chatterjee (1993) listed several characters of the taxon. The concave medial quadrate cotyle and medial dentary shelf are shared with Effigia. A deep and pointed premaxilla, and posteriorly placed external naris are shared with at least Effigia. The premaxilla is not hooked. The external naris is not necessarily smaller than ornithomimosaurs, as the anterior nasal is unpreserved (Lehane, 2005). The maxilla is not especially short, as the posterior portion is merely missing. Similarly, the nasal cannot be shown to be shorter than the frontal due to these unpreserved portions. Rauhut and Lehane have agreed no maxillary fenestra is present, so the nasal cannot participate.
Comments- Several elements have changed identifications and numbers between Chatterjee's (1993) original description and Lehane's (2005) thesis. The dorsal vertebra in TTUP 9281 was originally listed under 9280. Similarly, the scapula originally part of TTUP 9281 was separated as TTUP 11045. Originally, a left atlantal neural arch was listed under TTUP 9281, which may have been reidentified as the maxillary fragment, frontal, squamosal or quadrate. A left squamosal was listed under 9282 by Chatterjee, figured as such by Lehane, but listed under 2981 by Lehane in his text and table 1. An additional frontal was identified in TTUP 9282.
Shuvosaurus' holotype skull was discovered in 1984, and originally described as an ornithomimosaur by Chatterjee (1993). Rauhut (1997) rejected this based on the lack of an expanded cultriform process, long elongate basipterygoid processes, small endocranial cavity, lack of (pro)maxillary fenestrae, and ventral ectopterygoid groove (instead of an expanded fossa). This is unsurprising given the lack of any known Triassic tetanurines, let alone coelurosaurs. He proposed it was a non-tetanurine, though still a theropod. Rauhut (2000; published in 2003) later found Shuvosaurus to fall out in Coelophysoidea based on the elongate basispenoid and posteriorly forked premaxilla, shared with Coelophysis rhodesiensis. Lehane (2005) also found Shuvosaurus to be a coelophysoid sister to rhodesiensis, while Tykoski (2005) recovered it as the basalmost theropod. Lehman and Chatterjee (2005) agreed Shuvosaurus was likely not ornithomimosaurian, but did state "additional cranial and postcranial material suggests that Shuvosaurus is more derived than ceratosaurs."
Long and Murry (1995) on the other hand proposed Shuvosaurus was synonymous with their new pseudosuchian Chatterjeea, with both holotypes discovered in the same quarry and even potentially belonging to the same individual. They referred a premaxilla originally described by Parrish and Carpenter (1986) as Reptilia indet. to Shuvosaurus, and noted a Chatterjeea ilium was found with it (described by Parrish and Carpenter as Poposaurus). Nesbitt et al. (2007) referred the dorsal vertebrae of the Gojirasaurus holotype from this same quarry to Shuvosaurus. While the pubis and tibia are coelophysoid, some of the other elements of the specimen may belong to Shuvosaurus as well. Rauhut (1997) and Lehane both raised objections to the pseudosuchian identification, noting it possesses dinosaurian, saurischian and theropod characters. The issue was decided by the description of Effigia (Nesbitt and Norell, 2006), which has a Shuvosaurus-like skull on a pseudosuchian body. Both taxa do share numerous theropod synapomorphies, but these are seen as convergences. Note both the non-shuvosaurid scapula and avepod tibia which Long and Murry stated raised questions about their proposed synonymy have since been removed from the genus (see below).
Chatterjeea's holotype was originally identified as a juvenile Postosuchus (Chatterjee, 1985), along with nine other specimens. None have been described in detail, and the numerous additional fragments referred to this taxon by Long and Murry may not belong. Specimens from other formations referred to Chatterjeea by Long and Murry may belong to other shuvosaurids so are not listed here.
Not Shuvosaurus- Chatterjee (1993) originally included a proximal scapula in the paratype TTUP 9281, but this was separated as TTUP 11045 by Lehane (2005). Chatterjee also stated as correspondence at the end of his description "we have just found a diagnostic leg bone (tibia) from the Post quarry which confirms the presence of an ostrich dinosaur", though this was not technically referred to Shuvosaurus. The tibia was later assigned the number TTUP 11044, as revealed in Lehane's thesis. Lehane noted both elements are too small to belong to Shuvosaurus individuals known from that quarry. The scapula is here placed as Archosauria indet. pending further study, while the tibia has been reidentified as a probable coelophysoid (Nesbitt and Chatterjee, 2008; Martz et al., 2013).
Lehane listed several other postcranial specimens referred to Shuvosaurus. TTUP 10071 is an incomplete ilium identified as Coelophysis by Lehman and Chatterjee (2005) and agreed to be coelophysoid-grade by later authors (Nesbitt and Chatterjee, 2008; Martz et al., 2013). TTUP 10072 is a fragmentary skeleton (containing dorsal vertebrae and pelvic fragments which Lehane doesn't mention) which was also identified as Coelophysis by Lehman and Chjatterjee, but reidentified as a poorly supported sister to Avepoda by Nesbitt and Chatterjee. Based on my own analysis, it may be a sauropodomorph instead so is here placed as Saurischia incertae sedis. Finally, TTUP 10082 is a partial pelvis yet again referred to Coelophysis by Lehman and Chatterjee, but which was reidentified as herrerasaurid by later authors (Nesbitt and Chatterjee, 2008; Martz et al., 2013) and may belong to Chindesaurus.
References- Chatterjee, 1985. Postosuchus, a new thecodontian reptile from the Triassic of Texas and the origin of tyrannosaurs. Philosophical Transactions of the Royal Society of London, Series B. 309(1139), 395-460.
Parrish and Carpenter, 1986. A new vertebrate fauna from the Dockum Formation (Late Triassic) of eastern New Mexico. In Padian (ed.). The Beginning of the Age of Dinosaurs. Cambridge University Press, NewYork. 151-160.
Murry and Long, 1989. Geology and paleontology of the Chinle Formation, Petrified Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils of the southwestern Upper Triassic. In Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. 29-64.
Chatterjee, 1993. Shuvosaurus, a new theropod. National Geographic Research and Exploration. 9(3), 274-285.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. Bulletin of the New Mexico Museum of Natural History and Science. 4, 1-254.
Rauhut, 1997. Zur Schädelanatomie von Shuvosaurus inexpectatus (Dinosauria, Theropoda). In Sachs, Rauhut and Weigert (eds.). Erstes Treffen der deutschsprachigen Paläoherpetologen, Extended Abstracts. Terra Nostra. 9, 17-21.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis, University of Bristol. 440 pp.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 96 pp.
Lehane, 2005. Anatomy and relationships of Shuvosaurus, a basal theropod from the Triassic of Texas. Masters thesis, Texas Tech University. 92 pp.
Lehman and Chatterjee, 2005. Depositional setting and vertebrate biostratigraphy of the Triassic Dockum Group of Texas. Journal of Earth System Science. 114(3), 325-351.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. Journal of Vertebrate Paleontology. 25(3), 96A.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD thesis, University of Texas at Austin. 553 pp.
Nesbitt and Norell, 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B. 273, 1045-1048.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.
Nesbitt and Chatterjee, 2008. Late Triassic dinosauriforms from the Post Quarry and surrounding areas, west Texas, U.S.A. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen. 249(2), 143-156.
Martz, Mueller, Nesbitt, Stocker, Parker, Atanassov, Fraser, Weinbaum and Lehane, 2013. A taxonomic and biostratigraphic re-evaluation of the Post Quarry vertebrate assemblage from the Cooper Canyon Formation (Dockum Group, Upper Triassic) of southern Garza County, western Texas. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103, 1-26.

unnamed shuvosaurid (Cope, 1887)
Norian, Late Triassic
Petrified Forest Member of the Chinle Formation, New Mexico, US

Material- (AMNH 2725; syntype of Coelurus bauri) distal femur (Cope, 1887)
Comments- This specimen is a syntype of Coelophysis bauri and was collected with the lectotypes of C. bauri, C. longicollis, C. willistoni and the holotype of Longosaurus (Cope, 1887, 1887; Padian, 1986). Most recently, it was viewed as generically indeterminate (along with the types of the above species), and Coelophysis was assigned a neotype from a different locality (Hunt and Lucas, 1991; ICZN, 1996). Sullivan and Lucas (1999) erected a new species of coelophysid from the Coelophysis lectotype locality, Eucoelophysis baldwini, and felt AMNH 2725 was probably referrable to Eucoelophysis, but could not definitively assign it due to its undiagnostic nature. Nesbitt et al. (2005, 2007) determined Eucoelophysis was not a theropod, however, and that AMNH 2725 is not either. While Eucoelophysis is a basal dinosauriform, AMNH 2725 is a shuvosaurid based on the small distal sulcus and absent fourth trochanter (Nesbitt et al., 2007). Additional material originally assigned to Coelophysis by Cope may also turn out to belong to shuvosaurids, although some is coelophysoid (e.g. AMNH 2705, 2706, 2708, 2722).
References- Cope, 1887. The dinosaurian genus Coelurus. Amer. Naturalist xxi pp. 367-369.
Cope, 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proc. Amer. Philos. Soc. xxiv pp. 209-228, pls. i, ii.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic dinosaur from New Mexico (USA). Paläontol. Z. 65 p. 191-198.
International Commision on Zoological Nomenclature, 1996. Opinion 1842. Coelurus bauri Cope, 1887 (currently Coelophysis bauri; Reptilia, Saurischia): lectotype replaced by a neotype. Bulletin of Zoological Nomenclature. 53, 142-144.
Sullivan and Lucas, 1999. Eucoelophysis baldwini, a new theropod dinosaur from the Upper Triassic of New Mexico, and the status of the original types of Coelophysis. Journal of Vertebrate Paleontology 19(1): 81-90.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

Paracrocodyliformes Weinbaum and Hungerbuhler, 2007
Definition- (Batrachotomus wildi, Postosuchus kirkpatricki, Saurosuchus galilei, Crocodylus niloticus <- Poposaurus gracilis) (Weinbaum and Hungerbuhler, 2007)
= Loricata Merrem, 1820 preoccupied Vicq d'Azyr, 1792 and Schumacher, 1817
Definition- (Crocodylus niloticus <- Poposaurus gracilis, Ornithosuchus longidens, Aetosaurus ferratus) (Nesbitt, 2011)
Comments- Though Nesbitt (2011) ressurected Merrem's (1820) name Loricata for this clade, Merrem used the term only for living crocodilians (he did not comment on fossils). Vicq d'Azyr (1792) used Loricata earlier for armadillos, Schumacher (1817) used it for chitons, and Ehrenberg (1830) later used it for a polyphyletic group of rotifers.
References- Vicq d'Azyr, 1792. Systeme anatomique des quadrupedes. Encyclopedic methodique. Volume 2. Paris.
Schumacher, 1817. Essai d'un Nouveau Systeme des Habitations des Vers Testaces. Copenhagen. 287 pp.
Merrem, 1820. Versuch eines Systems der Amphibien. Marburg. 191 pp.
Ehrenberg, 1830. Sur l'organisation des Infusoires. Isis. 2, 168. 8, 758.

Prestosuchidae Romer, 1966
= Prestosuchia Parrish, 1994

Apatosuchus
References- Knoll, 2010. What is "Halticosaurus" orbitoangulatus? Journal of Vertebrate Paleontology. Program and Abstracts 2010, 116A.

unnamed paracrocodyliform (Ellenberger and Ellenberger, 1954)
Hettangian-Sinemurian, Early Jurassic
Upper Elliot Formation, South Africa
Material
- (SAM 383) maxillary fragment, teeth
Comments- Ellenberger and Ellenberger (1954) mentioned a carnosaur maxilla, which Smith believes is SAM 383, as reported by Nesbitt (2009). The latter described it and identified the material as a paracrocodylomorph similar to Fasolasuchus.
References- Ellenberger and Ellenberger, 1954. Le gisement de Dinosauriens de Maphutseng (Basutoland, Afrique du Sud). Compte Rendus de la Société Géologique de France. 241, 99-101.
Nesbitt, 2009. The early evolution of archosaurs: Relationships and the origin of major clades. PhD thesis, Columbia University. 665 pp.

Teratosauroidea Cope, 1871
= Rauisuchoidea Huene, 1942
= Teratosauria Colbert, 1970
= Rauisuchia Bonaparte, 1975

Teratosauridae Cope, 1871
= Rauisuchidae Huene, 1942
Definition- (Rauisuchus tiradentes <- Aetosaurus ferratus, Prestosuchus chiniquensis, Poposaurus gracilis, Crocodylus niloticus) (Nesbitt, 2011)

Teratosaurus

Crocodylomorpha Walker, 1968
Definition- (Crocodylus niloticus <- Postosuchus kirkpatricki) (modified from Kischlat, 2000)
Other definitions- (Crocodylus niloticus <- Poposaurus gracilis, Gracilisuchus stipanicicorum, Prestosuchus chiniquensis, Aetosaurus ferratus) (Fiorelli and Calvo, 2007)
(Crocodylus niloticus <- Rauisuchus tiradentes, Poposaurus gracilis, Gracilisuchus stipanicicorum, Prestosuchus chiniquensis, Aetosaurus ferratus) (Nesbitt, 2011)
= Crocodylomorphi Hay, 1930
= Pedeticosauria Walker, 1968

Clarensea

Apatosuchus orbitoangulatus

Sphenosuchia Bonaparte, 1971
Definition- (Sphenosuchus acutus, Terrestrisuchus gracilis <- Crocodylus niloticus) (Sereno, in prep.)
Other definitions- (Terrestrisuchus gracilis <- Crocodylus niloticus) (Fiorelli and Calvo, 2007)

Hallopoda Marsh, 1881
Definition- (Hallopus victor <- Crocodylus niloticus) (modified from Kischlat, 2000)

Hallopus

Macelognathus

Crocodyliformes Hay, 1930
Definition- (Crocodylus niloticus + Protosuchus richardsoni) (Sereno et al., 2001; modified from Kischlat, 2000)

Kemkemia Cau and Maganuco, 2009
K. auditorei Cau and Maganuco, 2009
Cenomanian, Late Cretaceous
Kem Kem Beds, Morocco
Holotype
- (MSNM V6408) distal caudal vertebra (60.48 mm)
Diagnosis- (modified after Cau and Maganuco, 2009) robust distal caudal neural spine (mediolateral width at apex at least 30% of anterior central width); mediolaterally concave dorsal surface of distal caudal neural spine.
Other diagnoses- Lio et al. (2012) found most of the diagnostic characters proposed by Cau and Maganuco were typical of crocodyliforms- inflated neural canal on distal caudal vertebra, broader than width of centrum at mid-length; strongly reduced finger-like distal caudal prezygapophyses lacking articular facets and failing to reach level of anterior end of centrum; shallow fossa on posterior half of lateral surface of distal caudal neural spine bounded distally by postspinal lamina.
Comments- Cau and Maganuco (2009) believed this specimen resembled ceratosaurs such as Ilokelesia, Ligabueino and especially the Lameta abelisaur vertebra GSI K27/705. However, Lio et al. (2012) reidentified it as a crocodyliform vertebra. Due to the poor description of most crocodyliforms, it is not possible to determine its relationships further than it being outside Eusuchia, and dissimilar to Baurusuchus, Simosuchus, Notosuchus and Uruguaysuchus. It may be synonymous with another named taxon from the area, like Araripesuchus, Aegisuchus, Aegyptosuchus, Hamadasuchus, Kaprosuchus, Laganosuchus or Stomatosuchus. Chiarenza and Cau (2016) mentioned mid caudals of Spinosaurus/Sigilmassasaurus share Kemkemia's neural spine anatomy, reduced zygapophyses, non-grooved ventral surface and tall neural spine on a vertebra lacking transverse processes. They thus proposed Kemkenia may be a spinosaurine distal caudal vertebra. I feel this is doubtful as spinosaurine mid caudals lack the expanded neural canal, neural spine fossa or pre/postspinal laminae, and each of these characters are expected to be present proximally and end distally in theropod tails. Thus the broad and dorsally concave neural spine anatomy would be more parsimoniously convergent in Kemkemia and spinosaurines.
References- Cau and Maganuco, 2009. A new theropod dinosaur, represented by a single unusual caudal vertebra, from the Kem Kem Beds (Cretaceous) of Morocco. Atti della Società Italiana di Scienze Naturali e del Museo di Storia Naturale di Milano. 150(II), 239-257.
Lio, Agnolin, Cau and Maganuco, 2012. Crocodyliform affinities for Kemkemia auditorei Cau and Maganuco, 2009, from the Late Cretaceous of Morocco. Atti della Società Italiana di Scienze Naturali e del Museo di Storia Naturale di Milano. 153(I), 119-126.
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.

"Maroccanoraptor" Singer, 2015 online
"M. elbegiensis" Singer, 2015 online
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco

Material- (JP Cr683) coracoid (proximodistal 150 mm)
Comments- This was briefly described online by Singer (2015), on the website of a theme park chain based on a specimen collected and owned by the chain. He assigns this unofficial taxon to Unenlagiinae based on resemblence to the coracoid of Buitreraptor. However, Cau (online, 2015) convincingly showed a greater resemblence to crocodyliforms such as Simosuchus. While several taxa are known from the Kem Kem beds (Aegisuchus, Arraripesuchus? rattoides, Elosuchus, Hamadasuchus, Laganosuchus, etc.), none have preserved coracoids, and very few coracoid characters have been used in crocodyliform analyses so far.
References- Cau, 2015 online. http://theropoda.blogspot.it/2015/12/maroccanoraptor.html
Singer, 2015 online. JuraPark na tropie nowych dinozaurow z Maroka. http://jurapark.pl/jurapark-na-tropie-nowych-dinozaurow-z-maroka/

Protosuchia Mook, 1934
Definition- (Protosuchus richardsoni <- Crocodylus niloticus) (Sereno et al., 2001)

Protosuchidae Brown, 1934
Definition- (Protosuchus richardsoni + Hemiprotosuchus leali) (Fiorelli and Calvo, 2007)
= Protosuchoidea Brown, 1934 sensu Deraniyagala, 1939
= Protosuchinae Brown, 1934 sensu Busbey and Gow, 1982

Mesoeucrocodylia Whetstone and Whybrow, 1983
Definition- (Crocodylus niloticus <- Protosuchus richardsoni) (Sereno et al., 2001)
= Emydosauria Blainville, 1822

Lisboasaurus

Lusitanosuchus

Razanandrongobe

Sinocoelurus Young, 1942
S. fragilis Young, 1942
Tithonian, Late Jurassic
Guangyuan (Kuangyuan) Formation, Sichuan, China

Syntypes- (IVPP V232-234) incomplete tooth, two partial teeth, tooth (? x 9 x 4.5; ? x 8.5 x 6; ? x 7.5 x 5, ? x 8 x 5.2 mm)
Comments- Dong et al. (1983) stated these could derive from the underlying Aalenian-Bajocian Qianfuyan Formation instead.
These teeth were discovered in 1941 and described by Young (1942) as a new taxon of coelurosaur (sensu Huene) based on "the general features of the teeth". Though none are complete, the crowns would have been ~40 mm long. Dong et al. (1983) incorrectly included IVPP V235 as a syntype, but this is a Szechuanosaurus specimen, showing two Sinocoelurus teeth must share one specimen number. The incomplete tooth is IVPP V232, but the identity of the two partial teeth which are illustrated, which number includes two teeth, and which specimen each set of FABL and BW measurements is from are unknown. The teeth are highly elongate, strongly compressed (BW/FABL = 50-70%) and slightly curved lingually and distally. There are both mesial and distal carinae, with the distal carina being stronger. The carinae lack serrations, which Young used to justify creation of a new genus. Wu et al. (2009) further described them as striated. Within Coelurosauria, Steel (1970) referred Sinocoelurus to Coeluridae, which was followed by most traditional authors. This was due to Coeluridae being a wastebasket taxon for small theropods, in addition to Sinocoelurus' name no doubt. More recently, Sinocoelurus has usually been referred to as Theropoda indet. (e.g. Norman, 1990), though Holtz et al. (2004) placed it in Tetanurae indet. without comment.
Wu et al. examined the material and considered it to most probably belong to a plesiosaur. However, plesiosaur teeth are circular in section (except for the tip in some taxa), including the contemporaneous Yuzhoupliosaurus, and their carinae are perpendicular to the plane of curvature.
Rozhdestvensky (1977) referred Sinocoelurus to Crocodylia without comment. A placement in Crocodyliformes seems plausible, as taxa such as Kaprosuchus, Shantungosuchus, Torvoneustes and Pehuenchesaurus have compressed, recurved teeth without serrations. Further study is necessary however.
References- Young, 1942. Fossil vertebrates from Kuangyuan, N. Szechuan, China. Bulletin of the Geological Society of China. 22(34), 293-309.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 87 pp.
Rozhdestvensky, 1977. The study of dinosaurs in Asia. Journal of the Palaeontological Society of India. 20, 102-119.
Dong, Zhou and Zhang, 1983. [Dinosaurs from the Jurassic of Sichuan]. Palaeontologica Sinica, New Series C. 162(23), 1-136.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press: Berkeley. 280-305.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. The Dinosauria (second edition). University of California Press. 71-110.
Wu, Currie, Dong, Pan and Wang, 2009. A new theropod dinosaur from the Middle Jurassic of Lufeng, Yunnan, China. Acta Geologica Sinica. 83(1), 9-24.

unnamed crocodyliform (Sauvage, 1876)
Early Cretaceous
Blacourt, Oise, France

Material- distal femur
Comments- This was initially described as Megalosaurus by Sauvage (1876), then referred to Erectopus superbus by Huene (1926). Chure (2000) determined it was crocodyliform.
References- Sauvage, 1876. Notes sur les reptiles fossiles. Bulletin of the Geological Society of France (ser. 3). 4, 435-442.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista Museo de La Plata. 29, 35-167.
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.

undescribed crocodyliform (Carrano et al., 2002)
Middle Maastrichtian, Late Cretaceous
Anembalemba Member of Maevarano Formation, Madagascar

Material- (FMNH PR 2204) proximal caudal vertebra
Comments- This was identified as Masiakasaurus (Carrano et al., 2002) before being reidentified as a crocodyliform (Carrano et al., 2011).
References- Carrano, Sampson and Forster, 2002. The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology. 22(3), 510-534.
Carrano, Loewen and Sertich, 2011. New materials of Masiakasaurus knopfleri Sampson, Carrano, and Forster, 2001, and implications for the morphology of the Noasauridae (Theropoda: Ceratosauria). Smithsonian Contributions to Paleobiology. 95, 53 pp.

Gobiosuchidae Osmolska, 1972
Definition- (Gobiosuchus kielanae + Zaraasuchus shepardi) (Fiorelli and Calvo, 2007)

Hsisosuchidae

Metasuchia Benton and Clark, 1988
Definition- (Notosuchus terrestris + Crocodylus niloticus) (Sereno et al., 2001)

Trematochampsidae Buffetaut, 1974

Notosuchia Gasparini, 1971
Definition- (Notosuchus terrestris <-Crocodylus niloticus) (Sereno et al., 2001)
= Sebecia Larsson and Sues, 2007
Definition- (Sebecus icaeorhinus <- Crocodylus niloticus) (Larsson and Sues, 2007)

Notosuchiformes Carvalho et al., 2004
Definition- (Notosuchus terrestris + Uruguaysuchus aznarezi) (modified from Carvalho et al., 2004)
= Notosuchia sensu Carvalho et al., 2004
Definition- (Peirosaurus torminni + Notosuchus terrestris + Libycosuchus brevirostris + Baurusuchus pachecoi) (modified)
= Notosuchimorpha Calvalho et al., 2004
Definition- (Notosuchus terrestris + Stolokrosuchus lapparenti) (modified from Carvalho et al., 2004)
= Peirosauromorpha Carvalho et al., 2004
Definition- (Peirosaurus torminni + Candidodon itapecurvense)

Brasileosaurus Huene, 1931
B. pachecoi Huene, 1931
Turonian, Late Cretaceous
Adamantina Formation of the Bauru Group, Brazil
Holotype
- incomplete humerus, incomplete femur
Comments- Assigned to Coelurosauria by Huene (1931) and to Coeluridae by Romer (1966). Huene (1933) later recognized it was a crocodile, possibly of the same genus as Uruguaysuchus aznarezi. Simpson (1937) assigned it to Notosuchidae.
References- Huene, 1931. Verschiedene mesozoische Wirbeltierreste aus Südamerika [Different Mesozoic vertebrate remains from South America]. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Abteilung A. 66, 181-198.
Huene, 1933. Ein Versuch zur Stammesgeschichte der Krokodile. Centralblatt für Mineralogie, Geologie und Paläontologie, Abteilung B. 11, 577-585.
Simpson. 1937. An ancient eusuchian crocodile from Patagonia. American Museum Novitates. 965, 1-20.
Kuhn, 1939. Saurischia. In Fossilium Catalogus I. Animalia. 87. 124 pp.
Price, 1950. Os crocodilídeos da fauna da formação Bauru, do Cretáceo terrestre do Brasil meridional. Anais da Academia Brasileira de Ciências. 22, 473-490.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press, Chicago. 1-468.
Bertini, 1994. Comments on the fossil amniotes from the Adamantina and Marília formations, continental Upper Cretaceous of the Paraná Basin, Southeastern Brazil (Part 1: Introduction, Testudines, Lacertilia, Crocodylomorpha). Boletim do 3° Simpósio sobre o Cretáceo do Brasil, Rio Claro, 1994, UNESP - Campus de Rio Claro/SP. 97-100.

Peirosauridae sensu Larsson and Sues, 2007
Definition- (Peirosaurus torminni, Uberabasuchus terrificus <- Sebecus icaeorhinus)

Peirosauriformes Carvalho et al., 2004
Definition- (Araripesuchus gomesii + Peirosaurus torminni) (modified from Carvalho et al., 2002)

Uruguaysuchidae Rusconi, 1933
Other definitions- (Uruguaysuchus) (Carvalho et al., 2004)
= Uruguaysuchinae Rusconi, 1933

Mahajangasuchini sensu Carvalho et al., 2004
Definition- (Uberabasuchus terrificus + Mahajangasuchus insignis) (modified from Carvalho et al., 2004)

Cunampaia

Hamadasuchus
H? sp.
(Canudo, Salgado, Barco, Bolatti and Ruiz-Omeñaca, 2004)
Late Cenomanian-Early Turonian, Late Cretaceous
Cerro Lisandro Formation, Rio Negro, Argentina
Material
- (Endemas-PV 6) lateral tooth (~21x9x7.6 mm)
Comments- Canudo et al. (2004) and Salgado et al. (2004) reffered this tooth to Spinosauridae and cf. Spinosauridae respectively. Salgado et al. (2009) indicated this differs from spinosaurids in having large serrations which are apically oriented, and Hasegawa et al. (2010) referred it to Hamadasuchus.
References- Canudo, Salgado, Barco, Bolatti and Ruiz-Omeñaca, 2004. Dientes de dinosaurios terópodos y saurópodos de la Formación Cerro Lisandro (Cenomaniense superior-Turoniense inferior, Cretácio superior) en Río Negro (Argentina). Geo-Temas. 6, 31-34.
Salgado, Canudo, Garrido, Ruiz-Omeñaca, Garcýa, de la Fuente, Barco and Bollati, 2009. Upper Cretaceous vertebrates from El Anfiteatro area, Rio Negro, Patagonia. Cretaceous Research. 30, 767-784.
Hasegawa, Tanaka, Takakuwa and Koike, 2010. Fine sculptures on a tooth of Spinosaurus (Dinosauria, Theropoda) from Morocco. Bulletin of Gunma Museum of Natural History. 14, 11-20.

"Mahajangasuchidae" Sereno and Larsson, 2009
Definition- (Mahajangasuchus insignis <- Baurusuchus pachecoi, Pholidosaurus schaumbergensis, Goniopholis crassidens, Crocodylus niloticus, Peirosaurus torminni, Simosuchus clarki, Araripesuchus gomesii, Notosuchus terrestris) (Sereno and Larsson, 2009)

Peirosauridae Gasparini, 1982
Definition- (Peirosaurus torminni + Lomasuchus palpebrosus) (modified from Carvalho et al., 2004)
Other definitions- (Peirosaurus torminni <- Notosuchus terrestris, Araripesuchus gomesii, Simosuchus clarki, Baurusuchus pachecoi, Crocodylus niloticus) (Fiorelli and Calvo, 2007)

Sebecidae sensu Larsson and Sues, 2007
Definition- (Sebecus icaeorhinus <- Peirosaurus torminni, Uberabasuchus terrificus)

Candidodontidae Carvalho et al., 2004
Definition- (Candidodon itapecurvense + Mariliasuchus amarili) (modified from Carvalho et al., 2004)

Ziphosuchia Ortega et al., 2000
Definition- (Notosuchus terrestris + Libycosuchus brevirostris + Baurusuchus pachecoi) (modified from Cavalho et al., 2004)
= Sebecidae sensu Carvalho et al., 2004
Definition- (Sebecus icaeorhinus + Libycosuchus brevirostris) (modified)

Libycosuchidae Stromer, 1914
= Libycosuchinae Stromer, 1914

Chimaerasuchidae sensu Carvalho et al., 2004
Definition- (Chimaerasuchus pardoxus + Simosuchus clarki) (modified from Carvalho et al., 2004)

Gondwanasuchia Carvalho et al., 2004
Definition- (Chimaerasuchus paradoxus + Notosuchus terrestris) (modified from Carvalho et al., 2004)

Notosuchidae Dollo, 1924
Other definitions- (Notosuchus) (Calvalho et al., 2004)
(Notosuchus terrestris <- Comahuesuchus brachybuccalis, Araripesuchus gomesii, Simosuchus clarki, Baurusuchus pachecoi, Crocodylus niloticus) (Fiorelli and Calvo, 2007)
= Notosuchinae Dollo, 1924 vide Nopcsa, 1928

Sphagesuchidae sensu Fiorelli and Calvo, 2007
Definition - (Sphagesaurus huenei <- Notosuchus terrestris, Comahuesuchus brachybuccalis, Araripesuchus gomesii, Simosuchus clarki, Baurusuchus pachecoi, Sebecus icaeorhinus, Crocodylus niloticus)

Sphagesauridae Kuhn, 1968
Definition- (Sphagesaurus huenei, Adamantinasuchus navae <- Chimaerasuchus paradoxus, Notosuchus terrestris, Mariliasuchus amarali, Uruguaysuchus aznarezi, Uruguaysuchus terrai, Comahuesuchus brachybuccalis, Candidodon itapecurvense, Simosuchus clarki, Baurusuchus pachecoi, Sebecus icaeorhinus) (Marinho and Carvalho, 2007)

Doratodon

Sebecosuchia Simpson, 1937
Definition- (Pehuenchesuchus enderi + Baurusuchus pachecoi) (modified from Martinelli and Pais, 2008)

Sebecoidea Simpson, 1937 vide Sereno in prep.
= Baurusuchoidea Price, 1945 vide Carvalho e al., 2004
Definition- (Sebecus icaeorhinus + Baurusuchus pachecoi) (modified from Carvalho et al., 2004)

Bergisuchidae Rossman, Ruhe and Ortega, 2000

Iberosuchidae Antunes, 1975

Sebecidae Simpson, 1937
Other definitions- (Sebecus icaeorhinus + Libycosuchus brevirostris) (modified from Carvalho et al., 2004)
(Sebecus icaeorhinus <- Peirosaurus torminni, Uberabasuchus terrificus) (Larsson and Sues, 2007)

Baurusuchidae Price, 1945
Definition- (Baurusuchus pachecoi <- Sebecus icaeorhinus, Crocodylus niloticus, Simosuchus clarki, Araripesuchus gomesii, Comahuesuchus brachybuccalis, Notosuchus terrestris) (Sereno, in prep.)
Other definitions- (Baurusuchus pachecoi + Stratiotosuchus maxhechti) (modified from Carvalho et al., 2005)

Pabwehshi Wilson, Malkani and Gingerich, 2001
= "Vitakridrinda" Malkani, 2003
= Vitakridrinda Malkani, 2006a
P. pakistanensis Wilson, Malkani and Gingerich, 2001
= "Vitakridrinda sulaimani" Malkani, 2003
= Vitakridrinda sulaimani Malkani, 2006a
Maastrichtian, Late Cretaceous
Vitakri Formation, Pakistan
Holotype-
(GSP-UM 2000) anterior skull, anterior mandibles
Paratype- (GSP-UM 2001) anterior mandibles
Referred- ?(MSM-53-2, MSM-54-2, MSM-55-2; paratypes of Vitakridrinda sulaimani) three fragmentary dorsal or caudal vertebrae (8.7 mm)
?(MSM-56-1; paratype of Vitakridrinda sulaimani) caudal vertebra
?(MSM-57-3; paratype of Vitakridrinda sulaimani) caudal vertebra (7.3 mm)
?(MSM-58-15; paratype of Vitakridrinda sulaimani) presacral vertebra (7.9 mm)
?(MSM-59-19; syntype of Vitakridrinda sulaimani) proximal ?femur
?....(MSM-60-19; syntype of Vitakridrinda sulaimani) proximal ?femur
?(MSM-61-19; syntype of Vitakridrinda sulaimani) posterior ?braincase
?(MSM-62-19; syntype of Vitakridrinda sulaimani) ?tooth fragment (FABL 22 mm)
(MSM-155-19; syntype of Vitakridrinda sulaimani) anterior skull, anterior mandibles
?(MSM-282-15) partial vertebra (Malkani, 2009)
?(MSM-706-19) partial presacral centrum (Malkani, 2009)
?(MSM-765-19) partial presacral centrum (Malkani, 2009)
?(MSM-780-2) partial vertebra (Malkani, 2009)
?(MSM-984-2) femoral fragment (Malkani, 2009)
?(MSM-1027-2) limb bone fragment (Malkani, 2009)
?(MSM-1039-19) femoral fragment (Malkani, 2009)
?(MSM-1040-16) partial vertebra (Malkani, 2009)
?(MSM-1041-2) limb bone fragment (Malkani, 2009)
?(MSM-1042-4) limb bone fragment (Malkani, 2009)
?(MSM-1043-16) limb bone fragment (Malkani, 2009)
?(MSM-1044-16) limb bone fragment (Malkani, 2009)
?(MSM-1048) partial vertebra (Malkani, 2009)
?(MSM-1049-K) proximal femur or nodule (Malkani, 2009)
?(MSM-1050) coprolite (Malkani, 2009)
?(MSM-1051) coprolite (Malkani, 2009)
?(MSM-1052) coprolite (Malkani, 2009)
?(MSM-1053) coprolite (Malkani, 2009)
?(MSM-1054-2) coprolite (Malkani, 2009)
?(MSM-1057-K) femoral fragment or nodule (Malkani, 2009)
?(MSM-1059) long bone fragment (Malkani, 2009)
Diagnosis- (after Wilson et al., 2001) dorsally inset first premaxillary alveolus (unknown in Wargosuchus); prominent premaxillary lip overhanging the external nares (unknown in Cynodontosuchus); diverticulum in the posterodorsal comer of the external naris (unknown in Campinasuchus, Cynodontosuchus and Pissarrachampsa); diastematic fossa positioned just posterior to the second maxillary tooth (unknown in Cynodontosuchus and Wargosuchus); enlarged, caniniform sixth dentary tooth (unknown in Campinasuchus, Cynodontosuchus, Pissarrachampsa, Stratiotosuchus and Wargosuchus); reduced fourth and fifth dentary teeth (unknown in Cynodontosuchus, Stratiotosuchus and Wargosuchus).
Other diagnoses- Malkani (2006a) lists many characters for Vitakridrinda in his diagnosis. Those involving the snout (rostrum; labiolingually compressed tooth) are plesiomorphic for baurusuchids. Though Malkani (2006a, b) originally reported the naris was surrounded by the premaxilla, he later (2009) indicated the nasal participates too. The tooth has a low crown according to Malkani (CHR ~1-1.5), but this falls within the range of Rugops, Majungasaurus and other unnamed abelisaurids (e.g. MGUP MEGA002), in addition to baurusuchids like Baurusuchus and Cynodontosuchus.
For the supposed braincase, Malkani listed "occipital condyle articulated with posterior braincase" which is plesiomorphic for archosaurs, as well as thick and long occipital condyle. Given the lack of any obvious foramen magnum above it, any character based on this object is suspect. Contra Malkani (2009), the braincase does not show more closely spaced 'basipterygoid processes' (basal tubera, if braincase is correctly identified) than Abelisaurus.
Another character listed by Malkani is "amphicoelous vertebrae", but this is based on referred specimens and plesiomorphic for archosaurs.
Finally, the syntype femora are diagnosed as having- greater trochanter; inturned femoral head; and femur hollow with thick walls. The former is plesiomorphic for archosaurs, the second plesiomorphic for dinosaurs, and the last apparently untrue according to Malkani (2009).
Comments- Pabwehshi was described in 2001 based on two snouts, with Wilson et al. (2001) assigning it to Baurusuchidae.
Turner and Calvo (2005) added Pabwehshi to a phylogenetic analysis (127 characters, 32 taxa; founded in Clark's 1994 analysis). They recovered it sister to the only included baurusuchid, Baurusuchus. After ordering several characters (1, 3, 15, 37, 45, 49, 67, 77, 111, 113; mostly based on Turner, 2006), much resolution is lost, but Pabwehshi's placement is identical once the wildcard Iberosuchus is deleted a posteriori. Note Eremosuchus is in the cladogram, but not the data matrix. Enforcing Sues and Larsson's Sebecia (see below) is 8 steps longer, and moving Pabwehshi there (it emerges sister to Bretesuchus) is 9 steps longer. If Hamadasuchus is added, that genus is sister to Araripesuchus and the tree otherwise similar except Libycosuchus is now sister to Notosuchus+Comahuesuchus+Anatosuchus. Now, Sebecia is only 4 steps longer and moving Pabwehshi there 5 steps longer (sister to Bretesuchus again).
Turner (2006) slightly changed the analysis (adding two characters and replacing Libycosuchus with Araripesuchus wegeneri) and found similar results to Turner and Calvo, with Pabwehshi sister to Baurusuchus. The same ordering was modified, and enforcing Sebecia is still 8 steps longer, but moving Pabwehshi there (again sister to Bretesuchus) is 11 steps longer. Hamadasuchus is again sister to Araripesuchus when added, and the tree is otherwise similar. Sebecia is 7 steps longer, and including Pabwehshi in it is 10 steps longer.
Larsson and Sues (2007) entered it into a phylogenetic analysis (158 characters, 33 taxa) and found it sister to a clade of sebecids, Hamadasuchus and peirosaurids instead. They named this clade Sebecia. It takes 4 more steps to place Pabwehshi sister to Baurusuchus, though no other baurusuchids are included. As this is the only analysis to find Pabwehshi outside Baurusuchidae, its topology is tested versus other analyses discussed here.
Nascimento and Zaher (2011) added it to another analysis (262 characters, 63 taxa; founded in Gasparini et al.'s 2005 analysis) which placed it in Baurusuchidae sister to Stratiotosuchus. In this tree, sebecids (here used in a broad sense to also include Bergisuchus and/or Iberosuchus) are sister to baurusuchids, peirosaurids are closer to crocodylians, and Hamadasuchus is not included. 10 more steps are needed to make Larsson and Sues' Sebecia and 14 to include Pabwehshi in it (it falls out sister to Iberosuchus). After adding Hamadasuchus, it takes 8 steps to make that Sebecia and 11 steps to include Pabwehshi.
Montefeltro et al. (2011) examined the phylogeny of Baurusuchidae but excluded Pabwehshi a priori because "it has a sagittal torus on its maxillary palatal shelves, absent in the putative baurusuchids" (referencing Larsson and Sues' paper), but this is only one character. If added to their matrix (66 characters, 10 taxa), Pabwehshi is sister to other baurusuchids. As Larsson and Sues stated "P. pakistanensis shares numerous features with Sebecia in general and H. rebouli in particular", Hamadasuchus was added to the matrix as well. The latter emerges sister to Pabwehshi and other baurusuchids, and enforcing a Pabwehshi-Hamadasuchus clade is two steps longer. Notably, Montefeltro et al.'s analysis is completely cranial and does not include baurusuchid sister taxa (sebecids, then Pehuenchesuchus which can only be coded for two characters) as outgroups.
Pol and Powell (2011) included Pabwehshi in a comparable analysis (285 characters, 89 taxa; founded in Pol and Norell's 2004 analysis) where it again fell out sister to Stratiotosuchus in Baurusuchidae. Sebecids are again sister to baurusuchids, but Hamadasuchus is sister to Peirosauridae (including Mahajungasuchus) plus Stolokrosuchus and neosuchians. 2 more steps are needed to make Larsson and Sues' Sebecia and 6 more to place Pabwehshi there (where it falls out as a basal sebecid). 3 more steps are needed to place this Sebecia sister to Larsson and Sues' version of Neosuchia instead of being in Notosuchia, and 7 more to then include Pabwehshi.
Pol et al. (2012) ran a larger version of that analysis (347 characters, 88 taxa) which had similar results, though Baurusuchidae's monophyly is masked by the uncertain position of Cynodontosuchus. Sebecids are again sister to baurusuchids, but Hamadasuchus (plus Mahajungasuchus) is sister to pierosaurids in basal Notosuchia. 5 more steps are needed to make Larsson and Sues' Sebecia and 9 more to place Pabwehshi there (where it falls out as a basal sebecid). Yet 16 more steps are needed to place this Sebecia sister to Larsson and Sues' version of Neosuchia instead of being in Notosuchia, and 19 more to then include Pabwehshi. To test the importance of including Hamadasuchus, excluding it makes Sebecia 13 steps less likely instead of 5. Forcing a sebecian Pabwehshi without Hamadasuchus is still four steps longer yet (at 17 more steps).
The conclusion of these analyses is that Sebecia is slightly to moderately unparsimonious in most analyses (2-10 more steps), and the one which supported it is smaller than most which rejected it. Importantly, even when Sebecia is enforced, Pabwehshi still clades with baurusuchids with most analyses (including the largest ones) taking 4 extra steps to move it into Sebecia. This suggests there is no sebecian signal in Pabwehshi, as does the lack of a particular position within Sebecia when it is forced to be there (sister to either Bretesuchus, Iberosuchus, or a wildcard basal sebecid). Notably it is always attracted to sebecids though, which are sister to baurusuchids in non-sebecian phylogenies, as opposed to being isolated or close to Hamadasuchus as in Larsson and Sues' paper. While Hamadasuchus is important for making Sebecia itself more likely (by 1-6 extra steps), it only made Pabwehshi more likely to be a sebecian in one analysis, and then only by one step. Most recently, Pol et al. (2014) expanded on the last analysis (412 characters, 109 taxa) to find Pabwehshi as a basal baurusuchid, more derived than Cynodontosuchus but outside Pissarrachampsinae+Baurusuchinae.
Vitakridrinda- Malkani first mentioned Vitakridrinda in his 2003 description of Brohisaurus, referencing it as an abelisaurid described in "Malkani, 2004a". The only publication in the bibliography solely by Malkani is his saurischian biodiversity paper, listed as in progress, but not actually published until 2006. As the 2003 mention of Vitakridrinda lacks a description or definition (ICZN Article 13.1.1; note the reference to Malkani, 2004a doesn't count under 13.1.2 since it was not published yet), was not indicated to be a new taxon (16.1), and did not have a type specimen indicated (16.4), it was a nomen nudum at the time. The first description of Vitakridrinda is generally claimed to be the 2004 "Saurischian Dinosaurs from Late Cretaceous of Pakistan", as referenced in Malkani 2006a and the Paleobiology Database. Yet ICZN Article 9.9 lists "abstracts of articles, papers, posters, texts of lectures, and similar material when issued primarily to participants at meetings, symposia, colloquia or congresses" as not being published work, so the paper doesn't count. In addition, it still violates 16.4 in not indicating a type specimen. Note while Malkani's "Saurischian dinosaurs from the Late Cretaceous Pab Formation of Pakistan" has been listed as being published in 2005 (Malkani, 2006a) or "2004d" and in review (Malkani, 2006b), Malkani confirms (pers. comm. 2012) that it remains unpublished. Two 2006 publications of Malkani's do describe Vitakridrinda sufficiently, with 2006b being published in December, while 2006a was published in April. Thus Malkani's 2006a publication is the actual official description of Vitakridrinda.
The material was probably mostly discovered in 2001, as Malkani discovered 2700 bones and bone fragments from the area at that time. The femora MSM-59-19 and MSM-60-19 were found associated in one mass and seemingly belong to one individual. The braincase was found 100 meters away in the same horizon, while the snout and tooth were found later (though early enough to be mentioned in 2004) and 50 meters from the braincase. Malkani believes they all belong to the same individual based on supposed lack of transportation and similar size, but the area of separation is so large that this lacks merit. While Malkani refers to them all as the holotype, they may more properly be syntypes, and their uncertain derivation from one animal would make selection of a lectotype desirable. Regarding the supposed femora, figure 14a of Malkani 2006a does seem to show a hollow cross section of a long bone, but the perspective doesn't reveal which element or what kind of animal. Figures 5-8 of the femora are too small to say anything from. Based on Malkani's (2006a) description, figure 14b is supposed to be an anterior view of paroccipital processes and basipterygoid processes, which is similar in rough outline to Abelisaurus (assuming the supposed basipterygoid processes are basal tubera instead) except for the decurved and much taller paroccipital processes. The latter features match titanosaur braincases better, though in that case everything beneath the occipital condyle area would be broken off. In either case, it's only the shape that is similar, and since there are no obvious surface features or natural edges, it could just as easily be part of an ilium, vertebra, etc.. Figures 15a and b do have the rough shape of an occipital condyle, but no obvious foramen magnum above it. The possibility it could be a non-fossil rock with a rounded protruding area should not be excluded. Assuming the gray chevron-shape in the rock (figure 16a of 2006a) is the supposed tooth based Malkani's (2006a) description, it's the cross section of a carina. There are many other possibilities for its identification though, assuming it's even a fossil.
Despite labels for MSM-155-19, I don't see any distinction from the matrix or bone contacts in figure 5 of Malkani (2006b). The supposed anterior and ventral premaxillary edge is the right shape for an abelisaurid, and the supposed naris is darker (but so is the supposed nasal), but there are two steps in the lateral surface running longitudinally which make no sense for a maxilla. The photo does not show any unambiguous fossil features. However, Wilson (pers. comm., 2014) correctly observed the posterior view (fig. 2C in Malkani, 2010) is very similar to the snout cross section of Pabwehshi. Pabwehshi is from the same formation and about 50% larger, though the latter measurement will vary based on how far down the snout each cross section is. The nasal diverticulum is listed by Wilson et al. (2001) as an autapomorphy of Pabwehshi, but the internal anatomy of other baurusuchid snouts is poorly known, though Baurusuchus, Stratiotosuchus and Wargosuchus at least lack evidence of it in the external naris. Under Molnar's interpretation, Malkani's dorsal palatal process is the secondary palate, the ventral palatal process is the mandible, the cavity between these is the oral cavity, and the cavity in the ventral palatal process is the Meckelian canal. This leaves almost all of Malkani's supposed lateral surface as matrix which needs to be prepared to expose bone. Thus his supposed bite punctures are merely grooves excavated in the matrix, while his 'combat teeth' are tooth cross sections ventral to the dentary. Malkani's scenario of intraspecific combat causing these features is thus exposed as being fanciful. While most features of Vitakridrinda's snout as described by Malkani are based on misinterpretation, it should be noted Pabwehshi has labiolingually compressed tooth crowns with mesial and distal serrations and a more convex mesial edge as in abelisaurids. Of the other holotype material, the limb bone segments are possibly theropod due to their supposed inturned head, distinct neck, and hollowness, though Malkani (2009) states a section "has fibrous bone network in the hollow", so perhaps they are not actually hollow. The stated lack of an anterior trochanter may support a crocodiliform identity, or may be due to preservation. The supposed braincase matches Baurusuchus as poorly and ambiguously as it does abelisaurids, so remains an unidentified object. The supposed tooth cross section works as well for Pabwehshi as it does for abelisaurids.
The paratype vertebrae are from other localities and do not overlap the type material, so cannot be convincingly referred to the same taxon. Both abelisaurids and notosuchians can have amphicoelous centra, and while Malkani (2009) says the vertebral centra are hollow, he also says they have fibrous bone in the central cavity, so are not pneumatic. In 2006(a) he says they lack pleurocoels, which would also match either abelisaurs' posterior vertebrae or crocodiliform vertebrae. At least MSM-56-1 and 57-3 appear more similar to titanosaurs given their tall postzygapophyses and short, posteriorly placed neural arch. They are all illustrated in small photographs from oblique perspectives, making comparison difficult. The material referred by Malkani (2009) is far too fragmentary to be justified, including cross sections of long bones, fragments of long bone walls, material that Malkani himself lists as possibly non-fossil and coprolites. All non-snout material would most accurately be Archosauria indet. or something even less specific, but is retained here for ease of comparison pending further study.
The pes MSM-303-2 was referred to Vitakridrinda by Malkani (2009), but later made the holotype of Vitakrisaurus by Malkani (2010b).
References- Wilson, Malkani and Gingerich, 2001. New crocodyliform (Reptilia, Mesoeucrocodylia) from the Upper Cretaceous Pab Formation of Vitakri, Balochistan (Pakistan). Contributions from the Museum of Paleontology. The University of Michigan. 30(12), 321-336.
Malkani, 2003. First Jurassic dinosaur fossils found from Kirthar range, Khuzdar District, Balochistan, Pakistan. Geological Bulletin University of Peshawar. 36, 73-83.
Malkani, 2004. Saurischian dinosaurs from Late Cretaceous of Pakistan. in Hussain and Akbar (eds.). Abstract volume of Fifth Pakistan Geological Congress, Islamabad, Pakistan. 71-73.
Turner and Calvo, 2005. A new sebecosuchian crocodyliform from the Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 25, 87-98.
Malkani, 2006a. Biodiversity of saurischian dinosaurs from the Latest Cretaceous park of Pakistan. Journal of Applied and Emerging Sciences. 1(3), 108-140.
Malkani, 2006b. First rostrum of carnivorous Vitakridrinda (abelisaurid theropod dinosaur) found from the latest Cretaceous Dinosaur Beds (Vitakri) Member of Pab Formation, Alam Kali Kakor locality of Vitakri area, Barkhan District, Balochistan, Pakistan. Sindh University Research Journal (Science Series). 38(2), 5-24.
Turner, 2006. Osteology and phylogeny of a new species of Araripesuchus (Crocodyliformes: Mesoeucrocodylia) from the Late Cretaceous of Madagascar. Historical Biology. 18, 255-369.
Larsson and Sues, 2007. Cranial osteology and phylogenetic relationships of Hamadasuchus rebouli (Crocodyliformes: Mesoeucrocodylia) from the Cretaceous of Morocco. Zoological Journal of the Linnean Society. 149, 533-567.
Malkani, 2009. New Balochisaurus (Balochisauridae, Titanosauria, Sauropoda) and Vitakridrinda (Theropoda) remains from Pakistan. Sindh University Research Journal (Science Series). 41(2), 65-92.
Malkani, 2010a. Vitakridrinda (Vitakrisauridae, Theropoda) from the Latest Cretaceous of Pakistan. Journal of Earth Science. 21(Special Issue 3), 204-212.
Malkani, 2010b. Stratigraphy and mineral potential of Sulaiman (Middle Indus) basin, Pakistan. Sindh University Research Journal (Science Series). 42(2), 39-66.
Malkani, 2011. Vitakridrinda and Vitakrisaurus of Vitakrisauridae Theropoda from Pakistan. Proceedings of the 6th Symposium of IGCP 507 on Paleoclimates of the Cretaceous in Asia and their global correlation. Beijing, China. 59-66.
Montefeltro, Larsson and Langer, 2011. A new baurusuchid (Crocodyliformes, Mesoeucrocodylia) from the Late Cretaceous of Brazil and the phylogeny of Baurusuchidae. PLoS ONE. 6, e21916.
Nascimento and Zaher, 2011. The skull of the Upper Cretaceous baurusuchid crocodile Baurusuchus albertoi Nascimento & Zaher 2010, and its phylogenetic affinities. Zoological Journal of the Linnean Society. 163(S1), S116-S131.
Pol and Powell, 2011. A new sebecid mesoeucrocodylian from the Rio Loro Formation (Palaeocene) of north-western Argentina. Zoological Journal of the Linnean Society. 163, S7-S36.
Pol, Leardi, Lecuone and Krause, 2012. Postcranial anatomy of Sebecus icaeorhinus (Crocodyliformes, Sebecidae) from the Eocene of Patagonia. Journal of Vertebrate Paleontology. 32, 328-354.
Pol, Nascimento, Carvalho, Riccomini, Pires-Domingues and Zaher, 2014. A new notosuchian from the Late Cretaceous of Brazil and the phylogeny of advanced notosuchians. PLoS ONE. 9(4), e93105.
Malkani, unpublished. Saurischian dinosaurs from the Late Cretaceous Pab Formation of Pakistan. Geological Survey of Pakistan, Information Release. 823, 1-117.

Neosuchia Benton and Clark, 1988
Definition- (Crocodylus niloticus <- Notosuchus terrestris) (Sereno et al., 2001)
Other definitions- (Crocodylus niloticus <- Sebecus icaeorhinus) (Larsson and Sues, 2007)

Tethysuchia Buffetaut, 1982
Definition- (Pholidosaurus purbeckensis + Dyrosaurus phosphaticus) (Andrade et al., 2011)

Pholidosauridae sensu Sereno, in prep.
Definition- (Pholidosaurus schaumbergensis <- Alligatorellus beaumonti, Peirosaurus torminni, Araripesuchus gomesii, Notosuchus terrestris, Crocodylus niloticus)

Dyrosaurinae Zittel, 1923
Definition- (Dyrosaurus phosphaticus <- Sarcosuchus imperator, Pholidosaurus schaumbergensis, Goniopholis crassidens) (Sereno, in prep.)
= Dyrosauridae Zittel, 1923

Pholidosauridae Zittel and Eastman, 1902
Other definitions- (Pholidosaurus schaumbergensis <- Alligatorellus beaumonti, Peirosaurus torminni, Araripesuchus gomesii, Notosuchus terrestris, Crocodylus niloticus) (Sereno, in prep.)
= Pholidosaurinae Zittel and Eastman, 1902 sensu Nopcsa, 1928

Thalattosuchia Fraas, 1902
Definition- (Teleosaurus cadomensis, Metriorhynchus geoffroyii <- Pholidosaurus schaumburgensis, Dyrosaurus phosphaticus, Goniopholis crassidens) (Young and Andrade, 2009)
Other definitions- (Geosaurus giganteus <- Pholidosaurus schaumbergensis, Goniopholis crassidens, Crocodylus niloticus) (Sereno, in prep.)
= Teleosaurii Geoffroy, 1831
= Teleosaurina Bonaparte, 1840
= Teleosauri Owen, 1842
= Metriorhynchi Fitzinger, 1843
= Teleosauria Giebel, 1847

Cystosaurus

"Megalosaurus" mersensis

Teleosauroidea Geoffroy, 1831
Definition-
(Teleosaurus cadomensis <- Metriorhynchus geoffroyii) (Young and Andrade, 2009)
= Teleosauridae Cope, 1871
Definition- (Teleosaurus cadomensis <- Metriorhynchus geoffroyi, Crocodylus niloticus) (Sereno, in prep.)
= Teleosaurinae Cope, 1871 sensu Lydekker, 1887

unnamed teleosaur (Lapparent and Zbyszewski, 1957)
Kimmeridgian, Late Jurassic
Casal Labrusque / Areia Branca, Leiria, Portugal
Materia
l- (Geology Laboratory of the Faculty of Sciences of Lisbon coll.) two proximal caudal vertebrae (65 mm), five proximal caudal vertebrae (80 mm), distal caudal vertebra (75 mm)
Comments- These were referred to Megalosaurus insignis by Lapparent and Zbyszewski (1957). Chabli (1986) thought this series of caudals may be teleosaurian.
References- Lapparent and Zbyszewski, 1957. Les dinosauriens du Portugal [The dinosaurs of Portugal]. Mémoires des Services Géologiques du Portugal, nouvelle série. 2, 1-63.
Chabli, 1986. Données nouvelles sur un 'Dinosaurien' Jurassique Moyen du Maroc: Megalosaurus mersensis Lapparent 1955, et sur les Megalosaurides en général [New data on a Middle Jurassic 'dinosaur' from Morocco: Megalosaurus mersensis Lapparent 1955, and on megalosaurids in general]. Les Dinosaures de La Chine à La France, Colloque International de Paléontologie, Toulouse, France, 2-6 Septembre 1985. Muséum d'Histoire Naturelle de Toulouse, Toulouse. 65-72.

Metriorhynchoidea Fitzinger, 1843 sensu Deraniyagala, 1939
Definition- (Metriorhynchus geoffroyi <- Teleosaurus cadomensis) (Young and Andrade, 2009)
= Metriorhynchidae sensu Sereno, in prep.
Definiton- (Metriorhynchus geoffroyi <- Teleosaurus cadomensis, Crocodylus niloticus)

Metriorhynchidae Fitzinger, 1843
Definition- (Metriorhynchus geoffroyi + Geosaurus giganteus) (Young and Andrade, 2009)
Other definitions- (Metriorhynchus geoffroyi <- Teleosaurus cadomensis, Crocodylus niloticus) (Sereno, in prep.)
= Geosauridae Lydekker, 1889

Metriorhynchinae Fitzinger, 1843 sensu Lydekker, 1887
Definition- (Metriorhynchus geoffroyi <- Geosaurus giganteus) (Young and Andrade, 2009)

Geosaurinae Lydekker, 1889 sensu Nopcsa, 1928
Definition- (Geosaurus giganteus <- Metriorhynchus geoffroyi) (Young and Andrade, 2009)

Geosaurus Cuvier, 1824
= Halilimnosaurus Ritgen, 1826
= Brachytaenius Meyer, 1842
G. giganteus (Soemmerring, 1816)
= Lacerta gigantea Soemmerring, 1816
= Lacerta (Geosaurus) gigantea (Soemmerring, 1816) Cuvier, 1824
= Mosasaurus bavaricus Holl, 1829
= Geosaurus soemmerringii Meyer, 1831
= Brachytaenius perennis Meyer, 1842
= Geosaurus giganteus (Soemmerring, 1816) Cuvier, 1824 sensu Quenstedt, 1852
= Mosasaurus giganteus (Soemmerring, 1816) Cope, 1869
G. gracilis (Quenstedt, 1885) new combination
= Dakosaurus maximus var. gracilis Quenstedt, 1852
= Dakosaurus gracilis Quenstedt, 1885
= Megalosaurus gracilis (Quenstedt, 1885) Douville, 1885
= Dakosaurus lapparenti Debelmas and Strannolaubsky, 1957
= Geosaurus lapparenti (Debelmas and Strannolaubsky, 1957) Young and Andrade, 2009

Dakosaurus Quenstedt, 1856
= Plesiosuchus Owen, 1884
= Aggiosaurus Ambayrac, 1913
D. andiniensis Vignaud and Gasparini, 1996
D. lissocephalus Seeley, 1869
D. manselii (Hulke, 1870) Woodward, 1885
= Steneosaurus manselii Hulke, 1870
= Plesiosuchus manselii (Hulke, 1870) Owen, 1884
D. maximus (Plieninger, 1846) Quenstedt, 1856
= Geosaurus maximus Plieninger, 1846
= Liodon primaevum Sauvage, 1871
?= Megalosaurus schnaitheimii Bunzel, 1871
= Dakosaurus primaevus (Sauvage, 1871) Sauvage, 1873
D. nicaeensis (Ambayrac, 1913) Young and Andrade, 2009
= Aggiosaurus nicaeensis Ambayrac, 1913

"Dakosaurus" amazonicus

Stomatosuchidae Stromer, 1925
Definition- (Stomatosuchus inermis <- Crocodylus niloticus, Baurusuchus pachecoi, Peirosaurus torminni, Simosuchus clarki, Araripesuchus gomesii, Notosuchus terrestris) (Sereno and Larsson, 2009)
= Aegyptosuchidae Kuhn, 1936

Susisuchidae Salisbury, Frey, Martill and Buchy, 2003
Other definitions- (Susisuchus anatoceps + Susisuchus jaguaribensis) (Fortier and Schultz, 2009)

Goniopholididae Cope, 1875
Definition- (Goniopholis crassidens <- Crocodylus niloticus, Alligatorellus beaumonti, Pholidosaurus schaumbergensis, Peirosaurus torminni, Araripesuchus gomesii, Notosuchus terrestris) (Sereno, in prep.)
= Goniopholidinae Cope, 1875 sensu Lydekker, 1887
= Goniopholidiformes Hay, 1930

Bernissartiidae Dollo, 1883 emmend. Lydekker, 1888
=Bernissartiinae Dollo, 1883 sensu Nopcsa, 1928

Eusuchia Huxley, 1875
Definition- (Hylaeochampsa vectiana + Crocodylus niloticus) (Fiorelli and Calvo, 2007; modified from Brochu, 1999)
Other definitions- (Hylaeochampsa vectiana + Gavialis gangeticus + Alligator mississippiensis + Crocodylus niloticus) (Brochu, 2003)

Atoposauridae Gervais, 1871
Other definitions- (Atoposaurus jourdani <- Pierosaurus torminni, Araripesuchus gomesii, Notosuchus terrestris, Baurusuchus pachecoi, Crocodylus niloticus) (Sereno, in prep.)

Paralligatoridae Konzhukova, 1954

Hylaeochampsidae Williston, 1925
= Hylaeochampsinae Williston, 1925 sensu Nopcsa, 1928

Crocodylia Gmelin, 1789
Definition- (Alligator mississippiensis + Crocodylus niloticus + Gavialis gangeticus) (Brochu, 2003; modified from Benton and Clark, 1988)
Other definitions- crown(Crocodylus niloticus <- Passer domesticus) (Sereno, 2004)
(Crocodylus niloticus + Gavialis gangeticus) (Sereno, 2001)
= Brevirostres Zittel, 1887
Definition- (Alligator mississippiensis + Crocodylus niloticus) (modified from Brochu, 1999)

Alligatoroidea Cuvier, 1807 sensu Gray, 1844
Definition- (Alligator mississippiensis <- Gavialis gangeticus, Crocodylus niloticus) (Brochu, 2003; modified from Norell et al., 1994)

Gavialoidea Adams, 1854 sensu Case, 1930
Definition- (Gavialis gangeticus <- Alligator mississippiensis, Crocodylus niloticus) (Brochu, 2003; modified from Norell et al., 1994)

Crocodyloidea Covier, 1807 sensu Fitzinger, 1826
Definition- (Crocodylus niloticus <- Gavialis gangeticus, Alligator mississippiensis, Crocodylus niloticus) (Brochu, 2003; modified from Norell et al., 1994)

Avemetatarsalia Benton, 1999
Definition- (Vultur gryphus <- Crocodylus niloticus) (modified from Kischlat, 2000; modified from Benton, 1999)
Other definitions- (Passer domesticus <- Crocodilus niloticus) (Sereno, 2005)
= Panaves Gauthier and de Quieroz, 2001
Definition- (Vultur gryphus <- Crocodylus niloticus)
= Avemetatarsalia sensu Sereno, 2005
Definition- (Passer domesticus <- Crocodilus niloticus)

Avifilopluma Gauthier and de Queiroz, 2001
Definition- (hollow-based, filamentous epidermal appendages homologous with Vultur gryphus) (Gauthier and de Queiroz, 2001)
Comments- Avifilopluma was named a a theropod subclade defined by the presence of feathers (broadly defined). Yet not only has the feathered possible megalosauroid or basal coelurosaur Sciurumimus made this taxon broader (as foreseen by its namers), but the uncertain position of Sciurumimus within Orionides makes its application uncertain. Even worse, the ornithischians Tianyulong and Kulindadromeus show identical structures, so all dinosaurs may be avifiloplumans. Moving stemwards, pterosaurs possess this covering as well, termed pycnofibers. Intermediate taxa are only known to possess scales or naked skin when large, so while rampant convergence is possible, it's also possible large taxa lose feathers. Given all of this, Avifilopluma is a good example of why apomorphy-based definitions should not be used.

Ornithodira Gauthier, 1986
Definition- (Pterodactylus antiquus + Vultur gryphus) (modified from Kischlat, 2000)
Other definitions- (Scleromochlus taylori + Pterodactylus antiquus + Megalosaurus bucklandii) (modified from Sereno, 1991)
(Pterodactylus antiquus + Megalosaurus bucklandii) (modified from Benton, 2004)
(Pterodactylus antiquus + Passer domesticus) (Nesbitt, 2011)
= Ornithotarsi Gauthier, 1986

Avipes Huene, 1932
A. dillstedtianus Huene, 1932
Late Ladinian, Middle Triassic
Lettenkohlensandstein, Germany

Holotype- mid-portion of metatarsus (35 mm as preserved)
References- Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre entwicklung und geschichte. Monographien zur Geologia und Palaeontologie. 1, 1-362.
Rauhut and Hungerbuhler, 2000. A review of European Triassic theropods. Gaia 15, 75-88.

unnamed ornithodiran (Galton, Dyke and Kurochkin, 2009)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania

Material- (MTCO 17032; = MTCO-P 1912) distal ?radius
(MTCO 17637) proximal radius or distal femur
(MTCO 17661; = MTCO-P 9109) proximal radius or distal femur
Comments- Kessler and Jurcsak (1984) referred MTCO 17032 (= MTCO-P 1912) to Archaeopteryx sp. as a distal ulna. Dyke et al. (2011) considered it Archosauria indet., though noted it did resemble a pterosaur distal radius.
Galton et al. (2009) refer to basal hesperornithine elements from the Berriasian of Romania which are similar to Enaliornis. Dyke et al. only noted one element referred to Hesperornithes, a distal femur (MTCO 17637) said to be similar to Enaliornis in size and shape. However, Agnolin and Varricchio (2012) thought the very small and widely separated distal condyles, very wide, opened and pneumatized popliteal groove, and lack of an anterior extensor groove were dissimilar to birds and resembled the proximal radius of Azhdarcho.
MTCO 17661 (= MTCO-P 9109) was illustrated as a distal humerus of Aves indet. by Jurcsack and Kessler (1991), but was recently identified as a bird distal femur by Dyke et al.. Agnolin and Varricchio believed it was more similar to Azhdarcho's radius, for the same reasons as MTCO 17637.
References- Kessler, 1984. Lower Cretaceous birds from Cornet, Roumania. In Rief and Westphal (eds). Third Symposium on Mesozoic Terrestrial Ecosystems, Tubingen. 119-121.
Kessler and Jurcsak, 1984. Fossil bird remains in the bauxite from Cornet (Romania, Bihor County). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 25, 393-401.
Jurcsack and Kessler, 1991. The Lower Cretaceous paleofauna from Cornet, Bihor County, Romania. Nymphaea. 21, 5-32.
Galton, Dyke and Kurochkin, 2009. Re-analysis of Lower Cretaceous fossil birds from the UK reveals an unexpected diversity. Journal of Vertebrate Paleontology. 29(3), 102A.
Dyke, Benton, Posmosanu and Naish, 2011. Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology. 54(1), 79-95.
Agnolin and Varricchio, 2012 . Systematic reinterpretation of Piksi barbarulna Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of Western USA (Montana) as a pterosaur rather than a bird. Geodiversitas. 34(4), 883-894.

Pterosauromorpha Padian, 1997
Definition- (Pterodactylus antiquus <- Megalosaurus bucklandii) (modified from Kischlat, 2000; modified from Padian, 1997)
Other definitions- (Pterodactylus antiquus <- Passer domesticus) (Nesbitt, 2011)

undescribed possible pterosauromorph (Atanassov, 2001)
= Protoavis texensis (in part) Chatterjee, 1991
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, Texas, US

Material- (TTU P 9200; holotype of Protoavis texensis) ischium
(TTU P 9201; paratype of Protoavis texensis) tibia (~39 mm), fibula (~41 mm), four tarsals, metatarsal I (12 mm), metatarsal II (20 mm), metatarsal III (18 mm), metatarsal IV (14 mm)
Late Carnian, Late Triassic
Tecovas Formation of the Dockum Group, Texas, US

(TTU P 9367) tibia (Chatterjee, 1995)
(TTU P 9368) tibia (~58 mm) (Chatterjee, 1995)
(TTU P 9369) fibula (Chatterjee, 1995)
(TTU P 10110) partial maxilla, incomplete dentary, two sacral vertebrae, ilium, ischia, femur, tibia, fibula, astragalus, calcaneum, metatarsus, phalanx V-1
?(TTU P coll.) vertebrae
Comments- This taxon was named and described in a thesis (Atanassov, 2001, 2002) as a basal pterosauromorph. Examination of the figures leads me to believe material originally referred to Protoavis in fact belongs to Atanassov's taxon. The original (Chatterjee, 1991; reidentified as a vomer and pterygoid in 1999) sternum is an ischium, the radius and ulna are the tibia and fibula, and the metacarpus is the metatarsus. Sereno (1997) was the first to suggest Protoavis' supposed metacarpus was actually an archosaurian metatarsus. The new form's maxilla, dentary, ilium and femur seem unrepresented in Protoavis' material, while some characters of the vertebrae are a good match. Unfortunately, only two vertebrae (both sacrals) were associated with the appendicular and cranial elements of this new taxon, so it's possible the latter's vertebrae are from a drepanosaurid or other taxon.
References- Chatterjee, 1991. Cranial Anatomy and relationships of a new Triassic bird from Texas. Philosophical Transactions of the Royal Society of London Series B 332(1265): 277-342.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences. 25, 435-489.
Chatterjee, 1999. Protoavis and the early evolution of birds. Palaeontographica A. 254, 1-100.
Atanassov, 2001. Two new archosauromorphs from the Late Triassic of Texas. Journal of Vertebrate Paleontology. 21(3) 30A.
Atanassov, 2002. Two new archosaur reptiles from the Late Triassic of Texas. Unpublished thesis. Texas Tech University. 352 pp.

Scleromochloidea Huene, 1914 sensu Young, 1964
Scleromochlidae Huene, 1914
Scleromochlinae Huene, 1914 sensu Nopcsa, 1923
Scleromochlus

Pterosauria

"Archaeopteryx" "vicensensis"

Laopteryx

Rhamphorhynchus

Eurolimnornithiformes Bock and Buhler, 1996
Eurolimnornithidae Kessler and Jurcsak, 1986
Comments- Kessler and Jurcsak (1986) proposed the name Eurolimnornithidae for their new genus Eurolimnornis, but continued to place it in the order Limnornithiformes. Yet since Limnornis is a passeriform, using the latter order for the Cretaceous taxon would be confusing. Bock and Buhler (1996) accordingly named the Eurlimnornithiformes.
References- Kessler and Jurcsak, 1986. New contributions to the knowledge of the Lower Cretaceous bird remains from Cornet (Romania). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 28, 289-295.
Bock and Buhler, 1996. Nomenclature of Cretaceous birds from Romania. Cretaceous Research. 17, 509-514.
Eurolimnornis Kessler and Jurcsak, 1986 (non Kessler, 1987)
E. corneti Kessler and Jurcsak, 1986 (non Kessler, 1987)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania
Holotype
- (MTCO 17642; = MTCO-P 7896) distal humerus (12 mm wide)
Comments- This specimen has a complicated taxonomic history, which has been unraveled by Bock and Buhler (1996). Those authors note the relevent papers were published very close together and in a different order than their manuscripts were written, so often reference incorrect dates for each other. In addition, while the holotypes of each valid species were originally thought to be a humerus and femur, Agnolin and Varricchio (2012) have revealed both to be humeri, and both are now catalogued under different numbers than they were when most of the literature was written. Kessler (1984) originally described this specimen and another distal humerus (MTCO 14909; = MTCO-P 1637; described as a distal femur) as "Limnornis corneti", but did not provide a proper description, making it a nomen nudum. Kessler and Jurcsak (1984) soon officially described the species as Limnornis corneti with MTCO 14909 as the holotype, but learned the genus was preoccupied by a furnariid passeriform. Jurcsak and Kessler (1985) thus proposed the replacement name "Palaeolimnornis" for their species. However, "Palaeolimnornis" was merely introduced in a list of taxa, making it a nomen nudum. Kessler and Jurcsak (1986) again noted Limnornis was preoccupied, but this time proposed the name Eurolimnornis corneti for the taxon. Oddly, they did not discuss why they did not stick with "Palaeolimnornis" (though they did state it was a synonym), and unfortunately they designated MTCO 17642 as the holotype. This makes Eurolimnornis corneti a separate species from their original Limnornis corneti based on MTCO 14909. In 1987, Kessler officially proposed Eurolimnornis as a replacement name for Limnornis corneti, though he did not note "Palaeolimnornis" or Kessler and Jurcsak's earlier use of Eurolimnornis. This makes Eurolimnornis corneti of Kessler, 1987 (based on MTCO 14909) a separate taxon (and junior homonym) of Eurolimnornis corneti of Kessler and Jurcsak, 1986 (based on MTCO 17642). MTCO 14909 is now known under the name Palaeocursornis corneti. It seems Kessler and/or Jurcsak had intended to switch the holotype of corneti to make MTCO 17642 Palaeolimnornis corneti, later changed to Eurolimnornis corneti, and make MTCO 14909 Palaeocursornis biharicus, but various steps were not allowed by the ICZN, leading to the current situation.
Kessler and Jurcsak (1986) and Kurochkin (1995) considered this species to be an avian sensu lato (perhaps similar to grebes), but Hope (2002) noted the rounded distal condyles are known in most ornithuromorphs, while the well developed brachial fossa is present in Ichthyornis as well (it is also seen in Longicrusavis, Gansus and several non-ornithuromorphs). Most recently, Agnlin and Varricchio (2012) have reinterpreted it as pterosaurian. They noted it differed from Ornithocheiroidea and Azhdarchoidea, but did not identify it past Pterosauria indet.. Here it is placed in Caelidracones, as only this clade extended into the Cretaceous.
A supposed ulnar shaft fragment (MTCO 17956; = MTCO-P 6966) and supposed distal carpometacarpus (MTCO 17558; = MTCO-P 207) were made paratypes of this species by Kessler and Jurcsak (1986), but Hope finds there is no evidence for this and Dyke et al. (2011) have reidentified them as an archosaur long bone shaft and bird proximal scapula respectively.
References- Kessler, 1984. Lower Cretaceous birds from Cornet, Roumania. In Rief and Westphal (eds). Third Symposium on Mesozoic Terrestrial Ecosystems, Tubingen. 119-121.
Kessler and Jurcsak, 1984. Fossil bird remains in the bauxite from Cornet (Romania, Bihor County). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 25, 393-401.
Jurcsak and Kessler, 1985. La paleofaune de Cornet - implications phylogenetiques et ecologiques. Evolution et Adaptation. 2, 137-147.
Kessler and Jurcsak, 1986. New contributions to the knowledge of the Lower Cretaceous bird remains from Cornet (Romania). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 28, 289-295.
Kessler, 1987. New contributions to the knowledge about Lower and Upper Cretaceous birds from Romania. In Currie and Koster (eds). Fourth Symposium on Terrestrial Ecosystems. Occasional Papers, Tyrrell Museum of Paleontology. 3, 133-135.
Jurcsack and Kessler, 1991. The Lower Cretaceous paleofauna from Cornet, Bihor County, Romania. Nymphaea. 21, 5-32.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of class Aves. Archaeopteryx. 13, 47-66.
Bock and Buhler, 1996. Nomenclature of Cretaceous birds from Romania. Cretaceous Research. 17, 509-514.
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.
Hope, 2002. The Mesozoic radiation of Neornithes. In Chiappe and Witmer (eds). Mesozoic birds: Above the heads of dinosaurs. Berkeley: University of California Press. 339-388.
Dyke, Benton, Posmosanu and Naish, 2011. Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology. 54(1), 79-95.
Agnolin and Varricchio, 2012 . Systematic reinterpretation of Piksi barbarulna Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of Western USA (Montana) as a pterosaur rather than a bird. Geodiversitas. 34(4), 883-894.

Cimoliornis diomedeus

Pterodactylus

Samrukia Naish, Dyke, Cau, Escuillie and Godefroit vide Buffetaut, 2011
= "Samrukia" Naish, Dyke, Cau, Escuillie and Godefroit, online 2011
S. nessovi Naish, Dyke, Cau, Escuillie and Godefroit vide Buffetaut, 2011
= "Samrukia nessovi" Naish, Dyke, Cau, Escuillie and Godefroit, online 2011
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Holotype
- (WDC Kz-001) posterior mandibles
Other diagnoses- (after Naish et al., 2011) large size (mandibular length >275 mm); presence of deep mediodorsal sulcus in the post-dentary region; prominent and raised anterior margin of the mandibular cotyla.
Comment- This specimen was placed in Ornithuromorpha when analyzed in the theropod matrix of Naish et al. (2012), but Buffetaut (2011) subsequently identified it as a pterosaur. Oddly, the latter restudy appeared in print before the actual description.
References- Buffetaut, 2011. Samrukia nessovi, from the Late Cretaceous of Kazakhstan: A large pterosaur, not a giant bird. Annales de Paléontologie. 97(3-4), 133-138.
Naish, Dyke, Cau, Escuillie and Godefroit, 2011. A gigantic bird from the Upper Cretaceous of Central Asia. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 165.
Naish, Dyke, Cau, Escuillie and Godefroit, 2012. A gigantic bird from the Upper Cretaceous of Central Asia. Biology Letters. 8(1), 97-100.

Piksi Varricchio, 2002
P. barbarulna Varricchio, 2002
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US

Holotype- (MOR 1113) distal humerus, proximal radius, proximal ulna, distal ulna
Diagnosis- (after Agnolin and Varricchio, 2012) distal margin of humerus oblique relative to the main axis of the shaft, with trochlea strongly distally extended; trochlea bulbous and subspherical; acute and well defined transverse crest connecting the entepicondyle and the lateral ridge that delimitates the olecranal fossa, in posterior view.
Comments- Originally described as a ornithothoracine of uncertain, but probably basal, relationship. Agnolin and Varricchio (2012) later redescribed it as an ornithocheiroid pterosaur.
References- Varricchio, 2002. A new bird from the Upper Cretaceous Two Medicine Formation of Montana. Canadian Journal of Earth Sciences. 39(1), 19-26.
Agnolin and Varricchio, 2012 . Systematic reinterpretation of Piksi barbarulna Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of Western USA (Montana) as a pterosaur rather than a bird. Geodiversitas. 34(4), 883-894.

Cretornis hlavaci

Palaeornis clifti

Herbstosaurus

Palaeocursornithiformes Kessler and Jurcsak, 1986
= Limnornithiformes Kessler and Jurcsak, 1984
= Cursornithiformes Jurcsak and Kessler, 1985
Palaeocursornithidae Jurcsak and Kessler, 1988
= "Limnornithidae" Kessler, 1984
= Limnornithidae Kessler and Jurcsak, 1984
= "Cursornithidae" Jurcsak and Kessler, 1985
Comments- Kessler (1984) originally proposed the Limnornithidae, but this was without an official description, making it a nomen nudum. Shortly thereafter, Kessler and Jurcsak (1984) officially named the Limnornithidae and Limnornithiformes for their new taxon Limnornis corneti, yet because Limnornis is preoccupied by a recent passeriform, neither family or order name can be used for the Cretaceous taxon. In 1985, Jurcsak and Kessler proposed Cursornithidae and Cursornithiformes for "Paleocursornis biharicus", but the family is not valid as there is no genus "Cursorius" to which it refers. The order is not covered by the ICZN, so could technically be used but would be misleading. Kessler and Jurcsak (1986) finally named Palaeocursornithiformes and Jurcsak and Kessler named Palaeocursornithidae, both for Palaeocursornis biharicus, which is now a junior synonym of Palaeocursornis corneti. If Palaeocursornis is an azhdarchid, Azhdarchidae named in 1984 still has priority.
References- Kessler, 1984. Lower Cretaceous birds from Cornet, Roumania. In Rief and Westphal (eds). Third Symposium on Mesozoic Terrestrial Ecosystems, Tubingen. 119-121.
Kessler and Jurcsak, 1984. Fossil bird remains in the bauxite from Cornet (Romania, Bihor County). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 25, 393-401.
Jurcsak and Kessler, 1985. La paleofaune de Cornet - implications phylogenetiques et ecologiques. Evolution et Adaptation. 2, 137-147.
Kessler and Jurcsak, 1986. New contributions to the knowledge of the Lower Cretaceous bird remains from Cornet (Romania). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 28, 289-295.
Jurcsak and Kessler, 1988. Evolutia avifaunei pe teritoriul Romaneie. III . Filogenie si sistematica. Cistia. 18, 647-688.
Palaeocursornis Kessler and Jurcsak, 1986
= "Limnornis" Kessler, 1984
= Limnornis Kessler and Jurcsak, 1984 (preoccupied Gould, 1839)
= "Palaeolimnornis" Jurcsak and Kessler, 1985
?= "Palaeocursornis" Jurcsak and Kessler, 1985
= Palaeocursornis Kessler and Jurcsak, 1986
= Eurolimnornis Kessler, 1987 (non Kessler and Jurcsak, 1986)
P. corneti (Kessler and Jurcsak, 1984) Bock and Buhler, 1996
= "Limnornis corneti" Kessler, 1984
= Limnornis corneti Kessler and Jurcsak, 1984
= "Palaeolimnornis" corneti (Kessler and Jurcsak, 1984) Jurcsak and Kessler, 1985
?= "Palaeocursornis biharicus" Jurcsak and Kessler, 1985
= Palaeocursornis biharicus Kessler and Jurcsak, 1986
= Eurolimnornis corneti (Kessler and Jurcsak, 1984) Kessler, 1987 (non Kessler and Jurcsak, 1986)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania
Holotype
- (MTCO 14909; = MTCO-P 1637) distal humerus (12 mm wide)
Comments- This specimen has an incredibly complicated taxonomic history, which has been unraveled by Bock and Buhler (1996). Those authors note the relevent papers were published very close together and in a different order than their manuscripts were written, so often reference incorrect dates for each other. In addition, while the holotypes of each valid species were originally thought to be a humerus and femur, Agnolin and Varricchio (2012) have revealed both to be humeri, and both are now catalogued under different numbers than they were when most of the literature was written. Kessler (1984) originally described the holotype (MTCO 14909; = MTCO-P 1637; as a distal femur) and another distal humerus (MTCO 17642; = MTCO-P 7896) as "Limnornis corneti", but did not provide a proper description, making it a nomen nudum. Kessler and Jurcsak (1984) soon officially described the species as Limnornis corneti with the supposed femur as the holotype, but learned the genus was preoccupied by a furnariid passeriform. Jurcsak and Kessler (1985) thus proposed the replacement name "Palaeolimnornis" for their species. However, "Palaeolimnornis" was merely introduced in a list of taxa, making it a nomen nudum. Kessler and Jurcsak (1986) again noted Limnornis was preoccupied, but this time proposed the name Eurolimnornis corneti for the taxon. Oddly, they did not discuss why they did not stick with "Palaeolimnornis" (though they did state it was a synonym), and unfortunately they designated the other distal humerus MTCO 17642 as the holotype. This makes Eurolimnornis corneti a separate species from their original Limnornis corneti based on MTCO 14909. In 1987, Kessler officially proposed Eurolimnornis as a replacement name for Limnornis corneti, though he did not note "Palaeolimnornis" or Kessler and Jurcsak's earlier use of Eurolimnornis. This makes Eurolimnornis corneti of Kessler, 1987 (based on MTCO 14909) a separate taxon (and junior homonym) of Eurolimnornis corneti of Kessler and Jurcsak, 1986 (based on MTCO 17642). Jurcsak and Kessler (1985) listed "Palaeocursornis biharicus" in a faunal list, without reference to a replacement name or holotype, making it a nomen nudum. It is here used as a junior synonym for the species represented by MTCO 14909 because the next year, Kessler and Jurcsak (1986) officially described Palaeocursornis biharicus with MTCO 14909 as the holotype, making biharicus a junior synonym of corneti. However, Palaeocursornis is a valid genus name and thus forms the combination Palaeocursornis corneti, first used by Bock and Buhler (1996). It seems Kessler and/or Jurcsak had intended to switch the holotype of corneti to make MTCO 17642 Palaeolimnornis corneti, later changed to Eurolimnornis corneti, and make MTCO 14909 Palaeocursornis biharicus, but various steps were not allowed by the ICZN, leading to the current situation.
Kessler and Jurcsak (1984) believe the holotype belongs to a ratite, while Kurochkin (1995) assigns it to Palaeognathae. The latter assignment is due to the narrow medial condyle articular surface, wide anterior ridge and "unexpressed crest on the internal supracondyle." Hope (2002) finds the patellar sulcus and large tibiofibular crest indicate it belongs to Ornithurae (sensu Chiappe), and notes that among Aves, tinamiforms and lithornithids also have narrow and deep intercondylar sulci and patellar fossae. Most recently, Agnlin and Varricchio (2012) have reinterpreted it as a pterosaur distal humerus, possibly an azhdarchid.
Three additional specimens were made paratypes of Limnornis corneti by Kessler and Jurcsak (1984). MTCO 17642 (= MTCO-P 7896) is a distal humerus that is now the holotype of the pterosaur Eurolimnornis corneti (see above). MTCO 17956 (= MTCO-P 6966) was described as an ulnar shaft, but was most recently reinterpreted as an indeterminate archosaur long bone shaft by Dyke et al. (2011).
References- Gould, 1839. Birds. In Darwin (ed.). The Zoology of the Voyage of H.M.S. Beagle. 3(3). Smith Elder and Co.. 33-56.
Kessler, 1984. Lower Cretaceous birds from Cornet, Roumania. In Rief and Westphal (eds). Third Symposium on Mesozoic Terrestrial Ecosystems, Tubingen. 119-121.
Kessler and Jurcsak, 1984. Fossil bird remains in the bauxite from Cornet (Romania, Bihor County). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 25, 393-401.
Jurcsak and Kessler, 1985. La paleofaune de Cornet - implications phylogenetiques et ecologiques. Evolution et Adaptation. 2, 137-147.
Kessler and Jurcsak, 1986. New contributions to the knowledge of the Lower Cretaceous bird remains from Cornet (Romania). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 28, 289-295.
Jurcsak and Kessler, 1987. Evolutia avifaunei pe teritoriul Romaneie. II . Morphologia speciilor fosile. Crisia. 17, 583-609.
Kessler, 1987. New contributions to the knowledge about Lower and Upper Cretaceous birds from Romania. In Currie and Koster (eds). Fourth Symposium on Terrestrial Ecosystems. Occasional Papers, Tyrrell Museum of Paleontology. 3, 133-135.
Jurcsak and Kessler, 1988. Evolutia avifaunei pe teritoriul Romaneie. III . Filogenie si sistematica. Cistia. 18, 647-688.
Jurcsack and Kessler, 1991. The Lower Cretaceous paleofauna from Cornet, Bihor County, Romania. Nymphaea. 21, 5-32.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of class Aves. Archaeopteryx. 13, 47-66.
Bock and Buhler, 1996. Nomenclature of Cretaceous birds from Romania. Cretaceous Research. 17, 509-514.
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.
Hope, 2002. The Mesozoic radiation of Neornithes. In Chiappe and Witmer (eds). Mesozoic birds: Above the heads of dinosaurs. Berkeley: University of California Press. 339-388.
Dyke, Benton, Posmosanu and Naish, 2011. Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology. 54(1), 79-95.
Agnolin and Varricchio, 2012 . Systematic reinterpretation of Piksi barbarulna Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of Western USA (Montana) as a pterosaur rather than a bird. Geodiversitas. 34(4), 883-894.

Dinosauromorpha Benton, 1985
Definition- (Megalosaurus bucklandii <- Pterodactylus antiquus) (modified from Sereno, 1991; modified from Kischlat, 2000)
Other definitions- (Lagerpeton chanarensis + Lagosuchus talampayensis + Pseudolagosuchus major + Megalosaurus bucklandii) (modified from Sereno, 1991)
(Passer domesticus <- Pterodactylus antiquus) (modified from Benton, 2004)
(Passer domesticus <- Crocodylus niloticus, Ornithosuchus woodwardi, Pterodactylus antiquus) (Sereno, 2005)
= Dinosauromorpha sensu Benton, 2004
Definition- (Passer domesticus <- Pterodactylus antiquus) (modified)
= Dinosauromorpha sensu Sereno, 2005
Definition- (Passer domesticus <- Crocodylus niloticus, Ornithosuchus woodwardi, Pterodactylus antiquus)

Dinosauromorpha sensu Sereno, 1991
Definition- (Lagerpeton chanarensis + Lagosuchus talampayensis + Pseudolagosuchus major + Megalosaurus bucklandii) (modified)

Lagerpetonidae Arcucci, 1986
Definition- (Lagerpeton chanarensis <- Alligator mississippiensis, Eudimorphodon ranzii, Marasuchus lilloensis, Silesaurus opolensis, Triceratops horridus, Saltasaurus loricatus, Passer domesticus) (Nesbitt, 2011)
Comments- This has recently been commonly misspelled Lagerpetidae (beginning with Nesbitt et al., 2009).
References- Arcucci, 1986. Nuevos materiales y reinterpretacion de Lagerpeton chanarensis Romer (Thecodontia, Lagerpetonidae nov.) del Triasico Medio de La Rioja, Argentina. Ameghiniana. 23(3-4), 233-242.
Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. Journal of Vertebrate Paleontology. 29(2), 498-516.

Lagerpeton

Dromomeron Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007
Diagnosis- (after Nesbitt et al., 2009) concave posterolateral surface of crista tibiofibularis on distal femur; distinct scar on anterior surface of distal femur; posterolateral condyle of proximal tibia ventrally deflected or "hooked."
Comments- Only D. romeri is detailed here, as D. gregorii material has not been assigned to Theropoda.
References- Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science. 317, 358-361.
Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. Journal of Vertebrate Paleontology. 29(2), 498-516.
D. romeri Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007
Norian, Late Triassic
Petrified Forest Member of the Chinle Formation, New Mexico, US

Holotype- (GR 218) femur (98 mm)
Paratypes- ?...(GR 219) femur
?...(GR 220) tibia
(GR 221) partial femur
(GR 222) tibia
(GR 223) astragalocalcaneum
(GR 234) incomplete femur
Referred- (AMNH 2721) distal femur (Cope, 1887)
(AMNH 30648) distal tibia (Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009)
(AMNH 30649) distal tibia (Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009)
(GR 238; GR 235 in Nesbitt, 2011) partial skeleton including femur, tibia, partial fibula, phalanx II-2, pedal ungual II, phalanx III-2, phalanx III-3, phalanx IV-1 and phalanx IV-2 (Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009)
(GR 239; GR 236 in Nesbitt, 2011) tibia (Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009)
(NMMNH P-35379) astragalocalcaneum (Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007)
Diagnosis- (after Irmis et al., 2007) absence of a fourth trochanter; sharp ridge on anteromedial edge of distal femur; large crest on anteromedial edge of astragalus.
(after Nesbitt et al., 2009) lateral tuberosity on anterolateral edge of distal femur.
Comments- The type material of Dromomeron was discovered in 2006 and described by Irmis et al. (2007) as a new taxon of lagerpetonid. Bennett (2015) proposed that D. romeri is actually the juvenile of Tawa hallae based on the shape and development of femoral processes, but that D. gregorii was a valid basal dinosauriform (note lagerpetonids are actually dinosauromorphs however). He does not note that if this were true, Tawa hallae would be a junior synonym of Dromomeron romeri, and that gregorii would need a new genus. However, Bennett fails to engage in the larger picture in that D. romeri is also known from other hindlimb material (some of which is associated in partial skeleton GR 238), that this small material shares lagerpetonid characters with gregorii and Lagerpeton itself, and that is has e.g. fused astragalocalcanea unlike the larger Tawa material. Indeed, Parker (DML, 2015) urged "Please don't put too much stock in Bennett's (2015) ideas about Tawa hallae and Dromomeron romeri forming an ontogenetic series of the same taxon", noting "there is a large amount of unpublished material from the Hayden Quarry as well as other SW NA TR sites that clearly demonstrates that the femoral head anatomy is not due to crushing or a lack of ossification."
AMNH 2721 was originally a syntype of Coelurus bauri (Cope, 1887 first published it as a proximal tibia of Tanystropheus bauri) and was generally retained in Coelophysis bauri (e.g. Huene, 1906; Padian, 1986; though first illustrated by Huene 1915 as Coelophysis longicollis) until that taxon was given a neotype from a different locality in 1996. Since then it has been lumped with Cope's other original Coelophysis material as Coelophysidae indet., or even Archosauria indet. (Nesbitt et al., 2007). Most recently, Rauhut reidentified it as a distal femur and Nesbitt et al. (2009) referred it to Dromomeron romeri. AMNH 30648 and 30649 were also collected in the 1880s as Coelophysis material, and were reidentified as D. romeri by Nesbitt et al. (2009).
References- Cope, 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proceedings of the American Philosophical Society. 24(126), 209-228.
Huene, 1906. Ueber die Dinosaurier der Aussereuropaischen Trias. Geologische und Paläontologische Abhandlungen. 12, 99-156.
Huene, 1915. On reptiles of the New Mexican Trias in the Cope collection. Bulletin American Museum of Natural History. 34, 485-507.
Padian, 1986. On the type material of Coelophysis Cope (Saurischia: Theropoda) and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation). In Padian (ed.). The Beginning of the Age of Dinosaurs: Faunal Change Across the Triassic-Jurassic Boundary. Cambridge University Press. 45-60.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209-243.
Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science. 317, 358-361.
Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. Journal of Vertebrate Paleontology. 29(2), 498-516.
Nesbitt, 2011. The early evolution of archosaurs: Relationships and the origin of major clades. Bulletin of the American Museum of Natural History. 352, 292 pp.
Bennett, 2015. An external mandibular fenestra and other archosauriform characters in basal pterosaurs re-examined. Historical Biology. 27(6), 796-814.
Parker, DML 2015. http://dml.cmnh.org/2015Jul/msg000[xx].html
D. gregorii Nesbitt, Irmis, Parker, Smith, Turner and Rowe, 2009

Dinosauriformes sensu Benton, 2004
Definition- (Passer domesticus <- Lagerpeton chanarensis) (modified)

Dinosauriformes Novas, 1992
Definition-
(Lagosuchus talampayensis + Megalosaurus bucklandii) (modified from Novas, 1992; modified from Kischlat, 2000)
Other definitions-
(Marasuchus lilloensis + Pseudolagosuchus major + Megalosaurus bucklandii) (modified from Novas, 1996)
(Passer domesticus <- Lagerpeton chanerensis) (modified from Benton, 2004)
(Marasuchus lilloensis + Passer domesticus) (Sereno, 2005)
= Dinosauriformes sensu Novas, 1996
Definition- (Marasuchus lilloensis + Pseudolagosuchus major + Megalosaurus bucklandii) (modified)
= Dinosauriformes sensu Sereno, 2005
Definition- (Marasuchus lilloensis + Passer domesticus)

unnamed dinosauriform (Hunt, 1998)
Carnian, Late Triassic
Blue Mesa Member of Chinle Formation, Arizona, US
Material
- (NMMNH coll.) proximal tibia
Comments- Assigned to Theropoda by Hunt (1998), the cnemial crest and posteriorly divided medial and lateral condyles are seen in other dinosauriforms as well (Nesbitt et al., 2007).
References- Hunt, 1998. Preliminary results of the dawn of the dinosaurs project Petrified Forest National Park, Arizona. p. 135–137. in Santucci and McClelland (eds). National Park Service Paleontological Research. National Park Service Technical Report NPS/NRGRD/GRDTR-98/1.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

unnamed dinosauriform (Long and Murray, 1995)
Late Carnian, Late Triassic
Mesa Redondo Member of the Chinle Formation, Arizona, US

Material- (UCMP A269/25793) distal tibia
(UCMP 25834) distal femur (Nesbitt et al., 2007)
Comments- Long and Murry (1995) assigned UCMP 25793 to ?Prosauropoda indet., but Hunt et al. (1998) note it may be a herrerasaurid. Nesbitt et al. (2007) found it to be very similar to Silesaurus and Eoraptor in addition to Herrerasaurus, so referred it to Dinosauriformes indet..
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

Lagosuchia Chatterjee, 1982
Lagosuchidae Bonaparte, 1975
Lagosuchus

Marasuchus

Agnosphitys Fraser, Padian, Walkden and Davis, 2002
A. cromhallensis Fraser, Padian, Walkden and Davis, 2002
Late Triassic
Cromhall Quarry, England
Holotype
- (VMNH 1745) ilium
Paratypes- ?(AUP 11095A) tooth
?(VMNH 1748) astragalus
?(VMNH 1749) astragalus
?(VMNH 1750) humerus
?(VMNH 1751) maxilla
? humerus, humeral fragments, nine partial and incomplete ilia, astragalus
Comments- In the original description, the new genus is listed as Agnosphitys, but the new species is listed as Agnostiphys cromhallensis. The names Agnosphitys and Agnostiphys are then used interchangeably throughout the paper. Fraser (2002) quickly published a note saying Agnosphitys is the intended spelling.
Fraser et al. (2002) believed Agnosphitys to be a dinosauriform less closely related to dinosaurs than Herrerasaurus is. Yet they differ from nearly all other recent authors in placing the latter taxon outside Dinosauria and Saurischia, bringing their conclusions into doubt. More recently, Yates (2007) included Agnosphitys in a cladistic analysis which found it to be a theropod less closely related to Avepoda than Chindesaurus and Guaibasaurus, while Ezcurra (2010) modified that matrix and found it to be a guaibasaurid sauropodomorph. Nesbitt et al. (2015) found the genus to be a basal silesaurid related to Asilisaurus and Lewisuchus in their unpublished analysis.
References- Fraser, Padian, Walkden and Davis, 2002. Basal dinosauriform remains from Britain and the diagnosis of the Dinosauria. Palaeontology. 45(1), 79-95.
Fraser, 2002. Corrigendum. Palaeontology. 45, 843.
Yates, 2007. Solving a dinosaurian puzzle: the identity of Aliwalia rex Galton. Historical Biology. 19(1), 93-123.
Ezcurra, 2010. A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: A reassessment of dinosaur origin and phylogeny. Journal of Systematic Palaeontology. 8(3), 371-425.
Nesbitt, Sidor, Irmis, Stocker, Angielczyk and Smith, 2015. The anatomy of Asilisaurus kongwe (Dinosauriformes: Silesauridae) and closely-related taxa provides new insights into the anatomical and chronological evolution of dinosauriforms. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 187-188.

Nyasasaurus Nesbitt, Barrett, Werning, Sidor and Charig, 2013
= "Nyasasaurus" Charig, 1967 vide Appleby, Charig, Cox, Kermack and Tarlo, 1967
N. parringtoni Nesbitt, Barrett, Werning, Sidor and Charig, 2013
?= Thecodontosaurus alophos Haughton, 1932
= "Nyasasaurus cromptoni" Charig, 1967 vide Appleby, Charig, Cox, Kermack and Tarlo, 1967
Late Anisian, Middle Triassic
Lifua Member of Manda Beds, Tanzania
Holotype
- (BMNH R6856) (~2-3 m, subadult) two partial dorsal vertebrae (38 mm), two partial dorsal centra, partial first primordial sacral vertebra, partial inserted sacral vertrebra (39 mm), partial inserted sacral rib, partial second primordial sacral vertebra (39 mm), partial second primordial sacral ribs, three fragmentary centra, incomplete humerus (~150 mm)
Referred- ?(BMNH R6795; = S 507; material of "Teleocrater rahinus" in partim) anterior cervical vertebra, mid dorsal vertebrae
?(SAM-PKK10654; holotype of Thecodontosaurus alophos) two incomplete anterior cervical vertebrae (73 mm), fragmentary anterior cervical vertebra, two partial mid to posterior dorsal vertebrae (38 mm)
Diagnosis- (After Nesbitt et al., 2013) hyposphene-hypantrum intervertebral articulations in presacral vertebrae; at least three sacral vertebrae; dorsoventrally tall sacral ribs; ventrally elongated deltopectoral crest; laterally deflected apex of deltopectoral crest; distinct notch central to apex of deltopectoral crest; pointed expansion on proximal surface near dorsal extent of deltopectoral crest; proximal surface of humerus continuous with lateral
surface of deltopectoral crest; distinct fossa present on posterodorsal surface, just ventral to proximal surface.
Other diagnoses- Haughton (1932) distinguished Thecodontosaurus alophos from Coelophysis? longicollis and unspecified South African taxa based on its lack of a ventrally keeled anterior cervical centrum.
Charig in his original unpublished description of Nyasasaurus diagnosed it based on two characters (at least three sacral vertebrae; deltopectoral crest extends for 47% of humeral length) which Nesbitt et al. (2011) state are widespread in basal dinosaurs.
Comments- The holotype of Thecodontosaurus alophos was discovered in the early 1930s and described by Haughton (1932) as a thecodontosaurian theropod similar to Coelophysis? longicollis. It was believed to be synonymous with Nyasasaurus parringtoni by Nesbitt et al (2013) based on the similarity of the dorsal vertebrae, similar position when entered into Nesbitt's matrix and recovery from the same horizon. However, they share no published autapomorphies or combination of characters, so this is tentative. It should be noted alophos has priority over parringtoni in case the former proves to be diagnostic in the future.
Nyasasaurus was originally described by Charig (1957) in his thesis as Specimen 50b and referred to Prestosuchidae. Appleby et al. (1967) credited the name Nyasasaurus cromptoni to Charig as a thecodontosaurid or possibly prestosuchid, though without a diagnosis this is a nomen nudum. White (1973) listed it as an anchisaurid. Nesbitt et al. (2013) strated Charig's unpublished manuscript named this Nyasasaurus parringtoni and believed it to be a dinosaur or close relative. They later officially named the taxon, 56 years after Charig's thesis.
Nesbitt et al. (2013) note the vertebrae of "Teleocrater rhadinus" are very similar to Nyasasaurus, but the appendicular and other elements appear to belong to another individual.
Nesbitt et al. (2013) recovered the holotype as either the sister taxon of Dinosauria, most basal ornithischian or a dilophosaurid. The Thecodontosaurus alophos specimen was found as the sister taxon of Tawa+avepods. When combined, the resulting OTU is closer to Dinosauria than silesaurids, but outside examined ornithischians, sauropodomorphs and Tawa+avepods.
References- Haughton, 1932. On a collection of Karroo vertebrates from Tanganyika Territory. The Quarterly Journal of the Geological Society of London. 88, 634-671.
Charig, 1957. New Triassic archosaurs from Tanganyika, including Mandasuchus and Teleocrater. PhD hesis, Cambridge University.
Appleby, Charig, Cox, Kermack and Tarlo, 1967. Reptilia. In Harland, Holland, House, Hughes, Reynolds, Rudwick, Satterthwaite, Tarlo and Willey (eds.). The fossil record: A symposium with documentation. 695-731.
White, 1973. Catalogue of the genera of dinosaurs. Annals of the Carnegie Museum. 44, 117-155.
Nesbitt, Barrett, Werning, Sidor and Charig, 2013. The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania. Biology Letters. 9(1), 20120949.

Saltopus Huene, 1910
S. elginensis Huene, 1910
Early Norian, Late Triassic
Lossiemouth Sandstone Formation, Scotland

Holotype- (BMNH 2915) (~800-1000 mm) two centra, anterior dorsal centrum (7 mm), anterior dorsal centrum (7.5 mm), partial fifth dorsal centrum (9 mm), sixth dorsal centrum (9 mm), seventh dorsal centrum, eighth dorsal centrum, ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal centrum, thirteenth dorsal centrum, fourteenth dorsal centrum, fifteenth dorsal centrum, six partial dorsal ribs, first sacral vertebra (~10 mm), second sacral vertebra (~10 mm), sacral ribs, twenty-four caudal vertebrae (proximal 9-10 mm, distal 11 mm), chevrons, scapula (~28 mm), scapular or humeral fragment, humerus (36 mm), radius (25 mm), ulna (25 mm), centrale, three distal carpals, metacarpal I, metacarpal II, metacarpal III, partial phalanx III-1, metacarpal IV (7 mm), metacarpal V (5 mm), ilia (26, 23 mm), pubis (33 mm), ischia (~35-40 mm), femora (47 mm), tibiae (66 mm), fibulae, astragali, calcanea, distal tarsal IV, metatarsal II (35 mm), phalanx II-1, phalanx II-2, metatarsal III (38 mm), partial phalanx III-1 (12 mm), partial phalanx III-2 (8.5 mm), partial phaalnx III-3 (7 mm), partial pedal ungual III, metatarsal IV (35 mm), phalanx IV-1, phalanx IV-2 (5 mm), phalanx IV-3 (5 mm), partial phalanx IV-4 (4 mm), pedal ungual IV (4 mm), metatarsal V (15 mm)
Comments- The holotype was shown to Huene in 1909. Historically, Saltopus was generally classified as a primitive theropod until being ignored by most authors through the 90s and early 2000s. In an abstract, Benton and Walker (2009) determined it was an avemetatarsalian but could not exclude it from Dinosauria. Benton and Walker (2011) redescribed the holotype, and found it was between Pseudolagosuchus and Silesauridae+Dinosauria in one matrix, and sister to Marasuchus in another. When analyzed in Nesbitt's archosauriform matrix, it emerges as a member of the silesaurid+dinosaur clade excluded from Eucoelophysis plus more derived silesaurids, Eusaurischia and known Ornithischia.
References- Huene, 1910. Ein primitiver Dinosaurier aus der mittleren Trias von Elgin. Geologische und Palaontologische Abhandlungen, Neue Folgung. 8, 315-322.
Benton and Walker, 2009. The enigmatic Late Triassic reptile Saltopus. Journal of Vertebrate Paleontology. 29(3), 62A.
Benton and Walker, 2011. Saltopus, a dinosauriform from the Upper Triassic of Scotland. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 285-299.

Silesauridae Langer, Ezcurra, Bittencourt and Novas, 2010
Definition-
(Silesaurus opolensis <- Passer domesticus, Triceratops horridus, Alligator mississippiensis) (Nesbitt, 2011)
= "Lewisuchinae" Paul, 1988

Lewisuchus
= Pseudolagosuchus

Eucoelophysis Sullivan and Lucas, 1999
E. baldwini
Sullivan and Lucas, 1999
Norian, Late Triassic
Petrified Forest Member of the Chinle Formation, New Mexico, US

Holotype- (NMMNH P-22298) (subadult) two posterior cervical vertebrae (33.7 mm), dorsal vertebral fragments, nearly complete proximal caudal centrum, three incomplete distal caudal centra, incomplete scapulocoracoid, ilial fragment, incomplete pubis, partial ischium, partial femora (~210 mm), proximal tibia, incomplete metatarsal II, metatarsal III (91 mm), incomplete metatarsal IV, pedal phalanges
Paratype- ?(AMNH 2704; syntype of Coelurus longicollis) femur (215 mm) (Cope, 1887)
Diagnosis- (modified from Sullivan and Lucas, 1999) ischio-acetabular groove in proximal pubis that pierces the medial pubic wall; tibia has large appressed surface on distolateral side for contact with fibula.
(modified from Ezcurra, 2006) non-invasive pleurocoels in the cervical vertebrae (also in Avepoda and derived Sauropoda); fourth trochanter reduced in femur (preservational?); cnemial crest distinctively offset from the tibial shaft, cranially straight, and without lateral notch.
Comments- Hunt and Lucas (1991) felt Cope's original Coelophysis specimens (including the lectotypes of C. bauri, C. longicollis and C. willistoni) were indeterminate, and created the name Rioarribasaurus colberti for the abundant complete Ghost Ranch specimens. However, the ICZN later (1996) ruled a Ghost Ranch specimen is the neotype of Coelophysis bauri. Sullivan and Lucas (1999) described a new diagnostic specimen from the horizon of Cope's originals as Eucoelophysis baldwini, referring AMNH 2706 to the taxon and suggesting the rest of Cope's specimens may belong to it as well (though they were viewed as generically indeterminate). However, Nesbitt et al. (2005, 2007) and Ezcurra (2006) have determined the holotype of Eucoelophysis is from a basal dinosauriform, not a coelophysoid. It was assigned to Ceratosauria based on triangular dorsal transverse processes and a trochanteric shelf, both of which it lacks. Eucoelophysis also lacks the following characters- well developed, strongly inturned femoral head (Dinosauria); trochanteric fossa on femoral head (Dinosauria); oblique ligamental groove on caudal surface of femoral head (Avepoda); femoral medial epicondyle well developed (Avepoda); caudal cleft between medial and lateral condyle on the proximal end of the tibia (Dinosauriformes); tibial cnemial crest well developed (Avepoda); fibular crest on tibia (Avepoda).
The pubis AMNH 2706 referred to Eucoelophysis by Sullivan and Lucas appears to be coelophysoid as well based on the pubic foramen, despite the supposed ischio-acetabular groove (which is more similar to Saturnalia in not being open laterally).
The femur AMNH 2704 was originally a syntype of Coelurus longicollis (Cope, 1887a), later renamed Coelophysis longicollis. Though the specimen is generally identified as an indeterminate theropod (Hunt and Lucas, 1991), it seems to be a silesaurid. Cope noted the lack of an ectocondylar tuber, and in addition the head is not offset from the shaft and the anterior trochanter is poorly developed. Besides stratigraphy, the reduced fourth trochanter may indicate it is referrable to Eucoelophysis.
References- Cope, 1887. The dinosaurian genus Coelurus. American Naturalist. 21, 367-369.
Huene, 1915. On reptiles of the New Mexican Trias in the Cope collection. Bulletin American Museum of Natural History. 34, 485-507.
Hunt and Lucas, 1991. Rioarribasaurus, a new name for a Late Triassic dinosaur from New Mexico (USA). Paläontol. Z. 65 p. 191-198.
International Commision on Zoological Nomenclature, 1996. Opinion 1842. Coelurus bauri Cope, 1887 (currently Coelophysis bauri; Reptilia, Saurischia): lectotype replaced by a neotype. Bulletin of Zoological Nomenclature. 53, 142-144.
Sullivan and Lucas, 1999. Eucoelophysis baldwini, a new theropod dinosaur from the Upper Triassic of New Mexico, and the status of the original types of Coelophysis. Journal of Vertebrate Paleontology 19(1): 81-90.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. JVP. 25(3) 96A.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas. 28 (4),649-684.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

unnamed silesaurid (Long and Murray, 1995)
Late Triassic
Dockum Group, Texas, US
Material
- (TMM 31100-185) femur
Comments- Originally described by Long and Murray (1995) as a possible ornithosuchid, this was reidentified by Nesbitt et al. (2007) as a basal dinosauriform closely related to Silesaurus, Lewisuchus and Eucoelophysis.
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

unnamed silesaurid (Parker, Irmis and Nesbitt, 2006)
Carnian, Late Triassic
Blue Mesa Member of Chinle Formation, Arizona, US
Material
- (PEFO 34347) proximal femur
Reference- Parker, Irmis and Nesbitt, 2006. Review of the Late Triassic dinosaur record from Petrified Forest National Park, Arizona. Museum of Northern Arizona Bulletin. 62, 160-161.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

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)
Comments- This has a broader application than Theropoda due to the branched structures in Kulindadromeus.

Dinosauria Owen, 1842
Definition- (Hylaeosaurus armatus + Megalosaurus bucklandii) (modified from Kischlat, 2000)
Other definitions- (Herrerasaurus ischigualastensis + Allosaurus fragilis + Stegosaurus armatus) (modified from Novas, 1992)
(Triceratops horridus + Passer domesticus) (Sereno, 2004; modified from Padian and May, 1993)
(Iguanodon bernissartensis + Megalosaurus bucklandii) (Clarke et al., 2004; modified from Olshevsky, 2000)
= Pachypodes Meyer, 1845 preoccupied Erichson, 1840
= Pachypoda Troschel, 1867 preoccupied Gray, 1821
= Ornithoscelida Huxley, 1869
= Prodinosauria Haeckel, 1895
= Phytodinosauria Bakker, 1986
= Eudinosauria Novas, 1992
Definition- (Allosaurus fragilis + Stegosaurus armatus) (modified from Novas, 1992)
= Dinosauria sensu Novas, 1992
Definition- (Herrerasaurus ischigualastensis + Allosaurus fragilis + Stegosaurus armatus) (modified)
= Dinosauria sensu Padian and May, 1993
Definition- (Triceratops horridus + Passer domesticus) (modified)
= Dinosauria sensu Olshevsky, 2000
Definition- (Iguanodon bernissartensis + Megalosaurus bucklandii) (modified)
Comments- Although definitions using birds as a specifier have been widespread since Padian and May (1993), birds were not viewed as dinosaurs in their initial formulation by Owen, nor by the vast majority of workers until the 1970's and later. Thus, a definition involving two of Owen's original dinosaurs is preferred here.
Meyer (1845) used Pachypodes as an alternative to Dinosauria (for Megalosaurus, Plateosaurus, Hylaeosaurus and Iguanodon), but this is preoccupied by a group of beetles (Erichson, 1840). Similarly, Troschel (1867) and several later authors used Pachypoda as an alternative to Dinosauria, but the name is preoccupied by a bivalve group (Gray, 1821). Haeckel (1895) erected Prodinosauria for the most basal grade of dinosaurs, but did not indicate which taxa belonged.
References- Gray, 1821. A natural arrangement of Mollusca, according to their internal structure. London Medical Repository. 15, 229-239.
Erichson, 1840. Entomographien, Untersuchungen in dem Gebiete der Entomologie mit besonderer Benutzung der Königl. F. H. Morin, Berlin. 180 pp.
Meyer, 1845. System der fossilen Saurier. Neues Jahrbuch fur Mineralogie, Geologie und Palaontologie. 1845, 278-285.
Troschel, 1867. Bericht über die Leistungen in der Herpetologie während des Jahres 1866. Archiv Fur Naturgeschichte. 3(2), 33-41.
Haeckel, 1895. Systematische Phylogenie der Wirbelthiere: (Vertebrata). 660 pp.

Dinosauria incertae sedis

Dinosauria indet.

Apatodon Marsh, 1877
A. mirus Marsh, 1877
Tithonian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US
(Marsh-Felch Quarry 1)
Holotype- (YPM coll.; lost) partial neural spine
Comments- While Marsh only specified the locality to the "Rocky Mountains region" of the US, Monaco (1998) noted it came from Marsh-Felch Quarry 1 in Colorado.
Discovered in 1877, Marsh originally described Apatodon mirus that year as a partial dentary "with the last molar in place." Marsh states it was "about as large as a tapir" and that the supposed tooth resembled a suid most in form and crown surface, but that the structure was different and the "fangs" at least partially fused with the dentary. The molar was stated to be 8 mm apicobasally, 41 mm mesiodistally and 20 mm labiolingually. As for its identification, while Marsh said it was "very unlike any corresponding jaw of a dinosaur" and "suggests the mammalian type", he also stated it was "widely different from anything yet described, and its exact affinities are doubtful." The specimen has never been illustrated or described further.
Baur (1890) viewed the specimen, determining it to be a weathered partial dinosaur neural spine, "some parts of which looked something like a tooth of a hog." He states Marsh knew of this identification "long ago", but never published a correction.
Meyer (1890) also saw the holotype and stated he considered it "in the same way as other assistants of Professor Marsh do, not as the jaw of a mammal, but as a piece of a vertebrae of a Dinosaur." He also noted "Marsh is apparently of the same opinion, for in his "list of genera," printed as manuscript, Apatodon is enumerated as Dinosaur. The question is only Why has Professor Marsh not published corrections to his previous statements? and has he recognized himself the true nature of the fossil, or has it been pointed out to him? One of his assistants has made to me, and in my presence to somebody else, the positive statement that he has demonstrated this to the professor."
After this, we have no record of original viewings of the specimen. Hay (1902) listed it below Ornithopoda between Macelognathus and Brachyrophus while stating it was "a doubtful genus of doubtful position." Huene (1909; as incertae sedis), Abel (1910; as incertae sedis) and Mook (1916) also placed it in Ornithischia, while Zittel (1911) placed it specifically in Stegosauridae. Hay (1930) listed it under diplodocids, this time between Dystrophaeus and Brachyrophus. Kuhn (1939) listed it as a sauropod, between the titanosaur and diplodocid sections. Steel (1970) and Casanovas et al. (1987) listed it as a titanosaur. Chure and McIntosh (1989) incorrectly indicated Marsh originally identified Apatodon as a theropod. Olshevsky (1991) synonymized it with Allosaurus fragilis, as did Glut (1997). None of these authors justified their referrals in any way or indicated they had further resources for the specimen.
McIntosh (pers. comm. 1997 to Chure, 2000) and Glut both say the specimen cannot be found in the YPM collections.
A Textual Investigation- The phylogenetic assignments of every author before Chure and McIntosh are most easily explained by them copying Hay. Huene, Abel and Zittel kept the trio of Apatodon, Macelognathus and Brachyrophus together in their schemes, all following Hay's original placements. Mook also kept Apatodon and Macelognathus together, but had Camptosaurus inserted between them and Brachyrophus. Abel listed the three underneath the stegosaur genera, which Zittel probably misunderstood to indicate they were stegosaurs. Hay then kept it next to Brachyrophus 28 years later (Macelognathus having been moved to Theropoda) and before Alamosaurus, which Kuhn copied. Kuhn's pairing of Apatodon+Brachyrophus was placed between Campylodon and Alamosaurus, which probably inspired Steel to place Apatodon in Titanosauridae, as those two genera belong to that family. This was in turn copied by Casanovas et al. in their review of titanosaur genera. Notably, the latter two papers repeat Marsh's description of the material as a jawbone as if it were accurate, so widespread memory of Baur's and Meyer's reidentification had apparently faded by the 70s.
Given these patterns, the only real questions revolve around why Hay listed it in the places he did, as everyone's copying leads back to him. In fact, it seems likely Hay (1902) never intended to classify Apatodon as an ornithischian in the first place. Macelognathus, Apatodon and Brachyrophus (in that order) are the last dinosaurs listed in his paper, each with a comment stating their position is problematical or doubtful. Macelognathus is listed under Macelognathidae, and the lack of any other heading between it and the other two taxa could naively suggest Hay intended them all to be macelognathids. Yet they are known from incomparable parts of the skeleton (dentaries, a neural spine fragment, and centra respectively) and had no prior history together, so this seems unlikely (Macelognathus has since proven to be a basal crocodylomorph, while I believe Brachyrophus may be based on juvenile sauropod caudal centra). Given the lack of a heading separating Macelognathus/idae from Apatodon, it's likely the lack of a heading above Macelognathidae doesn't mean that family or the two subsequent genera were supposed to be in Ornithopoda, or even Ornithischia (as Huene assumed). Indeed, it may be none were even supposed to be in 'order' Dinosauria, as Hay mentions Marsh assigned Macelognathus to the order Macelognatha and states of Brachyrophus "even the ordinal position of this genus is doubtful." Pages 505 and 506 between Dinosauria and Pterosauria may have just been a convenient place to list three Morrison reptiles of uncertain ordinal affinities. Why did Hay then move Apatodon and Brachyrophus to Sauropoda in 1930? These two genera are the last saurischians listed before Orthopoda (=Ornithischia) except for Alamosaurus being listed below them. It's plausible Alamosaurus' placement was a mistake and that as in 1903, the two genera were supposed to be listed between groups, in this case Saurischia and Orthopoda. As such, they would be Dinosauria incertae sedis. In conclusion, Hay's confusing method of listing taxa led to incorrect assumptions by future workers, and all identifications more specific than Dinosauria are ultimately based on these errors.
Olshevsky (pers. comm., 2015) based his synonymy with Allosaurus on a tip from Molnar, which in turn was originally suggested by McIntosh (pers. comm. to Molnar, 2015). McIntosh is said to have located a paper claiming Marsh mistook an Allosaurus neural spine for a mammal jaw, which would be a probable reference to the Apatodon type. Unfortunately, neither Olshevsky nor Molnar remember the citation for this paper and I have not located it or heard back from McIntosh. As such, this is an intriguing idea that remains unconfirmed. It may also mean Chure and McIntosh's assignment of Apatodon to Theropoda was purposeful, even if the asterisk indicating Marsh thought this was a mistake. Notably, Allosaurus' mid dorsal neural spines do possess interspinous ligament protrusions which are somewhat molar-like, so could be confused for a tooth fused to the jawbone. Also, the holotype of Allosaurus was recovered from the same quarry during the same year, so it's possible Apatodon actually belongs to the holotype individual of Allosaurus fragilis. Even if we had the specimen though, it's unlikely a partial dorsal neural spine could be identified more precisely than Allosauridae.
Ignoring the so far unproven association with Allosaurus, Apatodon could be from any dinosaurian clade. Given a tapir's dentary is ~400 mm long, even a piece of such a bone would have to come from a large vertebra. No other large tetrapods are known from the Morrison Formation, so a dinosaurian identity is likely even considering the poor state of knowledge in the 1890s. Besides Allosaurus, the holotype of Diplodocus was also recovered from Marsh-Felch Quarry 1 that year, as well as material referred to Apatosaurus, Haplocanthosaurus and Stegosaurus.
References- Marsh, 1877. Notice of some new vertebrate fossils. American Journal of Arts and Sciences. 14, 249-256.
Baur, 1890. A review of the charges against the paleontological department of the U.S. Geological Survey, and of the defence made by Prof. O. C. Marsh. The American Naturalist. 24(1), 298-304.
Meyer, 1890. Some more nuts for Marsh to crack. Yew York Herald. January 26 1890, p. 25.
Hay, 1902. Bibliography and catalogue of the fossil Vertebrata of North America. Bulletin of the United States Geological Survey. 179, 1-868.
Huene, 1909. Skizze zu einer Systematik und Stammesgeschichte der Dinosaurier. Centralblatt für Mineralogie, Geologie und Paläontologie. 1909, 12-22.
Abel, 1910. Die Rekonstruktion des Diplodocus. Abhandlungen der kaiserlichen und koeniglichen Zoologisch-Botanischen Gesellschaft in Wien. 5, 1-60.
Zittel, 1911. Grundzüge der Paläontologie (Paläozoologie). II. Abteilung. Vertebrata. Druck und Verlag von R. Oldenbourg, München. 1-598.
Mook, 1916. A study of the Morrison Formation. Annals of the New York Academy of Sciences. 27(1), 39-191.
Hay, 1930. Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution of Washington. 390(II), 1-1074.
Kuhn, 1939. Saurischia. In Fossilium Catalogus I. Animalia. 87. 124 pp.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart 1-8.
Casanovas, Santafé, Sanz and Buscalioni, 1987. Arcosaurios (Crocodilia, Dinosauria) del Cretácico superior de la Conca de Tremp (Lleida, España). Estudios Geologicos, Volumen Extraordinario Galve-Tremp. 95-110.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Glut, 1997. Dinosaurs - The Encyclopedia. McFarland Press, Jefferson, NC. 1076 pp.
Monaco, 1998. A short history of dinosaur collecting in the Garden Park fossil area, Canon City, Colorado. Modern Geology. 23, 465-480.
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. PhD thesis. Columbia University. 964 pp.

Praeornithes Rautian, 1978
Praeornithiformes Rautian, 1978
Praeornithidae Rautian, 1978
Praeornis Rautian, 1978
P. sharovi Rautian, 1978
Late Callovian-Kimmeridgian, Middle-Late Jurassic
Karabastau (= Balabansai) Formation, Kazakhstan
Holotype
- (PIN 2585/32) feather?
Referred- ?(ZPAL V 32/967) feather? (Dzik, Sulej and Niedzwiedzki, 2010)
Comments- Discovered in 1971, Sharov labeled the holotype 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) thought 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. Most recently, another specimen was referred to Praeornis by Dzik et al. (2010). The authors found it differs from plant fossils in preservation and carbon isotope values. Yet it differs from feathers in having three vanes, an expanded distal tip to the shaft, and lack of basal narrowing. While seeming to support an animal identification for these structures, the resemblence to feathers seems limited to its basic shape. Considering the diversity of integument now known in ornithischians (e.g. ribbon-like tibial structures of Kulindadromeus, caudal quills of Psittacosaurus), Praeornis could easily belong to this clade as well. As such, it is assigned to Dinosauria incertae sedis at present.
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.
Dzik, Sulej and Niedzwiedzki, 2010. Possible link connecting reptilian scales with avian feathers from the early Late Jurassic of Kazakstan. 22(4), 394-402.

undescribed dinosaur (Lydekker, 1888)
Early-Middle Oxfordian, Late Jurassic
Upper Oxford Clay, England

Material- (BMNH 40517) distal fibula
Comments- This was first mentioned by Lydekker (1888) as Omosaurus? sp., then referred to Lexovisaurus durobrivensis by Galton (1985). Pickering (unpublished ms) has referred it to Metriacanthosaurus parkeri. It was most recently called Dinosauria indet. by Maidment et al. (2008).
References- Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamta, Rhynchocephalia, and Proterosauria: British Museum of Natural History, London. 309 pp.
Galton, 1985. British plated dinosaurs (Ornithischia, Stegosauridae). Journal of Vertebrate Paleontology. 5, 211-254.
Maidment, Norman, Barrett and Upchurch, 2008. Systematics and phylogeny of Stegosauria (Dinosauria: Ornithischia). Journal of Systematic Palaeontology. 6, 367-407.

unnamed dinosaur (Huene, 1929)
Campanian-Maastrichtian, Late Cretaceous
Allen Formation, Rio Negro, Argentina
Material
- (MLP CS 1240) metatarsal IV (100 mm)
Comments- Carnosaurus was listed by Huene (1929) in a faunal list for three specimens- a metapodial (MLP CS 1240) from the Allen Formation, a tooth (MACN coll.) perhaps from the Chubut group, and provisionally ("Cf. Carnosaurus") a tooth (MLP coll.) from the Plottier Formation. As Olshevsky (DML, 1999) noted, the name is probably a typographical error for Carnosauria made when translating the paper from German to Spanish. This is indicated by the fact he never attaches a name to these specimens in the description or plates, and indeed states on of the specimens is taxonomically indistinguishable from another named genus. Since "Carnosaurus" was apparently not meant as a valid name when it was published (ICZN Article 11.5), it is a nomen nudum.
MLP CS 1240 is listed as metatarsals in the faunal list, but described as a ?third metacarpal similar to Allosaurus. However, it is dissimilar to theropod metacarpals in several respects. The distal articular surface is not ginglymoid, unlike metacarpals I-III of ceratosaurs and basal tetanurines or I-II of avetheropods. There is no extensor pit dorsally unlike theropod metacarpals I and II, and ceratosaur metacarpal III. The shaft is much more robust than tetanurine metacarpal IIIs except for Torvosaurus, while the proximal end is less expanded than theropod metacarpal IIs, and the shaft is far more elongate than most theropod metacarpal Is. The form is more similar to a sauropod or ankylosaur metatarsal, both groups that are known from the Allen Formation. It is here assigned to Dinosauria indet. pending further comparisons.
References- Huene, 1929. Los saurisquios y ornitisquios del Cretacéo Argentino. Anales del Museo de La Plata (series 3). 3, 1-196.
Olshevsky, DML 1999. http://dml.cmnh.org/1999Nov/msg00507.html

unnamed dinosaur (Long and Murry, 1995)
Late Carnian, Late Triassic
Colorado City Member of Duckum Formation, Texas, US

Material- proximal femur
Description- Perpendicular and offset femoral head; similar to Chindesaurus in gross morphology.
Comments- Originally referred to Chindesaurus by Long and Murry (1995), but removed by Hunt et al. (1998). Other reported Coelophysis and coelurosaurian material from the formation is Trilophosaurus.
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.

undescribed dinosaur (Long and Murry, 1995)
Late Carnian, Late Triassic
Bluewater Creek Formation, Arizona, US

Material- teeth
Comments- Assigned by Long and Murry (1995) to cf. Coelophysis, but reassigned more generally to Dinosauria indet. by Hunt et al. (1998).
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.

unnamed dinosaur (Long and Murray, 1995)
Norian, Late Triassic
Petrified Forest Formation of the Chinle Group, Arizona, US

Material- (UCMP 126751) distal femur
Comments- Referred to Chatterjeea (= Shuvosaurus) by Long and Murray (1995), Parker et al. (2006) assigned it to Dinosauria indet. based on the fibular groove which opens with an obtuse angle and the strongly rounded margin of the fibular condyle in this groove.
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Parker, Irmis and Nesbitt, 2006. Review of the Late Triassic dinosaur record from Petrified Forest National Park, Arizona. Museum of Northern Arizona Bulletin. 62, 160-161.

unnamed dinosaur (Novas, Chatterjee, Ezcurra and Kutty, 2009; described by Novas, Ezcurra, Chatterjee and Kutty, 2011)
Late Norian-Early Rhaetian, Late Triassic
Upper Maleri Formation, India
Material
- (ISI R282) first sacral vertebra, third sacral vertebra, proximal caudal vertebra, ilium, pubes (one partial, one proximal)
Comments- Novas et al. (2009) said this resembles herrerasaurs, but Novas et al. (2011) later described it as a non-sauropodomorph dinosauriform. It emerged as a dinosaur outside Eoraptor+Chindesaurus+Tawa+Avepoda and Guaibasauridae+more derived sauropodomorphs when entered into Yates' sauropodomorph matrix.
References- Novas, Chatterjee, Ezcurra and Kutty, 2009. New dinosaur remains from the Late Triassic of Central India. Journal of Vertebrate Paleontology. 29(3), 156A.
Novas, Ezcurra, Chatterjee and Kutty, 2011. New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of Central India. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 333-349.

unnamed dinosaur (Novas, Chatterjee, Ezcurra and Kutty, 2009; described by Novas, Ezcurra, Chatterjee and Kutty, 2011)
Late Norian-Early Rhaetian, Late Triassic
Upper Maleri Formation, India
Material
- (ISI R284) ilium
Comments- This was mentioned by Novas et al. (2009) as a basal saurischian of uncertain affinities. It was later described even less securely by Novas et al. (2011) as a dinosauriform. They noted it resembled Nambalia so could belong to it, couldn't be compared to Jaklapallisaurus or Pradhania, but differed from Lamplughsaura, putative guaibasaurid ISI R277, and dinosaur ISI R282.
References- Novas, Chatterjee, Ezcurra and Kutty, 2009. New dinosaur remains from the Late Triassic of Central India. Journal of Vertebrate Paleontology. 29(3), 156A.
Novas, Ezcurra, Chatterjee and Kutty, 2011. New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of Central India. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 333-349.

Ornithischia
Definition- (Stegosaurus armatus <- Allosaurus fragilis) (modified from Kischlat, 2000)
Other definitions- (Triceratops horridus <- Passer domesticus) (Sereno, 2004; modified from Sereno, 1998; modified from Padian and May, 1993)
(Iguanodon bernissartensis <- Cetiosaurus medius) (Norman et al., 2004)
(Iguanodon bernissartensis <- Megalosaurus bucklandii, Cetiosaurus medius) (Wagner, 2004)
(Triceratops horridus <- Tyrannosaurus rex) (modified from Weishampel, 2004)
(Triceratops horridus <- Passer domesticus, Saltasaurus loricatus) (Sereno, 2005)
= Ornithischia sensu Padian and May, 1993
Definition- (Triceratops horridus <- Passer domesticus)
= Ornithischia sensu Norman et al., 2004
Definition- (Iguanodon bernissartensis <- Cetiosaurus medius)
= Ornithischia sensu Wagner, 2004
Definition- (Iguanodon bernissartensis <- Megalosaurus bucklandii, Cetiosaurus medius)
= Ornithischia sensu Weishampel, 2004
Definition- (Triceratops horridus <- Tyrannosaurus rex) (modified)
= Ornithischia sensu Sereno, 2005
Definition- (Triceratops horridus <- Passer domesticus, Saltasaurus loricatus)

Thecospondylus

Tichosteus
T. lucasanus
T? aequifacies

Genasauria

Sarcolestes

Struthiosaurus

Neornithischia
Definition- (Stegosaurus armatus + Triceratops horridus) (modified from Kischlat, 2000)

Fulgurotherium

Loncosaurus

Kulindadromeus Godefroit, Sinitsa, Dhouailly, Bolotsky, Sizov, McNamara, Benston and Spagna, 2014
= Daurosaurus Alifanov and Saveliev, 2014a
= Kulindapteryx Alifanov and Saveliev, 2014a
= Lepidocheirosaurus Alifanov and Saveliev, 2015
K. zabaikalicus Godefroit, Sinitsa, Dhouailly, Bolotsky, Sizov, McNamara, Benston and Spagna, 2014
= Daurosaurus olovus Alifanov and Saveliev, 2014a
= Kulindapteryx ukureica Alifanov and Saveliev, 2014a
= Lepidocheirosaurus natatilis Alifanov and Saveliev, 2015
Bajocian-Tithonian, Middle-Late Jurassic
Ukureyshaya Formation, Russia
Holotype
- (INREC K3/109) (juvenile) incomplete skull, incomplete mandible
Paratypes- (INREC K3/112) dentary or dorsal vertebra
(INREC K3/113) ilium
(INREC K3/114) incomplete pubis
(INREC K3/124) ischium
(INREC K3/134) incomplete humerus (mislabeled as scapula)
(INREC K3/200) dentary
(INREC K3/202) distal caudal vertebra
(INREC K3/203; PIN 5434/54) scapula, humerus
(INREC K3/204) partial scapula
(INREC K3/205) ulna
(INREC K3/206) femur
(INREC K3/207) tibia
(INREC K3/208) proximal femur
(INREC K3/209) proximal tibia
(INREC K4/22) partial skull, fragmentary dentary, sclerotic plates, feathers
(INREC K4/33) dorsal ribs, scapula, feathers
(INREC K4/42) maxilla
(INREC K4/44) proximal tibia, feathers
(INREC K4/57) tibia, tarsus, scales
(INREC K4/72) metatarsal I, partial metatarsal II, partial metatarsal III, partial metatarsal IV
(INREC K4/94) proximal caudal vertebrae, scales
(INREC K4/116) femur, scales, feathers
(INREC K4/117) at least six proximal caudal vertebrae, scales
(INREC K4/118) metatarsal fragments, pedal phalanges, scales
(INREC K4/150) five proximal caudal vertebrae
(INREC K4/159; PIN 5435/51; paratype of Lepidocheirosaurus natatilis) eleven distal caudal vertebrae, scales
(INREC K4/201) partial mandible
(INREC K coll.) four partial skulls, several hundred specimens including cranial elements, pedal phalanx I-1, pedal ungual I, phalanx II-1, pedal ungual II, phalanx III-1, pedal ungual III, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal claw sheaths
(PIN 5434/57; 5435/57 in Alifanov et al., 2014; = INREC K4/115) humerus, proximal radius, proximal ulna, scales, feathers
Referred- (PIN 5434/1; holotype of Daurosaurus olovus) ilium (100 mm), ischial fragment, femur (120 mm), tibia (155 mm)
(PIN 5434/2; paratype of Lepidocheirosaurus natatilis) distal caudal vertebra (Alifanov and Saveliev, 2015)
(PIN 5434/17; paratype of Daurosaurus olovus) ilium (Alifanov and Saveliev, 2014a,b)
(PIN 5434/19) partial dentary (Alifanov, 2014)
(PIN 5434/20; paratype of Daurosaurus olovus) ilium (Alifanov and Saveliev, 2014a,b)
(PIN 5434/24) pubis (Alifanov and Saveliev, 2014a,b)
(PIN 5434/25; holotype of Kulindapteryx ukureica) fragmentary proximal caudal vertebra, incomplete pubis, ischium (104 mm) (Alifanov and Saveliev, 2014a,b)
(PIN 5434/37; paratype of Daurosaurus olovus) ilium (Alifanov and Saveliev, 2014a,b)
(PIN 5434/39; paratype of Daurosaurus olovus) ischium (Alifanov and Saveliev, 2014a,b)
(PIN 5434/40; paratype of Daurosaurus olovus) ischium (90 mm) (Alifanov and Saveliev, 2014a,b)
(PIN 5434/41) ilium (Alifanov and Saveliev, 2014a,b)
(PIN 5434/43; paratype of Daurosaurus olovus) ischium (Alifanov and Saveliev, 2014a,b)
(PIN 5434/56; 5435/56 in Alifanov et al., 2014) humerus, radius, feathers (Alifanov and Saveliev, 2014a,b) or distal caudal vertebra (Alifanov and Saveliev, 2015)
(PIN 5434/63; paratype of Lepidocheirosaurus natatilis) distal caudal vertebra (Alifanov and Saveliev, 2015)
(PIN 5434/64; paratype of Lepidocheirosaurus natatilis) distal caudal vertebra (Alifanov and Saveliev, 2015)
(PIN 5434/65; paratype of Lepidocheirosaurus natatilis) distal caudal vertebra (Alifanov and Saveliev, 2015)
(PIN 5434/67; paratype of Lepidocheirosaurus natatilis) distal caudal vertebra (Alifanov and Saveliev, 2015)
(PIN 5435/1; holotype of Lepidocheirosaurus natatilis) incomplete pedal phalanx II-1, phalanx II-2 (18 mm), partial pedal ungual II, incomplete phalanx III-1, partial phalanx III-2 (~15 mm), fragmentary phalanx III-3, incomplete phalanx IV-1, phalanx IV-2 (12.5 mm), distal phalanx IV-3, phalanx IV-4 (13.5 mm), pedal ungual IV, pedal claw sheath, scales, feathers (Alifanov, 2014; described by Alifanov and Saveliev, 2015)
(PIN 5435/12; paratype of Lepidocheirosaurus natatilis) caudal vertebrae, scales (Alifanov and Saveliev, 2015)
(PIN 5435/13) feathers (Alifanov et al., 2014)
(PIN 5435/14) scales, feathers (Alifanov et al., 2014)
(PIN 5435/23) distal femur, proximal tibia, proximal fibula, scales, feathers (Alifanov et al., 2014)
(PIN 5435/47) feathers (Alifanov et al., 2014)
(PIN 5435/48) feathers (Alifanov et al., 2014)
(PIN 5435/50; paratype of Lepidocheirosaurus natatilis) caudal vertebrae, scales (Alifanov and Saveliev, 2015)
(PIN 5435/52; paratype of Lepidocheirosaurus natatilis) caudal vertebrae, scales (Alifanov and Saveliev, 2015)
(PIN 5435/54; paratype of Lepidocheirosaurus natatilis) caudal vertebrae, scales (Alifanov and Saveliev, 2015)
(PIN 5435/59) scales, feathers (Alifanov et al., 2014)
(PIN 5435/61) scales, feathers (Alifanov et al., 2014)
(PIN 5435/62; paratype of Lepidocheirosaurus natatilis) caudal vertebrae, scales (Alifanov and Saveliev, 2015)
(PIN 5435/63; paratype of Lepidocheirosaurus natatilis) caudal vertebrae, scales (Alifanov and Saveliev, 2015)
(PIN 5435/64; paratype of Lepidocheirosaurus natatilis) caudal vertebrae, scales (Alifanov and Saveliev, 2015)
Diagnosis- (after Godefroit et al., 2014) maxilla with anterior ascending process much lower than posterior ascending process; maxillary fenestra larger than antorbital fenestra; jugal with notched postorbital ramus; postorbital with dorsoventrally expanded posterior ramus; dorsoventrally slender postacetabular process; deep extensor fossae on metatarsals II-IV.
Comments- Kulindadromeus material was first discovered in 2010 (announced in Alifanov, 2012 and 2014a,b prior to naming of any taxa), with the type material corrected between 2010 and 2012. Two bonebeds are known in the Kulinda locality, each containing hundreds of specimens in various states of articulation. Godefroit et al. (2014) described the material as the basal neornithischian Kulindadromeus on July 25th (submitted March 13th), noting "Each individual skeletal element is represented by a single morphotype, and all of the observed morphological differences can easily be explained by ontogenetic and intraspecific variation, as confirmed by the detailed study of the partial skeletons" and that besides ornithischian material, only a single shed theropod tooth had been found. Alifanov (2014) wrote a preliminary paper proposing the Kulinda material belonged to a hypsilophodontid, a jeholosaurid and a theropod. Alifanov and Saveliev (2014a,b; submitted March 3rd) later described the two ornithischians as the hypsilophodontid Kulindapteryx and the jeholosaurid Daurosaurus, each based on pelvic and/or hindlimb material, with most other material only referred to Hypsilophodontia indet.. Godefroit (online 2014) alleges Alifanov stole INREC material and gave them PIN specimen numbers for his description of the taxa with Saveliev, and indeed INREC K4/115 is the same specimen as PIN 5434/57, while INREC K3/203 and PIN 5434/54 are part and counterpart. Neither Russian nor English version of Alifanov and Saveliev's paper has an established paper publication date (unlike Godefroit et al., who registered Kulindadromeus with ZooBank in their electronic publication), though they were available online by July 3rd and August 6th respectively. Of Godefroit et al.'s diagnostic characters for Kulindadromeus, only Daurosaurus' holotype can be evaluated- the dorsoventrally slender postacetabular process which is indeed present. Thus if the Russian version of Alifanov and Saveliev's paper was physically published prior to July 25th, it could be argued Daurosaurus is a diagnostic senior synonym of Kulindadromeus. Given the alleged ethical issues of Alifanov and Saveliev's paper, the poor quality of Daurosaurus' holotype (ilium, femur and tibia; the latter two of which have morphologies that are mostly "indiscernible" in their own words) compared to Kulindadromeus' (incomplete skull), the far more detailed and modern description in Godefroit's paper, and unestablished physical publication date of Daurosaurus, Kulindadromeus is used for the taxon here.
Lepidocheirosaurus- Alifanov's (2014) reported theropod material is PIN 5435/51 (distal caudal vertebrae with associated scales) and PIN 5435/1 (supposed distal metacarpals, phalanges and scales). While only referred to Theropoda fam. indet. at the time, these were later described as a new basal ornithomimosaur taxon Lepidocheirosaurus by Alifanov and Saveliev (2015), along with other distal caudals. These authors referred it to their new family Nqwebasauridae along with the eponymous genus. The supposed manus is the holotype, but is far more likely to be a pes with the supposed metacarpals I-III being phalanges II/III/IV-1. This is based on the lack of an asymmetrical ginglymus on 'metacarpal I', a short 'phalanx I-1', and short 'phalanx III-3'. These are expected in a pedal phalanx II-1, phalanx II-2 and phalanx IV-4 respectively though. As the only non-Kulindadromeus dinosaur material in the Kulinda beds reported by Godefroit et al. is a single shed theropod tooth, and the pes matches basal ornithischians, Lepidocheirosaurus is probably a junior synonym of Kulindadromeus. As for the referred caudal material, PIN 5435/51 is the counterslab to Kulindadromeus specimen INREC K4/159. Other specimens share the dorsal scales of Kulindadromeus and/or long postzygapophyses which are like basal ornithischians but unlike most theropods. As no other caudal morphologies are known from Kulinda, it would be very unlikely for only theropod caudals to be preserved but only ornithischian crania, pectoral and pelvic elements.
References- Alifanov, 2012. Kulinda, the first Late Jurassic dinosaur locality in Russia. Priroda. 2012(3), 53-54.
Alifanov, 2014a. On the discovery of Late Jurassic dinosaurs in Transbaikalia. Doklady Akademii Nauk. 455(4), 421-423.
Alifanov, 2014b. The discovery of Late Jurassic dinosaurs in Russia. Doklady Earth Sciences. 455(2), 365-367.
Alifanov and Saveliev, 2014a. Two new ornithischian dinosaurs (Hypsilophodontia, Ornithopoda) from the Late Jurassic of Russia. Paleontologicheskii Zhurnal. 48(4), 72-82.
Alifanov and Saveliev, 2014b. Two new ornithischian dinosaurs (Hypsilophodontia, Ornithopoda) from the Late Jurassic of Russia. Paleontological Journal. 48(4), 414-425.
Alifanov, Saveliev, Tereshchenko, Artemov and Seregin, 2014a. Skin structure in ornithischian dinosaurs (Hypsilophodontia, Ornithopoda) from the Late Jurassic of Transbaikalia. Paleontologicheskii Zhurnal. 48(5), 72-80.
Alifanov, Saveliev, Tereshchenko, Artemov and Seregin, 2014b. Integument structure in ornithischian dinosaurs (Hypsilophodontia, Ornithopoda) from the Late Jurassic of Transbaikalia. Paleontological Journal. 48(5), 523-533.
Godefroit, online 2014. http://dml.cmnh.org/2014Jul/msg00028.html
Godefroit, Sinitsa, Dhouailly, Bolotsky, Sizov, McNamara, Benston and Spagna, 2014a. A Jurassic ornithischian dinosaur from Siberia with both feathers and scales. Science. 345(6195), 451-455.
Godefroit, Sinitsa, Dhouailly, Bolotsky, Sizov, McNamara, Benston and Spagna, 2014b. Response to Comment on "A Jurassic ornithischian dinosaur from Siberia with both feathers and scales". Science. 346(6208), 434c.
Lingham-Soliar, 2014. Comment on "A Jurassic ornithischian dinosaur from Siberia with both feathers and scales". Science. 346(5208), 434b.
Saveliev and Alifanov, 2014a. A new type of skin derivatives in ornithischian dinosaurs from the Late Jurassic of Transbaikalia (Russia). Doklady Akademii Nauk. 456(2), 251-253.
Saveliev and Alifanov, 2014b. A new type of skin derivatives in ornithischian dinosaurs from the Late Jurassic of Transbaikalia (Russia). Doklady Biological Sciences. 456, 182-184.
Alifanov and Saveliev, 2015. The most ancient ornithomimosaur (Theropoda, Dinosauria), with cover imprints from the Upper Jurassic of Russia. Paleontological Journal. 49(6), 636-650.

Cerapoda

Ornithopoda

Iguanodontia

Hadrosauridae

unnamed hadrosaurid (Csiki and Grigorescu, 1998)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania

Material- (FGGUB R.351) distal metatarsal
Comments- This was first identified by Grigorescu and Kessler (1981) as a distal femur of Elopteryx, then by Csiki and Grigorescu (1998) as a neoceratosaur distal femur. It was most recently identified as a probably hadrosaurid distal metatarsal based on a matching complete specimen (MAFI Ob.3120a) (Kessler et al., 2005).
References- Grigorescu and Kessler, 1981. A new specimen of Elopteryx nopcsai from the dinosaurian beds of Hateg Basin. Revue Roumaine de Geologie, Geophysique et Geographie, Geologie. 24, 171-175.
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.
Kessler, Grigorescu and Csiki, 2005. Elopteryx revisited - a new bird-like specimen from the Maastrichtian of the Hateg Basin. Acta Palaeontologica Romaniae. 5, 249-258.

unnamed Hadrosauridae (Alifanov and Bolotsky, 2010a)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material
- (AEHM 2/419; paratype of Arkharavia heterocoelica) caudal vertebra (80 mm)
(AEHM 2/420; paratype of Arkharavia heterocoelica) caudal vertebra (85 mm)
(AEHM 2/720; paratype of Arkharavia heterocoelica) proximal caudal vertebra (80 mm)
(AEHM 2/997; paratype of Arkharavia heterocoelica) proximal caudal vertebra (93 mm)
Comments- These were referred to Alifanov and Bolotsky's (2010a, b) new supposed titanosauriform Arkharavia, but, Godefroit et al. (2012) stated the type vertebrae were likely hadrosaurid instead. Mannion et al. (2013) confirmed paratype AEHM 2/720 is hadrosaurid based on neural spine anatomy, and this would also hold true for the figured paratype AEHM 2/997. Another paratype (AEHM 2/420) is said to have the same kind of neural spine, so may also be hadrosaurid, but the final one (AEHM 2/419) is neither figured nor remarked upon, so we have no information to judge it from. Several hadrosaurid taxa are known from the Udurchukan Formation (Amurosaurus, Kerberosaurus, Olorotitan) and the correlated Yuliangze Formation (Charonosaurus, Mandschurosaurus, Sahaliyania, Wulagasaurus), but only a few of these have associated caudals, and none have been distinguished using caudal characters.
References- Alifanov and Bolotsky, 2010a. Arkharavia heterocoelica gen. et sp. nov., a new sauropod dinosaur from the Upper Cretaceous of far earstern Russia. Paleontologicheskii Zhurnal. 2010(1), 76-83.
Alifanov and Bolotsky, 2010b. Arkharavia heterocoelica gen. et sp. nov., a new sauropod dinosaur from the Upper Cretaceous of the far East of Russia. Paleontological Journal. 44(1), 84-91.
Godefroit, Bolotsky and Bolotsky, 2012. Osteology and relationships of Olorotitan arharensis, a hollow-crested hadrosaurid dinosaur from the latest Cretaceous of far eastern Russia. Acta Palaeontologica Polonica. 57(3), 527-560.
Mannion, Upchurch, Barnes and Mateus, 2013. Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of the Linnean Society. 168(1), 98-206.

"Troodon" isfarensis Nessov, 1995
= Saurornithoides isfarensis (Nessov, 1995) Olshevsky, 2000
Early Santonian, Late Cretaceous
Yalovach Formation, Tajikistan

Holotype- (CCMGE 484/12457; lost) prefrontal
Comments- Nessov (1995) identified this as a troodontid frontal, but though they could not locate it, Averianov and Sues (2007) reidentified the specimen as a hadrosaurid prefrontal.
References- 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.
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic Meanderings. 3, 1-157.
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.

Saurischia Seeley, 1887
Definition- (Allosaurus fragilis <- Stegosaurus armatus) (modified from Kischlat, 2000)
Other definitions- (Passer domesticus <- Triceratops horridus) (modified from Padian, 1997; modified from Gauthier, 1986)
(Megalosaurus bucklandii, Plateosaurus engelhardti <- Iguanodon bernissartensis) (Clarke et al., 2004)
(Tyrannosaurus rex <- Triceratops horridus) (modified from Holtz and Osmolska, 2004)
(Passer domesticus, Saltasaurus loricatus <- Triceratops horridus) (Sereno, 2005)
= Opisthocoelia Owen, 1860
= Pachypodosauria Huene, 1914
= Saurischia sensu Gauthier, 1986
Definition- (Passer domesticus <- Triceratops horridus) (modified)
= Saurischia sensu Clark et al., 2004
Definition- (Megalosaurus bucklandii, Plateosaurus engelhardti <- Iguanodon bernissartensis)
= Saurischia sensu Holtz and Osmolska, 2004
Definition- (Tyrannosaurus rex <- Triceratops horridus)
= Saurischia sensu Sereno, 2005
Definition- (Passer domesticus, Saltasaurus loricatus <- Triceratops horridus)
Comments- The most common definition includes birds as a specifier, but birds were not originally viewed as saurischians. Indeed, they were not accepted as such until the 1980's. Thus, a definition not necessitating their inclusion is preferred here. Kischlat's definition uses taxa which Seeley used to exemplify Saurischia and Ornithischia, so is accepted over Clarke et al.'s definition using taxa with historical priority and Holtz and Osmolska's less objective choices. Clarke et al.'s and Sereno's definitions do have the advantage of ensuring both sauropodomorphs and theropods are included however.
Owen (1860) named Opisthocoelia for a crocodylian group containing "Cetiosaurus" medius, Pelorosaurus and Streptospondylus. Huene (1914) erected Pachypodosauria for carnosaurs and sauropodomorphs, to the exclusion of coelurosaurs.
References- Owen, 1860. Palaeontology, or a Systematic Summary of Extinct Animals and their Geological Relations. Second Edition. Adam and Charles Black, Edinburgh. 463 pp.
Huene, 1914. Beiträge zur geschichte der Archosaurier [Contribution to the history of the archosaurs]. Geologie und Paläontologie Abhandlungen. 13(7), 1-56.

Saurischia incertae sedis

Alwalkeria Chatterjee and Creisler, 1993
= Walkeria Chatterjee, 1987
A. maleriensis (Chatterjee, 1987) Chatterjee and Creisler, 1993
= Walkeria maleriensis Chatterjee, 1987
Carnian, Late Triassic
Maleri Formation, India
Holotype
- (ISI R 306 in part) (?)dorsal vertebra (~20 mm), incomplete femur (122 mm), astragalus (25 mm wide, 13 mm tall)
Diagnosis- excavation of bases of dorsal neural arches; highly expanded femoral head; very prominant fourth trochanter.
Comments- Originally referred to Podokesauridae (Chatterjee, 1987), Alwalkeria was later assigned to Herrerasauridae by Paul (1988) based on unspecified femoral similarities, who viewed it as a link between traditional herrerasaurids and Protoavis. This latter link was due to the unserrated teeth though, which are no longer thought to be part of the holotype. Novas (1989) found the astragalus exhibited characters suggesting it was a Herrerasaurus-grade dinosaur, outside Avepoda. Norman (1990) viewed it as a theropod with no particularily close ties to coelophysoids. Rauhut and Remes (2005) find Alwalkeria to be chimaerical, with the anterior skull referrable to a crurotarsan, perhaps an ornithosuchid. This agrees with Rauhut's (2003) comments that the long mandibular symphysis is present in members of Crurotarsi. The partial cervical vertebra is said to resemble prolacertiforms, while the dorsal centra are highly variable in size and lamina development, suggesting they may belong to multiple taxa. The femur and astragalus are dinosaurian however, with the latter possessing saurischian characters. Rauhut and Remes state the femur lacks characters of sauropodomorphs or avepods though, suggesting a herrerasaurid- or Eoraptor-grade saurischian.
References- Chatterjee, 1987. A new theropod dinosaur from India with remarks on the Gondwana-Laurasia connection in the Late Triassic. Geophysical Monograph. 41, 183-189.
Paul, 1988. Predatory dinosaurs of the world. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". in Weishampel, et al. (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. p. 280-305.
Novas, 1989. The tibia and tarsus in Herrerasauridae (Dinosauria, incertae sedis) and the origin and evolution of the dinosaurian tarsus. Journal of Paleontology. 63, 677-690.
Chatterjee and Creisler, 1994. Alwalkeria (Theropoda) and Morturneria (Plesiosauria), new names for preoccupied Walkeria Chatterjee, 1987 and Turneria Chatterjee and Small, 1989. Journal of Vertebrate Paleontology. 14(1), 142.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Remes and Rauhut, 2005. The oldest Indian dinosaur Alwalkeria maleriensis Chatterjee revised: a chimera including remains of a basal saurischian. in Kellner, Henriques and Rodrigues (eds). II Congresso Latino-Americano de Paleontologia de Vertebrados, Boletim de Resumos. Museu Nacional, Rio de Janeiro. 218.
Novas, Ezcurra, Chatterjee and Kutty, 2011. New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram Formations of central India. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 333-349.

Herrerasauria Galton, 1985
Definition- (Herrerasaurus ischigualastensis <- Liliensternus liliensterni, Plateosaurus engelhardti) (modified from Langer, 2004)
= Staurikosauria Paul, 1988
= Herreravia Paul, 1988
= Herrerasauridae sensu Sereno, 1998
Definition- (Herrerasaurus ischigualastensis <- Passer domesticus) (modified)
Comments- Galton (1985) erected Herrerasauria to contain Staurikosaurus, Herrerasaurus and Aliwalia, within Theropoda. Paul (1988) later named Herreravia to contain Herrerasaurus, Frenguellisaurus, Aliwalia, Alwalkeria and possibly Protoavis. Both terms were seldomly used, as Herrerasauridae was soon defined to contain both Staurikosaurus and Herrerasaurus (Novas, 1992), making Herrerasauria redundant. Aliwalia is now known to be a junior synonym of the sauropodomorph Eucnemesaurus (Yates, 2007). Alwalkeria and Protoavis are both chimaeras whose theropod material seems more closely related to birds than Herrerasaurus (Remes and Rauhut, 2005; Nesbitt et al., 2007) and Frenguellisaurus is a synonym of Herrerasaurus itself (Novas, 1993). Langer (2004) recently reinstated Herrerasauria to include herrerasaurids and taxa more closely related to them than to sauropodomorphs and avepods. He suggested Chindesaurus may be a non-herrerasaurid herrerasaurian, but this has yet to be validated by a published phylogenetic analysis. In fact, that of Yates (2006) found Chindesaurus to be closer to Avepoda than Herrerasauridae. However, the unpublished analysis of Bittencourt and Kellner (2004) agrees with Langer. Hunt (1996) proposed Chindesaurus, Gojirasaurus (as "Revueltoraptor") and "Comanchesaurus" belonged to a herrerasaur clade separate from Staurikosaurus and Herrerasaurus, which would make them non-herrerasaurid herreravians under Langer's definition. The latter two taxa are now considered to be coelophysoids (Nesbitt et al., 2007). Novas et al. (2009) said ISI R282 resembles herrerasaurs, but Novas et al. (2011) later found it to emerge as a dinosaur outside Eoraptor+Chindesaurus+Tawa+Avepoda and Guaibasauridae+more derived sauropodomorphs when entered into Yates' sauropodomorph matrix.
References- Galton, 1985. The poposaurid thecodontian Teratosaurus suevicus v. Meyer, plus referred specimens mostly based on prosauropod dinosaurs, from the Middle Stubensandstein (Upper Triassic) of Nordwurttemberg. Stuttgart Beitrage zur Naturkunde (B). 116, 1-29.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster, New York. 464 pp.
Novas, 1992. Phylogenetic relationships of the basal dinosaurs, the Herrerasauridae. Palaeontology. 35, 51-62.
Novas, 1993. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina. Journal of Vertebrate Paleontology. 13, 400-423.
Hunt, 1996. A new clade of herrerasaur-like theropods from the Late Triassic of western North America. Journal of Vertebrate Paleontology 16(3), 43A.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 210(1), 41-83.
Bittencourt and Kellner, 2004. The phylogenetic position of Staurikosaurus pricei Colbert, 1970 from the Triassic of Brazil. Journal of Vertebrate Paleontology. 24(3).
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Remes and Rauhut, 2005. The oldest Indian dinosaur Alwalkeria maleriensis Chatterjee revised: a chimera including remains of a basal saurischian. in Kellner, Henriques and Rodrigues (eds). II Congresso Latino-Americano de Paleontologia de Vertebrados, Boletim de Resumos.Museu Nacional, Rio de Janeiro. 218.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.
Yates, 2007. Solving a dinosaurian puzzle: the identity of Aliwalia rex Galton. Historical Biology. 19(1), 93-123.
Novas, Chatterjee, Ezcurra and Kutty, 2009. New dinosaur remains from the Late Triassic of Central India. Journal of Vertebrate Paleontology. 29(3), 156A.
Novas, Ezcurra, Chatterjee and Kutty, 2011. New dinosaur species from the Upper Triassic Upper Maleri and Lower Dharmaram formations of Central India. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 101, 333-349.

Herrerasauridae Benedetto, 1973
= Staurikosauridae Galton, 1977
Definition- (Herrerasaurus ischigualastensis + Staurikosaurus pricei) (modified from Novas, 1992)
Other definitions- (Herrerasaurus ischigualastensis <- Passer domesticus) (Sereno, 2005; modified from Sereno, 1998)
Diagnosis- (after Novas, 1992; Langer, 2004) shortened posterior dorsal centra (dorsals 13-15 with length 80% or less of height); posterior dorsal and first sacral vertebrae with robust, squared neural spines; sacral ribs very deep and cover nearly 90% of the ilium; vertical neural spines on proximal caudals; elongate distal caudal prezygopophyses; brevis shelf and fossa reduced; reduced lesser trochanter.
(after Ezcurra and Novas, 2007) distal carpal V enlarged.
Comments- After being referred to Sauropodomorpha (Van Heerden, 1978; 1979; Cooper, 1981), herrerasaurids were generally excluded from Dinosauria (Gauthier, 1984, 1986; Brinkman and Sues, 1987; Paul, 1988; Novas, 1989; Benton, 1990; Sereno and Novas, 1990; Novas, 1992; also Langer et al., 1999 and Fraser et al., 2002) until Herrerasaurus was redescribed (Sereno and Novas, 1992; Novas, 1993; Sereno, 1993; Sereno and Novas, 1993) as a theropod. Additional studies placed Herrerasauridae in Theropoda (Sereno et al., 1993; Novas, 1996, 1997; Benton, 1999; Sereno, 1999; Kischlat, 2000; Rauhut, 2003; Tykoski, 2005; Ezcurra and Novas, 2007), or as saurischians outside the sauropodomorph-theropod clade (Eusaurischia) (Padian and May, 1993; Bonaparte and Pumares, 1995; Holtz, 1995; Langer et al., 1999; Langer, 2004; Ezcurra, 2006; Langer and Benton, 2006; Yates, 2007; Irmis et al., 2007; Smith et al., 2007; Bittencourt Rodrigues, 2010; Ezcurra, 2010). A position inside Theropoda is tentatively followed here, as it is more parsimonious when all the data of Rauhut (2003), Tykoski (2005), Ezcurra and Novas (2007), Langer and Benton (2006) and Smith et al. (2007) is combined, but only by four steps.
Benedetto (1973) erected Herrerasauridae to include Herrerasaurus and Staurikosaurus, though Staurikosaurus was generally excluded until Novas' 1992 study. Before then, Herrerasauridae was viewed as monotypic by most sources except for Paul (1988).
Not herrerasaurids- Paul (1988) included both Aliwalia and Alwalkeria in Herrerasauridae and considered Protoavis a possible herrerasaurid derivative. Aliwalia is now known to be a junior synonym of the sauropodomorph Eucnemesaurus (Yates, 2007). Alwalkeria and Protoavis are both chimaeras whose theropod material seems more closely related to birds than Herrerasaurus (Remes and Rauhut, 2005; Nesbitt et al., 2007). He also considered SMNS 51958, a proximal femur from the Lowenstein Formation of Germany, a herrerasaurid. While this may be true, it shows no particular similarity to herrerasaurid femora compared to those of other theropods. Novas (1989) referred part of Trialestes paratype PVL 2559 to Herrerasauridae indet., but this was retained in Trialestes by Clark et al. (2000).
Kirby (1993) referred to cf. Staurikosauridae? from the Owl Rock Member of the Chinle Formation of Arizona, but this was based on two limb bone fragments (MNA V6729) which Spielmann et al. (2007) could not identify past Archosauromorpha indet.. Long and Murry (1995) referred Chindesaurus to Herrerasauridae, which seems unlikely based on Langer (2004) and Yates (2006). Hunt (1994) referred Gojirasaurus (as "Revueltoraptor") and "Comanchesaurus" to a broader version of Herrerasauridae in his unpublished thesis. These are called herrerasaurids A and B by Hunt et al. (1998), but are coelophysoids (Nesbitt et al., 2007). Herrerasaurid C of Hunt et al. (1998) is based on dorsal and caudal centra from the Bull Canyon Formation (NMMNH P-16656) which were assigned to Archosauria indet. by Nesbitt et al (2007). Dzik (2001) identified a second sacral vertebra (ZPAL AbIII/284) from the Late Triassic of Poland as a possible herrerasaurid, but this was later referred to the silesaurid Silesaurus opolensis (Dzik, 2003). Nesbitt (2001) tentatively assigned a distal pubis from the Middle Triassic Moekopi Formation of Arizona to Herrerasauridae, but more recently (pers. comm., 2010) states its morphology is equivocal between poposauroids and theropods.
References- Colbert, 1970. A saurischian dinosaur from the Triassic of Brazil. American Museum Novitates. 2405, 1-39.
Benedetto, 1973. Herrerasauridae, nueva familia de saurisquios triasicos. Ameghiniana. 10(1), 89-102.
Galton, 1977. On Staurikosaurus pricei, an early saurischian dinosaur from the Triassic of Brazil, with notes on the Herrerasauridae and Poposauridae. Palaontologische Zeitschrift. 51, 234-245.
van Heerden, 1978. Herrerasaurus and the origin of sauropod dinosaurs. South African Journal of Science. 74, 187-189.
van Heerden, 1979. The morphology and taxonomy of Euskelosaurus (Reptilia: Saurischia; Late Triassic) from South Africa. Navorsinge van die Nasionale Museum, Bloemfontein. 4, 21-84.
Cooper, 1981. The prosauropod dinosaur Massospondylus carinatus Owen from Zimbabwe: Its biology, mode of life and phylogenetic significance. Occasional Papers of the National Museums and Monuments of Rhodesia (series B, Natural Sciences). 6, 689-840.
Gauthier, 1984. A cladistic analysis of the higher systematic categories of the Diapsida. PhD thesis. University of California, Berkeley. 564 pp.
Gauthier, 1986. Saurischian monophyly and the origin of birds. Memoirs of the Californian Academy of Sciences. 8, 1-55.
Brinkman and Sues, 1987. A staurikosaurid dinosaur from the Upper Triassic Ischigualasto Formation of Argentina and the relationships of the Staurikosauridae. Palaeontology. 30, 493-503.
Paul, 1988. Predatory dinosaurs of the world. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Novas, 1989. The tibia and tarsus in Herrerasauridae (Dinosauria, incertae sedis) and the origin and evolution of the dinosaurian tarsus. Journal of Paleontology. 63, 677-690.
Benton, 1990. Origin and interrelationships of dinosaurs. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press: Berkeley. 11-30.
Sereno and Novas, 1990. Dinosaur origins and the phylogenetic position of pterosaurs. Journal of Vertebrate Paleontology. 10(3), 42A.
Novas, 1992. Phylogenetic relationships of the basal dinosaurs, the Herrerasauridae. Palaeontology. 35, 51-62.
Sereno and Novas, 1992. The complete skull and skeleton of an early dinosaur. Science. 258, 1137-1140.
Kirby, 1993. Relationships of Late Triassic basin evolution and faunal replacement in the southwestern United States: Perspectives from the upper part of the Chinle Formation in northern Arizona. In Lucas and Morales (eds.). The Nonmarine Triassic. New Mexico Museum of Natural History and Science Bulletin. 3, 233-242.
Novas, 1993. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina. Journal of Vertebrate Paleontology. 13, 400-423.
Padian and May, 1993. The earliest dinosaurs. Bulletin of the New Mexico Museum of Natural History and Science. 3, 379-381.
Sereno, 1993. The pectoral girdle and forelimb of the basal theropod Herrerasaurus ischigualastensis. Journal of Vertebrate Paleontology. 13, 425-450.
Sereno and Novas, 1993. The skull and neck of the basal theropod Herrerasaurus ischigualastensis. Journal of Vertebrate Paleontology. 13, 451-476.
Sereno, Forster, Rogers and Monetta, 1993. Primitive dinosaur skeleton from Argentina and the early evolution of Dinosauria. Nature. 361, 64-66.
Hunt, 1994.
Bonaparte and Pumares, 1995. Notas sobre el primer craneo de Riojasaurus incertus (Dinosauria, Prosauropoda, Melanorosauridae) del Triasico Superior de La Rioja, Argentina. Ameghiniana. 32, 341-349.
Holtz, 1995. A new phylogeny of the Theropoda. Journal of Vertebrate Paleontology. 15(3), 35A.
Novas, 1996. Dinosaur monophyly. Journal of Vertebrate Paleontology. 16, 723-741.
Novas, 1997. Herrerasauridae. in Currie and Padian (eds). Encyclopedia of Dinosaurs. Academic Press, San Diego, California/London, UK. 303-311.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic Dinosaurs from the Western United States. Geobios 31, 4: 511-531.
Langer, Abdala, Richter and Benton, 1999. A sauropodomorph dinosaur from the Upper Triassic (Carnian) of souther Brazil. Comptes Rendus de l’Academie des Sciences, Paris. 329, 511-517.
Benton, 1999. Scleromochlus taylori and the origin of dinosaurs and pterosaurs. Philosophical Transactions of the Royal Society of London (Series B). 354, 1423-1446.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Clark, Sues and Berman, 2000. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology. 20(4), 683-704.
Heckert, Lucas and Sullivan, 2000. Triassic dinosaurs in New Mexico. in Lucas and Heckert (eds). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 17-26.
Kischlat, 2000. Tecodoncios: A aurora dos Arcosaurios no Triassico. in Holz and De Rose (eds.). Paleontologia do Rio Grande do Sol. 273-316.
Dzik, 2001. A new Paleorhinus fauna in the Early Late Triassic of Poland. Journal of Vertebrate Paleontology. 21(3), 625-627.
Nesbitt, 2001. New fossil vertebrate material from the Holbrook Member, Moenkopi Formation (Middle Triassic) from Northern Arizona. Journal of Vertebrate Paleontology. 21(3), 83A.
Fraser, Padian, Walkeden and Davis, 2002. Basal dinosauriform remains from Britain and the diagnosis of the Dinosauria. Palaeontology. 45, 78-95.
Dzik, 2003. A beaked herbivorous archosaur with dinosaur affinities from the Early Late Triassic of Poland. Journal of Vertebrate Paleontology. 23(3), 556-574.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Remes and Rauhut, 2005. The oldest Indian dinosaur Alwalkeria maleriensis Chatterjee revised: a chimera including remains of a basal saurischian. in Kellner, Henriques and Rodrigues (eds). II Congresso Latino-Americano de Paleontologia de Vertebrados, Boletim de Resumos.Museu Nacional, Rio de Janeiro. 218.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas. 28(4),649-684.
Langer and Benton, 2006. Early dinosaurs: A phylogenetic study. Journal of Systematic Palaeontology. 4(4), 309-358.
Ezcurra and Novas, 2007. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science. 317, 358-361.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Spielmann, Lucas and Heckert, 2007. Tetrapod fauna of the Upper Triassic (Revueltian) Owl Rock Formation, Chinle Group, Arizona. In Lucas and Spielmann (eds.). The Global Triassic, New Mexico Museum of Natural History and Science Bulletin. 41, 371-383.
Yates, 2007. Solving a dinosaurian puzzle: the identity of Aliwalia rex Galton. Historical Biology. 19(1), 93-123.
Bittencourt Rodrigues, 2010. Revisao filogenetica dos dinossauriformes basais: Implicacoes para a origem dod dinossauros. Unpublished Doctoral Thesis. Universidade de Sao Paulo. 288 pp.
Ezcurra, 2010. A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: A reassessment of dinosaur origin and phylogeny. Journal of Systematic Palaeontology. 8(3), 371-425.

undescribed possible herrerasaurid (Heckert, 1999)
Late Carnian, Late Triassic
Mesa Redondo Member of the Chinle Formation, New Mexico, US

Material- (NMMNH P-18391) proximal tibia
Comments- This was identified as a theropod by Heckert (1999), and was tentatively assigned to Herrerasauridae by Heckert et al. (2000).
References- Heckert. 1999. Upper Triassic tetrapods from the Lucero Uplift, central New Mexico. New Mexico Geological Society Guidebook, 50th Field Conference, Albuquerque 50:311-315
Heckert, Lucas and Sullivan, 2000. Triassic dinosaurs in New Mexico. in Lucas and Heckert (eds). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 17-26.

undescribed possible herrerasaurid (Kischlat and Barberena, 1999)
Late Norian, Late Triassic
Caturrita Formation, Brazil

Material- (FZB coll.) dorsal vertebra
....(UFPel coll.) sacral vertebrae
....(UFRGS coll.) pubis, ischium
Comments- Stated by Kischlat and Barberena to be from the Botucarai outcrop of the Santa Maria Formation, this is actually part of the younger Caturrita Formation.
Kischlat and Barberena imply this specimen is a herrerasaurid, as they compare it to both Herrerasaurus and Spondylosoma (which they treat as a herrerasaurid, though it is recognized here as a pseudosuchian). The pronounced pubic tubercle is said to be similar to Herrerasaurus and unlike the double tubercle of Spondylosoma, but the sacral vertebrae are stated to be similar to the third in Spondylosoma. As no herrerasaurid synapomorphies were mentioned and Spondylosoma is not referred to that family here, the specimen under consideration may belong to another kind of archosaur instead.
Reference- Kischlat and Barberena, 1999. Brazilian dinosaurs: New data. Paleontologia em Destaque, Boletim Informativo da Sociedade Brasileira de Paleontologia. 14(26), 56.

undescribed possible herrerasaurid (Ezcurra and Novas, 2007)
Late Carnian-Early Norian, Late Triassic
Cancha de Bochas Member of the Ischigualasto Formation, San Juan, Argentina

Material- (MACN-PV 18.649a) (small) vertebrae, distal ulna, carpus, manus, pedal phalanges
Diagnosis- (after Ezcurra and Novas, 2007) articular surfaces of ulna and ulnare subequal in size; manual unguals with a posteriorly bifurcated lateral groove.
(after Ezcurra, 2010) manual phalanx II-1 with conspicuous longitudinal ridge on its proximolateral border.
Comments- This specimen was discovered in the 1960s, but not noted until an abstract by Ezcurra and Novas (2007). That paper and Ezcurra (2010) placed it in Herrerasauridae, the latter using a version of Yates' sauropodomorph analysis. It was a herrerasaur based on- distal carpal V enlarged; manual phalanx I-1 longer than metacarpal I; strongly curved manual unguals; metacarpals IV-V ventral to the others. While these are shared with Herrerasaurus, the unknown forelimb of Staurikosaurus makes the relationship between these three taxa uncertain. Problematically, the recently described basal theropod Eodromaeus also has all four characters, and is also small and from the same formation. Indeed, MACN-PV 18.649a may be Eodromaeus, as the latter has a marked proximolateral projection on II-1, but whether this is a ridge or not is not described. The new taxon will be described in more detail by Ezcurra and Novas (in prep.).
References- Ezcurra and Novas, 2007. New dinosaur remains (Saurischia: Herrerasauridae) from the Ischigualasto Formation (Carnian) of NW Argentina. Ameghiniana. 44, 17R.
Ezcurra and Novas, 2008. A review of the dinosaur diversity of the Ischigualasto Formation (Carnian, NW Argentina): Insights on early dinosaur evolution. in Langer, Bittencourt and Castro (eds.). Boletim de Resumos, VI Simposio Brasileiro de Paleontologia de Vertebrados, Paleontologia, Edicao especial. Universidad de Sao Pablo: Ribeirao Preto. 88-89.
Ezcurra, 2010. A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: A reassessment of dinosaur origin and phylogeny. Journal of Systematic Palaeontology. 8(3), 371-425.
Ezcurra and Novas, in prep.

Staurikosaurus Colbert 1970
S. pricei Colbert 1970
Late Carnian-Early Norian, Late Triassic
Alemoa Member of Santa Maria Formation, Brazil
Diagnosis
- (after Novas, 1993) distal bevel on anterior margin of pubis.
(after Langer and Benton, 2006) anterior trochanter reduced to a scar.
(after Bittencourt and Kellner, 2009) distal end of tibia subcircular in distal view; dorsoventrally oriented sulcus on medial surface of proximal third of fibula.
Holotype- (MCZ 1669) (~2.25 m, 19 kg) mandibles (215 mm), fourth cervical vertebra (~33.4 mm), partial fifth cervical vertebra, incomplete sixth cervical vertebra, incomplete seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, first dorsal vertebra, fragmentary second dorsal vertebra, third dorsal vertebra, fourth dorsal vertebra, fifth dorsal vertebra, partial sixth dorsal vertebra (~31.9 mm), seventh dorsal centrum, partial eighth dorsal centrum, ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal vertebra, thirteenth dorsal vertebra, fourteenth dorsal vertebra, fifteenth dorsal neural spine, partial dorsal ribs, first sacral vertebra, second sacral vertebra (29 mm), incomplete third sacral vertebra, partial first caudal vertebra (28.5 mm), second caudal vertebra (24 mm), incomplete third caudal vertebra (22.5 mm), fifth caudal vertebra (23.5 mm), seventh caudal vertebra (24 mm), fourteenth caudal vertebra (25.5 mm), fifteenth caudal vertebra (31.5 mm), sixteenth caudal vertebra (28 mm), seventeenth caudal vertebra (28 mm), eighteenth caudal vertebra (28.5 mm), nineteenth caudal vertebra (29 mm), twenty-fourth caudal vertebra (29 mm), twenty-fifth caudal vertebra (29 mm), twenty-ninth caudal vertebra (30.5 mm), thirtieth caudal vertebra (30.5 mm), thirty-first caudal vertebra (30 mm), thirty-second caudal vertebra (26.5 mm), thirty-third caudal vertebra (29 mm), thirty-fourth caudal vertebra (28.5 mm), thirty-fifth caudal vertebra (28 mm), thirty-sixth caudal vertebra (27 mm), thirty-seventh caudal vertebra (27 mm), thirty-eighth caudal vertebra (22.5 mm), thirty-ninth caudal vertebra (22 mm), fortieth caudal vertebra (23.5 mm), forty-first caudal vertebra (23 mm), forty-second caudal vertebra (20 mm), forty-third caudal vertebra (20 mm), forty-fourth caudal vertebra (18.5 mm), forty-fifth caudal vertebra (21.5 mm), forty-sixth caudal vertebra (15.5 mm), forty-seventh caudal vertebra (16 mm), distal scapula, ilia (one incomplete; 98 mm), pubes (171 mm), incomplete ischia, femora (229 mm), tibiae (246 mm), fragments
Comments- Staurikosaurus' holotype was discovered in 1936, but not described until 1970. The scapula identified by Novas (1992) is an ischium (Bittencourt and Kellner, 2009), and the proximal humerus identified by Colbert (1970) is unidentifiable (Galton, 1977).
Staurikosaurus was originally assigned to Palaeosauriscidae (Colbert, 1970), a defunct family based largely on Efraasia. Van Heerden (1978; 1979) agreed. Galton (1973) suggested Staurikosaurus may be a theropod ancestor within Saurischia. He later (1977) erected the family Staurikosauridae for the genus, where it was placed by Brinkman and Sues (1987) and Paul (1988). These latter authors and Sues (1990) all viewed it as being a dinosauriform less closely related to dinosaurs than Herrerasaurus. Langer and Benton (2006) note the characters used to support this topology are invalid, though they find the poorly developed anterior trochanter could support it. Benedetto (1973) and Galton (1985) were the first to recognize Staurikosaurus and Herrerasaurus were more closely related to each other than to sauropodomorphs or avepods, placing them both in the Herrerasauridae and Herrerasauria respectively. This was confirmed by Novas (1992), and nearly every subsequent cladistic analysis agrees (e.g. Rauhut, 2003; Langer and Benton, 2006; Yates, 2007). One exception is Kischlat and Barbarena (1999) and Kischlat (2000), who find Staurikosaurus to be a basal sauropodomorph based on an unpublished phylogenetic analysis. This was due to the possible caudosacral vertebra, the lack of a trochanteric shelf, and a dorsolateral trochanter.
References- Colbert, 1970. A saurischian dinosaur from the Triassic of Brazil. American Museum Novitates. 2405, 1-39.
Benedetto, 1973. Herrerasauridae, nueva familia de saurisquios triasicos. Ameghiniana. 10(1), 89-102.
Galton, 1973. On the anatomy and relationships of Efraasia diagnostica (Huene) n. gen., a prosauropod dinosaur (Reptilia: Saurischia) from the Upper Triassic of Germany. Paläontologische Zeitschrift. 47(3/4), 229-255.
Galton, 1977. On Staurikosaurus pricei, an early saurischian dinosaur from the Triassic of Brazil, with notes on the Herrerasauridae and Poposauridae. Palaontologische Zeitschrift. 51, 234-245.
van Heerden, 1978. Herrerasaurus and the origin of sauropod dinosaurs. South African Journal of Science. 74, 187-189.
van Heerden, 1979. The morphology and taxonomy of Euskelosaurus (Reptilia: Saurischia; Late Triassic) from South Africa. Navorsinge van die Nasionale Museum, Bloemfontein. 4, 21-84.
Galton, 1985. The poposaurid thecodontian Teratosaurus suevicus v. Meyer, plus referred specimens mostly based on prosauropod dinosaurs, from the Middle Stubensandstein (Upper Triassic) of Nordwurttemberg. Stuttgart Beitrage zur Naturkunde (B). 116, 1-29.
Brinkman and Sues, 1987. A staurikosaurid dinosaur from the Upper Triassic Ischigualasto Formation of Argentina and the relationships of the Staurikosauridae. Palaeontology. 30, 493-503.
Paul, 1988. Predatory dinosaurs of the world. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Benton, 1990. Origin and interrelationships of dinosaurs. in Weishampel, et al. (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 11-30.
Sues, 1990. Staurikosaurus and Herrerasauridae. in Weishampel, et al. (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 143-147.
Novas, 1992. Phylogenetic relationships of the basal dinosaurs, the Herrerasauridae. Palaeontology. 35, 51-62.
Novas, 1993. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina. Journal of Vertebrate Paleontology 13 p. 400-423.
Kischlat and Barbarena, 1999. Brazilian dinosaurs: new data. Paleontologia em Destaque, Boletim Informativo da Sociedade Brasileira de Paleontologia. 14(26).
Galton, 2000, Are Spondylosoma and Staurikosaurus (Santa Maria Formation, Middle-Upper Triassic, Brasil) the oldest saurischian dinosaurs? Palaontologische Zeitschrift. 74(3), 393-423.
Kischlat, 2000. Tecodoncios: A aurora dos Arcosaurios no Triassico. in Holz and De Rose (eds.). Paleontologia do Rio Grande do Sol. 273-316.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Bittencourt and Kellner, 2004. The phylogenetic position of Staurikosaurus pricei Colbert, 1970 from the Triassic of Brazil. Journal of Vertebrate Paleontology. 24(3), 23A-24A.
Bittencourt, 2004. Revisao descritiva e posicionamento filogenetico de Staurikosaurus pricei Colbert 1970 (Dinosauria, Theropoda). Masters thesis, Universidade Federal do Rio de Janeiro.
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Grillo and Azevedo, 2005. Reconstruction of the skeleton of Staurikosaurus pricei Colbert, 1970 (Dinosauria, Theropoda): Use of 3D laser scanner and three-dimensional virtual modeling to reconstruct fossil vertebrates. Journal of Vertebrate Paleontology. 25(3), 66A.
Grillo and Azevedo, 2006. Pelvic and hind limb musculature of Staurikosaurus pricei Colbert, 1970 (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. 26(3), 70A.
Langer and Benton, 2006. Early dinosaurs: A phylogenetic study. Journal of Systematic Palaeontology. 4(4), 309-358.
Yates, 2007. Solving a dinosaurian puzzle: the identity of Aliwalia rex Galton. Historical Biology. 19(1), 93-123.
Bittencourt and Kellner, 2009. The anatomy and phylogenetic position of the Triassic dinosaur Staurikosaurus pricei Colbert, 1970. Zootaxa. 2079, 1-56.
Grillo and Azevedo, 2011a. Recovering missing data: Estimating position and size of caudal vertebrae in Staurikosaurus pricei Colbert, 1970. Anais da Academia Brasileira de Ciências. 83(1), 61-71.
Grillo and Azevedo, 2011b. Pelvic and hind limb musculature of Staurikosaurus pricei (Dinosauria: Saurischia). Anais da Academia Brasileira de Ciências. 83(1), 73-98.

"Herrerasaurinae" Benedetto, 1973 sensu Novas, 1989
Diagnosis- (proposed) narrow, U-shaped antorbital fossa (unknown in Staurikosaurus; also in Tawa); spine tables on posterior dorsal and first sacral vertebrae (also in Eoraptor); prominent acromion on scapula (unknown in Staurikosaurus); pubis with sinuous lateral margin in anterior view; longitudinal keel on anterior edge of proximal femur; anterolateral subcircular muscle scar on distal femur.
Comments- Novas (1989) proposed Herrerasaurinae in his unpublished thesis for a herrerasaurid subfamily containing Herrerasaurus, Ischisaurus and Frenguellisaurus but not Staurikosaurus. As Sanjuansaurus shares several characters with Herrerasaurus to the exclusion of Staurikosaurus, the subfamily may prove useful again.
References- Benedetto, 1973. Herrerasauridae, nueva familia de saurisquios triasicos. Ameghiniana. 10(1), 89-102.
Novas, 1989. Los dinosaurios carnivoros de la Argentina. PhD thesis. Universidad Nacional de La Plata. 510 pp.

Sanjuansaurus Alcober and Martinez, 2010
S. gordilloi Alcober and Martinez, 2010
Late Carnian-Early Norian, Late Triassic
Cancha de Bochas Member of the Ischigualasto Formation, San Juan, Argentina

Holotype- (PVSJ 605; adult) (skull ~384 mm) incomplete maxilla, axis (37 mm), third cervical vertebra (42.5 mm), fourth cervical vertebra (44 mm), fifth cervical vertebra (46.1 mm), sixth cervical vertebra (47.5 mm), seventh cervical vertebra (45.6 mm), eighth cervical vertebra (38.5 mm), ninth cervical vertebra (38 mm), first dorsal vertebra (33.6 mm), second dorsal vertebra (28.3 mm), posterior fifth dorsal vertebra, sixth dorsal vertebra (36.8 mm), seventh dorsal vertebra (38 mm), eighth dorsal vertebra (38.8 mm), ninth dorsal vertebra (36.6 mm), tenth dorsal vertebra (37.7 mm), eleventh dorsal vertebra (37.5 mm), twelfth dorsal vertebra (38 mm), thirteenth dorsal vertebra (36 mm), fourteenth dorsal vertebra (34.5 mm), eighth dorsal rib fragment, first sacral vertebra (38 mm), second sacral vertebra (48 mm), third sacral vertebra (46.2 mm), first caudal vertebra (35.8 mm), second caudal vertebra (36.8 mm), third caudal vertebra (35 mm), fourth caudal vertebra (37 mm), fifth caudal vertebra (33.2 mm), sixth caudal vertebra (32.3 mm), seventh caudal vertebra (38.5 mm), eighth caudal vertebra (38.5 mm), ninth caudal vertebra (37.8 mm), tenth caudal vertebra (37.8 mm), eleventh caudal vertebra (36.5 mm), twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra, anterior fifteenth caudal vertebra, incomplete scapulocoracoids (scapula 185 mm), ulna (178.2 mm), incomplete manual ungual III, ilial fragment fused to pubes (260 mm), femora (395 mm), tibiae (260 mm), fibula, astragalocalcaneum (83 mm wide), metatarsal II (147.5 mm)
Diagnosis- (after Alcober and Martinez, 2010) shelf-like, posterolaterally directed transverse processes on the posterior cervical vertebrae; neural spines of the sixth to eighth dorsal vertebrae, at least, bearing acute anterior and posterior processes; everted lateral margins of the glenoid; short pubis (63% of the femoral length); pronounced, rugose scar on the medial surface of the femur at the level of the fourth trochanter.
(suggested) articular surface for ulnare on ulna convex; very short tibia (66% of femoral length).
Comments- This specimen was discovered in 1994 and later mentioned in an abstract (Martinez and Alcober, 2007). Alcober and Martinez (2010) officially described the taxon as a herrerasaurid in a trichotomy with Staurikosaurus and Herrerasaurus using a version of Langer and Benton's matrix.
References- Martinez and Alcober, 2007. Un nuevo herrerasaurido (Dinosauria, Saurischia) de la Formacion Ischigualasto (Triasico Superior, Craniano). Ameghiniana. 44, 28-29R.
Alcober and Martinez, 2010. A new herrerasaurid (Dinosauria, Saurischia) from the Upper Triassic Ischigualasto Formation of northwestern Argentina. ZooKeys. 63, 55-81.

Herrerasaurus Reig, 1963
?= Ischisaurus Reig, 1963
?= Frenguellisaurus Novas, 1986
H. ischigualastensis Reig, 1963
?= Ischisaurus cattoi Reig, 1963
?= Frenguellisaurus ischigualastensis Novas, 1986
Late Carnian-Early Norian, Late Triassic
Cancha de Bochas Member of the Ischigualasto Formation, San Juan, Argentina

Diagnosis- (after Reig, 1963) pubis proximally curved and ventrally oriented; prominent greater trochanter; tibia shortened (87-91% of femur).
(after Novas, 1986) maxilla strongly convex ventrally (also in Eodromaeus).
(after Novas, 1993) premaxilla-maxilla fenestra (unknown in other herrerasaurids); ridge on lateral surface of jugal (unknown in other herrerasaurids); dorsal part of laterotemporal fenestra less than a third as wide as ventral part (unknown in other herrerasaurids); deeply incised supratemporal fossa that extends across medial postorbital process (unknown in other herrerasaurids); subquadrate ventral squamosal process with lateral depression (unknown in other herrerasaurids); quadratojugal overlaps posterodorsal quadrate face (unknown in other herrerasaurids); pterygoid process of quadrate with inturned, trough-shaped ventral margin (unknown in other herrerasaurids); slender ribbed posterodorsal dentary process (unknown in other herrerasaurids); surangular with forked anterior process for articulation with posterodorsal dentary process (unknown in other herrerasaurids); humerus internal tuberosity proximally projected and separated from the humeral head by a deep groove (also in coelophysoids; unknown in other herrerasaurids); humeral entepicondyle ridge-like with anterior and posterior depressions (unknown in other herrerasaurids); enlarged hands (60% of humerus+radius) (unknown in other herrerasaurids); posterior border of ilial peduncle forms right angle with dorsal border of shaft on ischium (unknown in Sanjuansaurus).
(after Sereno and Novas, 1993) broad subnarial process of premaxilla (also in Ornithischia; unknown in other herrerasaurids); broad supratemporal depression (unknown in other herrerasaurids); basal tuber and occipital condyle subequal in width (unknown in other herrerasaurids).
(after Sereno, 1993) circular pit on humeral ectepicondyle (also in Saturnalia; unknown in other herrerasaurids); saddle-shaped ulnar condyle of humerus (unknown in other herrerasaurids); articular surface for ulnare on ulna convex; articular surface of ulnare smaller than that of ulna (unknown in Staurikosaurus and Sanjuansaurus); centrale placed distal to radiale (unknown in other herrerasaurids).
Other diagnoses- Reig (1963) listed numerous characters as being diagnostic of Herrerasaurus. Several refer to the skull of PVL 2558b, which is now excluded from the genus (premaxilla robust and high; three premaxillary teeth; maxilla tall; antorbital fossa widely separated from external nares; eight maxillary teeth; maxillary teeth robust with round alveoli; dentary symphysis dorsoventrally expanded and robust; twelve dentary teeth). The humerus being more than half of femoral length is also based on this specimen. Most of the others are plesiomorphic for saurischians- three sacral vertebrae; short, tall ilium; ischium similar to Plateosaurus and shorter than pubis; gracile and sigmoid femur; astragalus wide and calcaneum small; pes similar to plateosaurids; elongate metatarsus; metatarsal I >66% of metatarsal II; metatarsal V ~66% of metatarsal IV. The preacetabular process is not more robust than in Staurikosaurus. The fourth trochanter is prominent, aliform and as proximally placed in other herrerasaurids. The pubic boot in smaller specimens is only slightly longer than Sanjuansaurus. The pubis is longer than the tibia, not almost as long. The astragalus does not lack an ascending process.
Reig (1963) also listed numerous characters in Ischisaurus' diagnosis. Most of these are plesiomorphic for saurischians- four premaxillary teeth; slender dentary; ~15 dentary teeth; teeth laterally compressed and recurved; relatively short cervical vertebrae; relatively short dorsal vertebrae without ventral keel; short, rounded coracoid; gracile humerus; prominent deltopectoral crest; gracile and sigmoid femur; gracile tibia; astragalus and pes similar to Herrerasaurus. Other characters are vague- premaxillary teeth large; low maxilla; typical olecranon on ulna. The fourth trochanter is as proximally placed in other herrerasaurids, while the "laterally shortened proximal end of tibia" is also present in Sanjuansaurus though not some other Herrerasaurus specimens. The humerus is not a third of humeral length, being 51-59% as in some other basal saurischians. The calcaneum is not larger than the Herrerasaurus holotype. If true, the lack of ventral keels on the cervicals would be distinctive.
Novas (1986) listed a posterodorsal surangular process lateral to the glenoid as an autapomorphy of Ischisaurus, but this is also present in Staurikosaurus.
Novas (1986) listed several characters as apomorphies of Frenguellisaurus. A long and low skull is primitive for dinosauriforms. The anterior maxilla also has enlarged 'canine' teeth in Sanjuansaurus and Tawa. Having a maxilla lower under the antorbital fenestra than Dilophosaurus is plesiomorphic, as is having a smaller antorbital fenestra than that genus. Sanjuansaurus and Eoraptor both have rounder anterior margins on their antorbital fenestrae than Dilophosaurus. Tawa also has a tall jugal, which is also possibly primitive for avepods, while Eoraptor's jugal ventral margin is also placed at a high angle to the maxilla. The dentary is also about half mandibular length (52%; measured to tip of posteroventral process) in Staurikosaurus, and only slightly longer in Tawa (58%) than PVSJ 407 (56%), though all are longer than the Frenguellisaurus holotype (49%). Other herrerasaurids have distal caudal prezygapophyses over half of centrum length and with lateral ridges that also run along the centrum.
Novas (1993) listed many supposed autapomorphies of Herrerasaurus. A narrow, U-shaped antiorbital fossa is now known in Sanjuansaurus and Tawa. Posterior dorsal neural spine tables are also present in Sanjuansaurus and Eoraptor. The pubis also has a sinuous lateral margin in Sanjuansaurus. The femur also has an anteroproximal keel and distal anterolateral scar in Sanjuansaurus.
Sereno (1993) listed several additional pectoral and forelimb autapomorphies of Herrerasaurus. The acromion also exhibits an abrupt increase in width in Sanjuansaurus. Sereno claimed disatal carpal IV is enlarged, but this is actually distal carpal V, which is also enlarged in undescribed herrerasaurid MACN-PV 18.649a.
Holotype- (PVL 2566) (3.5 m, 210 kg) seventh dorsal vertebra (50 mm), eighth dorsal vertebra (45 mm), ninth dorsal vertebra (45 mm), tenth dorsal vertebra (45 mm), eleventh dorsal vertebra (45 mm), twelfth dorsal vertebra (45 mm), thirteenth dorsal vertebra (42 mm), fourteenth dorsal vertebra (44 mm), dorsal rib fragments, first sacral vertebra (42 mm), second sacral vertebra (56 mm), incomplete third sacral vertebra (65 mm), incomplete first caudal vertebra (49 mm), incomplete second caudal vertebra (45 mm), incomplete third caudal vertebra (42 mm), incomplete fourth caudal vertebra (47 mm), incomplete fifth caudal vertebra (46 mm), sixth caudal vertebra (47 mm), seventh caudal vertebra (50 mm), eighth caudal vertebra (50 mm), ninth caudal vertebra (50 mm), tenth caudal vertebra (50 mm), eleventh caudal vertebra, twelfth caudal vertebra (47 mm), thirteenth caudal vertebra (56 mm), fourteenth caudal vertebra (46 mm), fifteenth caudal vertebra (53 mm), sixteenth caudal vertebra (47 mm), seventeenth caudal vertebra (47 mm), eighteenth caudal vertebra (45 mm), nineteenth caudal vertebra (46 mm), twentieth caudal vertebra (48 mm), twenty-first caudal vertebra (48 mm), twenty-second caudal vertebra, twenty-third caudal vertebra (51 mm), twenty-fourth caudal vertebra (48 mm), twenty-fifth caudal vertebra (50 mm), twenty-sixth caudal vertebra (52 mm), twenty-seventh caudal vertebra (43 mm), twenty-eighth caudal vertebra (53 mm), twenty-ninth caudal vertebra (60 mm), thirtieth caudal vertebra (36 mm), thirty-first caudal vertebra (46 mm), thirty-second caudal vertebra (48 mm), thirty-fourth caudal vertebra (37 mm), thirty-fifth caudal vertebra (50 mm), thirty-sixth caudal vertebra (43 mm), thirty-seventh caudal vertebra (37 mm), thirty-eighth caudal vertebra (43 mm), thirty-ninth caudal vertebra (43 mm), fortieth caudal vertebra (43 mm), forty-first caudal vertebra (43 mm), forty-second caudal vertebra (43 mm), chevrons 1-39, ilia (240 mm), pubes (430 mm; one partial), ischia (330 mm), femur (473 mm), tibia (413 mm), fibula (415 mm), astragali (95 mm wide), calcaneum, metatarsal I (~134 mm), phalanx I-1 (~53 mm), metatarsal II (~195 mm), phalanx II-1 (~64 mm), metatarsal III (223 mm), phalanx III-1 (~75 mm), metatarsal IV (~203 mm), phalanx IV-1 (~67 mm), phalanx IV-2 (~42 mm), proximal phalanx IV-3, metatarsal V (~140 mm), phalanx V-1 (~31 mm)
Paratypes- ?(MLP 61-VIII-2-2) pelvises including ilium
(PVL 2054; PVL 2045 in Reig, 1963) (3.17 m, 114 kg) gastralia, fragments of pubes, femora (385 mm), tibiae (327 mm), fibulae, several metatarsals including metatarsal III (176 mm), pedal phalanges
?(PVL 2558a) manual ungual, fragments of femora (~440 mm), tibiae
Referred- (MACN 18.060; holotype of Ischisaurus cattoi) (2.36 m, 47 kg) (skull ~250 mm) fragmentary premaxilla, maxillae, incomplete frontals, (mandible ~255 mm) dentaries, splenials, articular end of mandible, teeth, atlantal intercentrum, axis, fourth cervical vertebra (37 mm), fifth cervical vertebra (35 mm), dorsal vertebrae (27 mm), caudal vertebrae, humeri (154, 146 mm), proximal ulnae, radiale, partial ilium, pubis, femur (286 mm), tibiae (280 mm), astragalus (43 mm wide), calcaneum, proximal metatarsal I, metatarsal II, metatarsal III (134 mm), pedal phalanges (Reig, 1963)
?(MACN 18.688) (juvenile) partial pelvis including ilium and pubis (Novas, 1993)
?(MCZ 7063) incomplete skull (Sereno and Novas, 1993)
....(MCZ 7064) axis (~65 mm), five partial dorsal vertebrae including posterior dorsal centrum (~43 mm), partial scapulocoracoids (scapula ~194 mm), partial humeri (~266 mm), partial ilium, distal pubes, distal femur, partial tibia, proximal fibula, few partial pedal phalanges (Brinkman and Sues, 1987)
?(MLP 61-VIII-2-3; paratype of Ischisaurus cattoi) (2.77 m, 75 kg) maxillary fragments, partial mandibles, vertebral fragments, coracoid, humeri (197 mm), fragmentary ulnae, femur (335 mm), tibia, pedal elements including metatarsal III (156 mm) (Reig, 1963)
(PVSJ 53; holotype of Frenguellisaurus ischigualastensis) partial skull (560 mm), mandibles (~572 mm), axis, fragments of cervical vertebrae, partial dorsal neural spine, twentieth caudal vertebra (61 mm), twenty-first caudal vertebra (59 mm), twenty-second caudal vertebra (61 mm), twenty-third caudal vertebra (60 mm), twenty-fourth caudal vertebra (59 mm), incomplete twenty-seventh caudal vertebra (60 mm), incomplete twenty-eighth caudal vertebra (55 mm), incomplete twenty-ninth caudal vertebra (56 mm), thirtieth caudal vertebra (60 mm), thirty-first caudal vertebra (60 mm), thirty-second caudal vertebra (59 mm), thirty-fourth caudal vertebra (55 mm), thirty-fifth caudal vertebra (55 mm), thirty-sixth caudal vertebra (55 mm), thirty-seventh caudal vertebra (55 mm), thirty-ninth caudal vertebra (55 mm), fortieth caudal vertebra (55 mm), forty-fifth caudal vertebra (46 mm), forty-seventh caudal vertebra (45 mm), incomplete scapulocoracoid (scapula 285 mm), distal humerus, ulnae (one partial, one distal), distal radius (Novas, 1986)
?(PVSJ 104) fragmentary pelvis, fragmentary sacrum, fragmentary hindlimb including tibia (373 mm) (Novas, 1993)
(PVSJ 373) cervical vertebrae, partial third dorsal vertebra (34 mm), fourth dorsal vertebra (34 mm), fifth dorsal vertebra (35 mm), sixth dorsal vertebra (35 mm), seventh dorsal vertebra (38 mm), eighth dorsal vertebra (38 mm), ninth dorsal vertebra (38 mm), tenth dorsal vertebra (36 mm), eleventh dorsal vertebra (37 mm), twelfth dorsal vertebra (39 mm), thirteenth dorsal vertebra (42 mm), fourteenth dorsal vertebra (36 mm), first sacral vertebra (36 mm), second sacral vertebra (43 mm), third sacral vertebra (39 mm), partial scapula, partial humerus (~175 mm), radius (153 mm), ulna (167 mm), radiale, ulnare, centrale, distal carpal II, distal carpal III, distal carpal IV, distal carpal V, metacarpal I (37 mm), phalanx I-1 (42 mm), manual ungual I (~36 mm), metacarpal II (58 mm), phalanx II-1 (36 mm), phalanx II-2 (37 mm), manual ungual II (~38 mm), metacarpal III (62 mm), phalanx III-1 (34 mm), phalanx III-2 (~28 mm), phalanx III-3 (32 mm), manual ungual III (~38 mm), metacarpal IV (33 mm), phalanx IV-1 (10 mm), metacarpal V (15 mm), ischium, femur (345 mm), tibia (315 mm), fibula (315 mm), astragalus (30 mm wide), calcaneum, distal tarsal III, distal tarsal IV, metatarsal I (100 mm), phalanx I-1 (42 mm), pedal ungual I (30 mm), metatarsal II (142 mm), phalanx II-1 (50 mm), phalanx II-2 (40 mm), pedal ungual II (37 mm), metatarsal III (165 mm), phalanx III-1 (55 mm), phalanx III-2 (38 mm), phalanx III-3 (34 mm), pedal ungual III (35 mm), metatarsal IV (143 mm), phalanx IV-1 (40 mm), phalanx IV-2 (~25 mm), partial phalanx IV-3, phalanx IV-4 (24 mm), pedal ungual IV (30 mm), metatarsal V (85 mm), partial phalanx V-1 (Sereno and Novas, 1992)
(PVSJ 380) scapula, radiale, ulnare, centrale, distal carpal I, distal carpal II, distal carpal III, distal carpal IV, distal carpal V, metacarpal I (44 mm), phalanx I-1 (56 mm), manual ungual I (42 mm), metacarpal II (66 mm), phalanx II-1 (47 mm), phalanx II-2 (51 mm), manual ungual II (47 mm), metacarpal III (74 mm), phalanx III-1 (44 mm), phalanx III-2 (35 mm), phalanx III-3 (40 mm) (Sereno, 1993)
(PVSJ 407) incomplete skeleton including skull (300 mm), several sclerotic ossicles, stapes, mandibles (~290 mm), proatlases (26 mm), atlantal intercentrum (10 mm), atlantal neurapophyses (34 mm), axis (44 mm), third cervical vertebra (40 mm), fourth cervical vertebra (~43 mm), fifth cervical vertebra (~44 mm), sixth cervical vertebra (~43 mm), seventh cervical vertebra, eighth cervical vertebra (31 mm), ninth cervical vertebra, tenth cervical vertebra, dorsal vertebrae, scapulocoracoid, humerus (170 mm), radius, ulna (168 mm), radiale, ulnare, centrale, distal carpal I, distal carpal II, distal carpal III, distal carpal IV, metacarpals (Sereno and Novas, 1992)
?(PVSJ 409) partial scapulocoracoid (Sereno, 1993)
?(PVSJ 461) incomplete skeleton including dorsal vertebrae (Novas, 1993)
Comments- The holotype was discovered in 1961, while the paratypes were discovered between 1959 and 1961. MCZ 7064 was discovered in 1958 and described by Brinkman and Sues in 1987 as cf. Staurikosaurus sp., though it and an undescribed skull associated with the specimen were later assigned to Frenguellisaurus (Paul, 1988), excluded from Staurikosaurus (Novas, 1992), and finally referred to Herrerasaurus (Novas, 1993). Novas noted the bones described by Brinkman and Sues as proximal ischia are distal pubes. Sereno et al. (1988) reported new remains discovered that year, with PVSJ 373, 407 and 461 identified as herrerasaurid (incorrectly stating cervical epipophyses are absent) and PVSJ 380 as theropod. These were preliminarily described by Sereno and Novas (1992), and in detail in a series of monographs for the skull and neck (Sereno and Novas, 1993), pectoral girdle and forelimb (Sereno, 1993) and rest of the postcrania (Novas, 1993). The complete skull of PVSJ 407 and manus of PVSJ 373 now form much of the common perception of Herrerasaurus. Sereno (2007) noted that three sacral vertebrae are present in Herrerasaurus, with the first being identified as a dorsal by earlier authors. Ezcurra (2010) noted that a distal carpal I is present in PVSJ 380, making the carpals identified by Sereno (1993) as I-IV actually II-V.
Herrerasaurus a theropod? As a carnivorous dinosaur, Herrerasaurus has almost always been associated with theropods, though that group was often thought to be diphyletic (with separate origins for coelurosaurs and carnosaurs, both sensu Huene) and often contained prosauropods the first two decades after the genus was described. Bakker and Galton (1974) placed Herrerasaurus as a basal theropod in the sense recognized today, though without discussion. This hypothesis was common after Sereno and Novas' (1992) description of new material and was nearly always combined with placing Staurikosaurus in Herrerasauridae, so it will be discussed in more detail in the comments for that family. One exception is Kischlat (2000), who used an unpublished analysis to place Herrerasaurus as a theropod more derived than Eoraptor, while Staurikosaurus was a sauropodomorph.
Herrerasaurus a carnosaur sensu Huene? Reig (1963) assigned Herrerasaurus to the Carnosauria, noting most characters were similar to sauropodomorphs except the supposed reduction in tooth number (based on an incorrectly referred skull) and large pubic boot which resembled allosaurids and tyrannosaurids. The pubic boot has since been recognized as nonhomologous with that of neotheropods, as it involves folding of the pubic blade instead of anteroposterior expansion.
Herrerasaurus a sauropodomorph? Herrerasaurus has always been recognized as being similar to what are today considered basal sauropodomorphs. In the 1960s and 1970s, the taxonomy of that group was chaotic and hindered by their frequent misassociation with pseudosuchian jaw elements and teeth. Among the earliest referrals are Rozhdestvensky and Tatarinov (1964), who placed it in their Gryponychidae, a family traditionally viewed as including prosauropod-like basal carnosaurs but now known to be based on a massopod. Reig (1970) also placed it in Gryponychidae (or a related family), which in his scheme was a family of superficially carnosaur-like prosauropods. Similar was Colbert's (1970) referral of Herrerasaurus to Teratosauridae, which was thought to be a family of basal sauropodomorphs with macropredatory skulls but is now known to be based on prosauropod-grade sauropodomorphs associated with pseudosuchian cranial elements and teeth. Ignoring the large skull and teeth (which were incorrectly referred to both Herrerasaurus and sauropodomorphs), Colbert's suggested characters include robust hindlimb, tibia shorter than femur, and broad pes. Walker (1964) thought Herrerasaurus was similar in most respects to "Plateosaurus type" saurischians but viewed the pubic boot as indicating a relationship with Poposaurus and a pubis later referred to Postosuchus, two pseudosuchians. Both Romer (1966) and Steel (1970) listed it as a plateosaurid. Van Heerden (1978) placed Herrerasaurus as a basal sauropod, with prosauropods the sister taxon to Sauropoda. He later (1979) moved it to Prosauropoda (his Palaeopoda), which he viewed as basal carnosaurs sensu Huene. Cooper (1980, 1981) thought herrerasaurids were basal prosauropods, though also the grade from which theropods arose. While these historical comparisons of Herrerasaurus to sauropodomorphs were generally vague, unpublished analysis suggests herrerasaurids can be moved to a position as basalmost sauropodomorphs without adding many extra steps.
Herrerasaurus outside Dinosauria? Gauthier (1984) was the first to place Herrerasaurus outside Saurischia+Ornithischia, which in today's nomenclature would exclude it from Dinosauria (though he and other authors advocating this arrangement still viewed it as dinosaurian). However, this was in the context of it being a herrerasaurid along with Staurikosaurus so is discussed further under the comments for that family. Brinkman and Sues (1987) excluded Herrerasaurus from Dinosauria based on the incompletely perforated acetabulum and large pedal digit V. Benton (1990) excluded it based on less than three sacral vertebrae, anterior trochanter not a spike or crest, and distal tibia not transversely expanded. Sereno and Novas (1990) presented the results of an unpublished analysis in an abstract, citing a shorter preacetabular process as evidence for excluding Herrerasaurus from Dinosauria. Fraser et al. (2002) listed the rudimentary brevis fossa as evidence Herrerasaurus is non-dinosaurian. They also listed characters supposedly excluding it from Saurischia and Theropoda, but these actually only exclude it from Eusaurischia (manual digit I not offset; manual ungual I not enlarged; manual digit III longest) and Avepoda (short preacetabular process; five tibiotarsal characters listed by Novas, 1989a) and are thus congruent with placing Herrerasaurus as a basal saurischian or basal theropod as well. No recent cladistic analysis has excluded Herrerasaurus from Dinosauria.
Ischisaurus- The holotype of Ischisaurus cattoi was found in 1960 and only briefly described and illustrated, leading to general uncertainty regarding its relationships. Manual elements and parts of the right pes were reported by Reig, but not listed by Novas (1993). Ischisaurus was first thought to represent a taxon intermediate between coelurosaurs sensu Huene and pachypodosaurs (Huene's carnosaur+sauropodomorph clade) (Reig, 1963). Some authors placed it in families that were thought to be carnivorous prosauropods at the time (Gryponychidae- Rozhdestvensky and Tatarinov, 1964; Palaeosauriscidae- Colbert, 1970), while others placed it in families containing small standard prosauropods (Thecodontosauridae- Reig, 1970; Anchisauridae- Steel, 1970). Few provided evidence, though Colbert stated it was more similar to Staurikosaurus (which he also considered a palaeosauriscid) than Herrerasaurus in femoral curvature, the position of the femoral head, and the size and position of the fourth trochanter. Another common placement was as a theropod (Demathieu, 1970; Bakker and Galton, 1974; Galton and Cluver, 1976), with Bakker and Galton specifying a very basal placement in the clade. It was assigned to Herrerasauridae by Gauthier (1984, 1986), Novas (1986, 1989a, 1989b), Sues (1990), Olshevsky (1991) and Bonaparte (1997), and Staurikosauridae by Paul (1988).
The diagnosis of Ischisaurus contains several characters intended to distinguish the genus from Herrerasaurus, and Reig notes he initially considered the former a juvenile of the latter. Of these characters, the cranial ones (four premaxillary teeth; slender dentary; ~15 dentary teeth; teeth laterally compressed and recurved), supposedly short humerus and "normal olecranon" on the ulna are based on differences from PVL 2558b (Novas, 1993), which is no longer referred to Herrerasaurus. The smaller size and slightly larger tibiofemoral ratio are potentially ontogenetic. "Laterally shortened proximal end of tibia" is rather vague, but Novas' (1993) measurements do indicate Ischisaurus' tibia is only 6% wider proximally than distally, compared to 36% in the Herrerasaurus holotype and 58% in PVSJ 373. Contra Reig's diagnosis, Ischisaurus' calcaneum is actually smaller (37% of astragalar transverse width) than the Herrerasaurus holotype's (47%), though slightly larger than PVSJ 373's (32%). The cervicals are said to lack ventral keels, unlike PVSJ 407 (and Sanjuansaurus). Novas (1986) noted Ischisaurus differs from Frenguellisaurus in having a posterodorsal surangular process lateral to the glenoid (which he felt was an autapomorphy), but he later (1993) stated this was due to deformation and preparation differences (PVSJ 407 also has the process, so presumably Frenguellisaurus' condition is the deformed one). Romer (1966) and Cooper (1981) were the first to synonymize it with Herrerasaurus, though without stated evidence. Novas (1992) and Sereno and Novas (1992) synonymized it with Herrerasaurus based on unspecified information from the specimens found in 1988. Novas (1993) elaborated, stating that supposed differences from Herrerasaurus were due to PVL 2558b as noted above, that the two shared autapomorphies, and "do not show any differences." This has been accepted uncritically by most workers ever since. However, the tibial and calcaneal differences noted by Reig were based on PVL 2558a (still referred to Herrerasaurus by Novas) and Novas states on the same page that Herrerasaurus' holotype has shorter posterior dorsal centra than Ischisaurus' holotype or the specimens discovered in 1988 (PVSJ 373, 407 and 461). Bonaparte (1997) noted the latter and found the synonymy had not been convincingly demonstrated, retaining Ischisaurus as a distinct genus a decade later (Bonaparte et al., 2007). Langer and Benton (2006) stated Ischisaurus did not preserve either of the two apomorphies they diagnosed Herrerasaurus with (as it lacks a pubis or ischium) and has less compressed dorsals, but noted it resembles Herrerasaurus more than Staurikosaurus "in the longer ischiadic peduncle of the ilium, the more dorsally expanded cnemial crest and the square distal tibia." They also noted the holotypes of Ischisaurus and Herrerasaurus both have a platform anterior to the astragalar ascending process, but PVSJ 373 does not. They concluded it was probable Ischisaurus was synonymous with Herrerasaurus, but Langer et al. (2009) only considered it a possible synonym. Complicating matters is the recent discovery of Sanjuansaurus, which shares almost all the supposed Herrerasaurus autapomorphies listed by Sereno and Novas which it can be evaluated for. Unfortunately, the only valid Herrerasaurus apomorphy which can be evaluated for Ischisaurus in the literature is the comparatively large tibiofemoral ratio, which supports synonymizing the two. The lack of cervical ventral keels, more elongate dorsal centra and transversely narrow proximal tibia may suggest a taxonomic distinction, but this cannot be pursued further without access to the material given the meager description and illustration in the literature. Notably, the variation noted above and incompleteness of most specimens suggests that if Ischisaurus is valid, the referral of most known individuals to Herrerasaurus would be questionable.
Frenguellisaurus- Frenguellisaurus ischigualastensis was discovered in 1975 and described by Novas (1986) as a primitive saurischian, probably with theropod affinities. It was later assigned to Herrerasauridae by Paul (1988), Novas (1989b) and Bonaparte (1997), and to Staurikosauridae by Olshevsky (1991), while Molnar (1990) suggested it might be a dinosaur outgroup related to Staurikosaurus and herrerasaurids. Paul viewed it as a basal herrerasaurid but placed it in that family instead of Staurikosauridae based on the dorsal centra of MCZ 7064, which has since been referred to Herrerasaurus.
Novas (1992) and Sereno and Novas (1992) synonymized Freguellisaurus with Herrerasaurus based on unspecified information from the specimens found in 1988. Novas elaborated in 1993, stating that the separation from Herrerasaurus was based on the incorrect assumption that genus had short distal caudal prezygapophyses (though Novas never noted distinguishing characters in his 1986 paper) and that the two genera share autapomorphies. Bonaparte (1997) felt the synonymy had not been demonstrated, though he did not provide supporting evidence. Langer and Benton (2006) state that while Frenguellisaurus does not preserve any of the Herrerasaurus apomorphies they recognized, it was similar to PVSJ 407 in having "a narrow U-shaped maxillary antorbital fossa, a lateral ridge on the jugal, a squared ventral ramus of the squamosal, a dorsally narrow laterotemporal fenestra and a mediocaudally expanded quadratojugal." However, they also noted it has a shorter dentary and stronger lateral ridges on its distal caudals than either Staurikosaurus or Herrerasaurus, as well as larger 'canine' maxillary teeth. These differences were said to be possibly autapomorphic, but also possibly ontogenetic due to Frenguellisaurus' large size. As with Ischisaurus, they consider Frenguellisaurus probably synonymous with Herrerasaurus, but Langer et al. (2009) only considered it a possible synonym. Similar to the situation noted above for Ischisaurus, Sanjuansaurus complicates the referral of Frenguellisaurus to Herrerasaurus. An additional issue is the lack of much overlap between holotypes, which share only a dorsal neural spine and caudal vertebrae. While Frenguellisaurus is more similar to PVSJ 407 than Sanjuansaurus in having a highly convex ventral maxillary margin, PVSJ 407 itself only shares dorsal vertebrae with the Herrerasaurus holotype (which differ from it and are like Ischisaurus in being relatively elongate). The variation described above between specimens is here viewed as interspecific, but if it is taxonomic the referral of Frenguellisaurus and other specimens like PVSJ 407 to Herrerasaurus is in question. Further analysis requires access to the specimens, especially the Ischisaurus types and MACN 7063, which all contain cranial material.
Not Herrerasaurus- PVL 2558 was a paratype of Herrerasaurus, but includes two individuals. Bonaparte (1970) first recognized the cranial elements were probably different. The individual they belong to was recatalogued as PVL 2558b, excluded from Herrerasauridae by Novas (1986, 1989a) and referred to Archosauria indet. by Novas (1993). It includes a premaxilla, maxillae, dentaries, presacral and caudal vertebrae, humeri, ulnae and proximal pubes. Subtracting the material Novas (1993) retains in Herrerasaurus, this leaves a posterior mandible, partial ilium, fibulae, partial metatarsals and proximal pedal phalanges unaccounted for. Note Novas (1989a) retains the humeri, ulnae and fibulae in Herrerasauridae, contra his later work.
PVL 2264 is a partial femur from the Los Colorados Formation that was another paratype of Herrerasaurus, but was referred to Archosauria indet. by Novas (1993).
References- Reig, 1963. La presencia de dinosaurios saurisquios en los "Estratos de Ischigualasto" (Mesotriasico Superior) de las provincias de San Juan y La Rioja (República Argentina) [The presence of saurischian dinosaurs in the "Ischigualasto beds" (upper Middle Triassic) of San Juan and La Rioja Provinces (Argentine Republic)]. Ameghiniana. 3, 3-20.
Rozhdestvensky and Tatarinov, 1964. Order Saurischia. in Orlov (ed.). Osnovy Paleontologii. 529-552.
Romer, 1966. Vertebrate Paleontology, 3rd edition. University of Chicago Press, Chicago. 1-468.
Bonaparte, 1970. Annotated list of the South American Triassic tetrapods. Second Gondwana Symposium, Proceedings and Papers. 665-682.
Reig, 1970. The Proterosuchia and the early evolution of the archosaurs; an essay about the origin of a major taxon. Bulletin of The Museum of Comparative Zoology. 139, 229-292.
Steel, 1970. Part 14. Saurischia. Handbuch der Paläoherpetologie/Encyclopedia of Paleoherpetology. Gustav Fischer Verlag, Stuttgart. 1-87.
Benedetto, 1973. Herrerasauridae, nueva familia de saurisquios triasicos. Ameghiniana. 10(1), 89-102.
Bakker and Galton, 1974. Dinosaur monophyly and a new class of vertebrates. Nature. 248, 168-172.
Galton and Cluver, 1976. Anchisaurus capensis (Broom) and a revision of the Anchisauridae (Reptilia, Saurischia). Annals of the South African Museum. 69(6), 121-159.
van Heerden, 1978. Herrerasaurus and the origin of sauropod dinosaurs. South African Journal of Science. 74, 187-189.
van Heerden, 1979. The morphology and taxonomy of Euskelosaurus (Reptilia: Saurischia; Late Triassic) from South Africa. Navorsinge van die Nasionale Museum, Bloemfontein. 4, 21-84.
Cooper, 1980. The prosauropod ankle and dinosaur phylogeny. South African Journal of Science. 76, 176-178.
Cooper, 1981. The prosauropod dinosaur Massospondylus carinatus Owen from Zimbabwe: Its biology, mode of life and phylogenetic significance. Occasional Papers of the National Museums and Monuments of Rhodesia (series B, Natural Sciences). 6, 689-840.
Gauthier, 1984. A cladistic analysis of the higher systematic categories of the Diapsida. PhD thesis. University of California, Berkeley. 564 pp.
Novas, 1986. Un probable teropodo (Saurischia) de la Formación Ischigualasto (Triásico superior), San Juan, Argentina. [A probable theropod (Saurischia) from the Ischigualasto Formation (Upper Triassic), San Juan, Argentina.] [in Spanish, with English summ.]. Actas del Congreso Argentino de Paleontología y Bioestratigrafía. 4, 1-6.
Brinkman and Sues, 1987. A staurikosaurid dinosaur from the Upper Triassic Ischigualasto Formation of Argentina and the relationships of the Staurikosauridae. Palaeontology. 30, 493-503.
Paul, 1988. Predatory dinosaurs of the world. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Sereno, Novas, Arcucci and Yu, 1988. New evidence on dinosaur and mammal origins from the Ischigualasto Formation (Upper Triassic, Argentina). Journal of Vertebrate Paleontology. 8(3), 26A.
Novas, 1989a. The tibia and tarsus in Herrerasauridae (Dinosauria, incertae sedis) and the origin and evolution of the dinosaurian tarsus. Journal of Paleontology. 63, 677-690.
Novas, 1989b. Los dinosaurios carnivoros de la Argentina. PhD thesis. Universidad Nacional de La Plata. 510 pp.
Benton, 1990. Origin and interrelationships of dinosaurs. in Weishampel, Dodson and Osmolaska (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 11-30.
Molnar, 1990. Problematic Theropoda: "Carnosaurs". in Weishampel, Dodson and Osmolaska (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 306-317.
Sereno and Novas, 1990. Dinosaur origins and the phylogenetic position of pterosaurs. Journal of Vertebrate Paleontology. 10(3), 42A.
Sues, 1990. Staurikosaurus and Herrerasauridae. in Weishampel, Dodson and Osmolaska (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 143-147.
Novas, 1992. Phylogenetic relationships of the basal dinosaurs, the Herrerasauridae. Palaeontology. 35, 51-62.
Sereno and Novas, 1992. The complete skull and skeleton of an early dinosaur. Science. 258, 1137-1140.
Novas, 1993. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina. Journal of Vertebrate Paleontology. 13, 400-423.
Sereno and Novas, 1993. The skull and neck of the basal theropod Herrerasaurus ischigualastensis. Journal of Vertebrate Paleontology. 13, 451-476.
Sereno, 1993. The pectoral girdle and forelimb of the basal theropod Herrerasaurus ischigualastensis. Journal of Vertebrate Paleontology. 13, 425-450.
Franzosa, 2001, Herrerasaurus ischigualastensis (On-line), Digital Morphology. Accessed October 21, 2010 at http://digimorph.org/specimens/Herrerasaurus_ischigualastensis/.
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Sereno, 2007. The phylogenetic relationships of early dinosaurs: a comparative report. Historical Biology. 19(1), 145-155.
Paulina-Carabajal, 2009. El neurocráneo de los dinosaurios Theropoda de la Argentina: Osteología y sus implicancias filogenéticas. PhD thesis, Universidad Nacional de La Plata. 554 pp.
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Chindesaurus Long and Murry, 1995
= "Chindesaurus" Long vide Miller, 1985
= "Holbrookosaurus" Anonymous?, 1986
= Caseosaurus Hunt, Lucas, Heckert, Sullivan and Lockley, 1998
C. bryansmalli Long and Murry, 1995
= "Holbrookosaurus smallii" Anonymous?, 1986
= Caseosaurus crosbyensis Hunt, Lucas, Heckert, Sullivan and Lockley, 1998
Late Norian, Late Triassic
Petrified Forest Member of Chinle Formation, Arizona, New Mexico, US
Holotype
- (PEFO 10395) (~2.4 m) tooth (22 mm), incomplete cervical centrum (~32 mm), four posterior dorsal centra (27.5 mm), rib fragments, first sacral vertebra (32 mm), second sacral vertebra (38 mm), proximal caudal centrum, several mid caudal vertebrae, chevron, fragmentary ilia, proximal pubis, pubic shaft, ischial fragments, femora (one proximal) (265 mm), fragmentary tibia, astragalus (31 mm wide)
Paratype- ?(UCMP coll.) (>2.4 m) two sacral centra
Referred- (GR 210) femur (Whiteside et al., 2015)
(GR 226) femur (Irmis et al., 2007)
(H2-001-080622) proximal femur (Whiteside et al., 2015)
(NMMNH P-35995) ilium (Nesbitt et al., 2007)
(PEFO 33982) nine vertebrae, ilium fragment, proximal femur, bone fragments (Nesbitt et al., 2007)
Late Carnian, Late Triassic
Colorado City Formation? (Otis Chalk) of the Dockum Group, Texas, US
Paratype
- ?(TMM 31100-523) (~2.4 m) proximal femur (~265 mm)
Late Carnian, Late Triassic
Tecovas Formation of Dockum Group, Texas, US
(UMMP 8870; holotype of Caseosaurus crosbyensis) partial ilium (140.5 mm) (Case, 1927)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, New Mexico, US

?(NMMNH P-4882) sacral vertebra (Heckert, Lucas and Sullivan, 2000)
?(NMMNH P-16844) dorsal centrum (Heckert, Lucas and Sullivan, 2000)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, Texas, US

(TTUP 10082) partial ilium, incomplete pubis, ischial fragments (Lehane, 2005)
Diagnosis- (after Nesbitt et al., 2007) triangular rugosity on the postacetabular process; proximal tibial intercondylar groove that is strongly situated medially; posterior edge of the fibular condyle of the proximal tibia that is straight in proximal view; ventral cleft on astragalus.
Comments- When first discovered in 1984, the holotype was referred to Prosauropoda in the popular literature (Meyer, 1986). Then assigned to Herrerasauridae by Long and Murray (1995), Hunt (1996), Novas (1997) and Hunt et al. (1998), Sereno (1999) found Chindesaurus to be the sister taxon of Staurikosaurus within Herrerasauridae in his phylogenetic analysis, while Bittencourt and Kellner's (2004) analysis found it to be a non-herrerasaurid herrerasaurian. Langer noted characters congruent with the latter assessment. Rauhut (2003) found it difficult to assign Chindesaurus to a less inclusive taxon than Dinosauriformes, though he agrees with Langer that the dorsals are less shortened than herrerasaurids. Furthermore, he finds the medially expanded brevis shelf resembles pseudosuchians more than it does the laterally expanded brevis shelf of most dinosaurs. Rauhut believes the holotype may pertain to multiple individuals and/or taxa, though no later author has agreed. Yates (2006) include Chindesaurus in a phylogenetic analysis of sauropodomorphs and find it to be a theropod closer to Avepoda than Guaibasaurus or Agnostiphys, while Ezcurra (2010) modified the matrix and found it in a trichotomy with sauropodomorphs and theropods. Nesbitt et al. (2007) note a range of characters shared with various saurischians, complicating its phylogenetic relationships. Irmis et al. (2007) find Chindesaurus to be a herrerasaurid in their analysis, though they have a small taxon sample. More recently, Bittencourt Rodrigues (2010) also found it to be a herrerasaurid.
First identified as Coelophysis sp. by Case (1927), Long and Murray (1995) made UMMP 8870 a paratype of Chindesaurus bryansmalli. Hunt et al. (1998) later made it the holotype of a new taxon of herrerasaurid- Caseosaurus crosbyensis. This was based on the shallow brevis shelf, a more ventrally placed medial longitudinal ridge and a transversely thinner postacetabular blade. Langer (2004) has synonymized the taxa without comment, though Nesbitt et al. noted both the Chindesaurus and Caseosaurus holotypes lack brevis shelves and the other two differences are probably size related. Although they found an apomorphy shared by both specimens, they decided not to officially synonymize them due to their fragmentary condition. I follow Langer's decision to synonymize them here.
Nesbitt et al. find an ilium (NMMNH P-35995) originally assigned by Heckert et al. (2000, 2003) to Eucoelophysis sp. is extremely similar to the Caseosaurus holotype based on several characters. They and Stocker (2013) also substantiate the referral of a proximal femur (TMM 31100-523) to Chindesaurus by Long and Murry based on a unique combination of characters, contra Hunt et al. (1998). Nesbitt and Ezcurra (2015) state this femur is also indistinguishable from Herrerasaurus, but its locality still suggests Chindesaurus is a more probable identification.
Hunt et al. (1998) described a tooth in the holotype which wasn't noted by Long and Murry (1995).
Heckert et al. (2000) described a sacral (NMMNH P-4882) from the Bull Canyon Formation of New Mexico as possibly being Chindesaurus, a dorsal centrum (NMMNH P-16844) as generally conforming to the genus. Whether these assessments are correct requires further study. Lehane (2005) referred a partial pelvis (TTUP 10082) to Shuvosaurus, while Lehman and Chatterjee (2005) referred it to Coelophysis. Nesbitt and Chatterjee (2008) and Martz et al. (2013) identified it as herrerasaurid but did not assign it to a genus. Though they believed the presence of three sacral vertebrae was notable, both Herrerasaurus and Staurikosaurus have since been discovered to have this as well. The anterodistal bevel was previously considered an autapomorphy of Staurikosaurus and is otherwise unpreserved in North American herrerasaurids. The posterolateral rugosity on the postacetabular process has been considered an autapomorphy of Chindesaurus however, which also matches the general morphology and stratigraphy. TTUP 10082 is thus referred to that genus here. If properly assigned, TTUP 10082 would cement Chindesaurus' position as a herrerasaurian, though exactly where it falls in the clade would depend on including the pevis in phylogenetic analyses.
Not Chindesaurus- Long and Murry (1995) refer five dorsal centra (UCMP 177316) from the Mesa Redondo Member of the Chinle Formation to Chindesaurus, which were later assigned to Herrerasauridae indet. by Hunt et al. (1998). As Chindesaurus may not be a herrerasaurid, the centra are assigned to Saurischia indet. here.
PEFO 4849 is a dorsal centrum from the Petrified Forest Formation which was referred to Chindesaurus by Murry and Long (1989) and Long and Murry (1995), then assigned to Herrerasauridae indet. by Hunt et al. (1998). As Chindesaurus may not be a herrerasaurid, the centrum is assigned to Saurischia indet. here.
NMMNH P4415 is a proximal femur from the Bull Canyon Formation which was referred to Chindesaurus by Long and Murry (1995) and assigned to herrerasaurid A by Hunt (1994) and Hunt et al. (1998). It was described and illustrated by Heckert et al. (2000) as herrerasaurid B, noted to be similar to Chindesaurus in its medially angled and squared-off femoral head, but these characters are also seen in shuvosaurids and other basal dinosaurs. Hence this femur is listed here under Archosauria indet..
NMMNH P16656 consists of dorsal and caudal centra from the Bull Canyon Formation which were referred to Chindesaurus by Murry and Long (1989) and Long and Murry (1995), assigned to herrerasaurid C by Hunt (1994) and Hunt et al. (1998), but were found to be unidentifiable past Archosauria indet. by Nesbitt et al. (2007).
NMMNH P17325 consists of a dorsal centrum from the Bull Canyon Formation which was referred to Chindesaurus by Long and Murry (1995) and assigned to (?)Theropoda by Hunt (1994) and Hunt et al. (1998). Nesbitt et al. (2007) note dorsal centra referred to Chindesaurus are not diagnostic past Archosauria indet..
References- Case, 1927. The vertebral column of Coelophysis Cope. Contributions from the Museum of Geology, University of Michigan. 10, 209-222.
Miller, 1985. Ghost from the dawn of the dinosaurs - oldest dinosaur skeleton found in Arizona. Science News. 126(21), 325.
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Meyer, 1986. D-Day on the Painted Desert. Arizona Highways. 62(7), 3-13.
Paul, 1988. Predatory dinosaurs of the world. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Murry and Long, 1989. Geology and paleontology of the Chinle Formation, Petrified Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils of the southwestern Upper Triassic. In Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. 29-64.
Hunt, 1994. Vertebrate paleontology and biostratigraphy of the Bull Canyon Formation (Chinle Group: Norian), east-central New Mexico with revisions of the families Metoposauridae (Amphibia: Temnospondyli) and Parasuchidae (Reptilia: Archosauria). PhD thesis, University of New Mexico. 403 pp.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, 1996. A new clade of herrerasaur-like theropods from the Late Triassic of western North America. Journal of Vertebrate Paleontology 16(3), 43A.
Novas, 1997. Herrerasauridae. pp. 303–311. in Currie and Padian (eds). Encyclopedia of Dinosaurs. Academic Press, San Diego.
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Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Heckert, Lucas and Sullivan, 2000. Triassic dinosaurs in New Mexico. in Lucas and Heckert (eds). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin. 17, 17-26.
Heckert, Zeigler, Lucas, Rinehart and Harris, 2000. Preliminary description of coelophysoids (Dinosauria: Theropoda) from the Upper Triassic (Revueltian: early-mid Norian) Snyder quarry, north-central New Mexico. New Mexico Museum of Natural History & Science Bulletin. 17, 27-32.
Heckert, Zeigler, Lucas and Rinehart, 2003. Coelophysids (Dinosauria: Theropoda) from the Upper Triassic (Revueltian) Snyder quarry. New Mexico Museum of Natural History & Science Bulletin. 24, 127-132.
Rauhut, 2003. The interrelationship and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-215.
Bittencourt and Kellner, 2004. The phylogenetic position of Staurikosaurus pricei Colbert, 1970 from the Triassic of Brazil. Journal of Vertebrate Paleontology. 24(3).
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Lehane, 2005. Anatomy and relationships of Shuvosaurus, a basal theropod from the Triassic of Texas. Masters thesis, Texas Tech University. 92 pp.
Lehman and Chatterjee, 2005. Depositional setting and vertebrate biostratigraphy of the Triassic Dockum Group of Texas. Journal of Earth System Science. 114(3), 325-351.
Nesbitt, Irmis and Parker, 2005. Critical review of the Late Triassic dinosaur record, part 3: Saurischians of North America. Journal of Vertebrate Paleontology. 25(3), 96A.
Irmis, Nesbitt, Padian, Smith, Turner, Woody and Downs, 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science. 317, 358-361.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.
Yates, 2007. Solving a dinosaurian puzzle: the identity of Aliwalia rex Galton. Historical Biology. 19(1), 93-123.
Nesbitt and Chatterjee, 2008. Late Triassic dinosauriforms from the Post Quarry and surrounding areas, west Texas, U.S.A. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen. 249(2), 143-156.
Bittencourt Rodrigues, 2010. Revisao filogenetica dos dinossauriformes basais: Implicacoes para a origem dod dinossauros. Unpublished Doctoral Thesis. Universidade de Sao Paulo. 288 pp.
Ezcurra, 2010. A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: A reassessment of dinosaur origin and phylogeny. Journal of Systematic Palaeontology. 8(3), 371-425.
Martz, Mueller, Nesbitt, Stocker, Parker, Atanassov, Fraser, Weinbaum and Lehane, 2013. A taxonomic and biostratigraphic re-evaluation of the Post Quarry vertebrate assemblage from the Cooper Canyon Formation (Dockum Group, Upper Triassic) of southern Garza County, western Texas. Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103, 1-26.
Stocker, 2013. Contextualizing vertebrate faunal dynamics: New perspectives from the Triassic and Eocene of western North America. PhD thesis, University of Texas at Austin. 297 pp.
Nesbitt and Ezcurra, 2015. The early fossil record of dinosaurs in North America: A new neotheropod from the base of the Upper Triassic Dockum Group of Texas. Acta Palaeontologica Polonica. 60(3), 513-526.
Whiteside, Lindstrom, Irmis, Glasspool, Schaller, Dunlavey, Nesbitt, Smith and Turner, 2015. Extreme ecosystem instability suppressed tropical dinosaur dominance for 30 million years. Proceedings of the National Academy of Sciences. 112(26), 7909-7913.

Eoraptor Sereno, Forster, Rogers and Monetta, 1993
E. lunensis Sereno, Forster, Rogers and Monetta, 1993
Late Carnian-Early Norian, Late Triassic
Cancha de Bochas Member of the Ischigualasto Formation, San Juan, Argentina
Holotype
- (PVSJ 512) (~1.2 m young adult) incomplete skull (123 mm), mandibles (110 mm), proatlas, atlas, partial axis, incomplete third cervical vertebra, incomplete fourth cervical vertebra (24 mm), fifth cervical vertebra (24 mm), incomplete sixth cervical vertebra (23 mm), seventh cervical vertebra (20 mm), eighth cervical vertebra (18 mm), ninth cervical vertebra (17 mm), partial cervical ribs, first dorsal vertebra (17 mm), second dorsal vertebra (16 mm), third dorsal vertebra (16 mm), fourth dorsal vertebra (17 mm), fifth dorsal vertebra (~19 mm), sixth dorsal vertebra (20 mm), seventh dorsal vertebra (20 mm), eighth dorsal vertebra (21 mm), ninth dorsal vertebra (20 mm), tenth dorsal vertebra (21 mm), eleventh dorsal vertebra (22 mm), twelfth dorsal vertebra (23 mm), thirteenth dorsal vertebra (21 mm), fourteenth dorsal vertebra (20 mm), fifteenth dorsal vertebra (18 mm), eleven partial dorsal ribs, gastralia, first sacral vertebra (~19 mm), second sacral vertebra (~19 mm), third sacral vertebra (20 mm), first caudal vertebra (17 mm), second caudal vertebra (20 mm), third caudal vertebra (22 mm), fourth caudal vertebra (20 mm), fifth caudal vertebra (20 mm), sixth caudal vertebra (21 mm), seventh caudal vertebra (20 mm), eighth caudal vertebra (19 mm), ninth caudal vertebra (19 mm), tenth caudal vertebra (18 mm), eleventh caudal vertebra (18 mm), twelfth caudal vertebra (17 mm), thirteenth caudal vertebra (17 mm), fourteenth caudal vertebra (17 mm), fifteenth caudal vertebra (17 mm), sixteenth caudal vertebra (17 mm), seventeenth caudal vertebra (20 mm), eighteenth caudal prezygapophyses, fifteen partial chevrons (~20-35 mm), scapulae (81 mm; one fragmentary), coracoid (22 mm), humeri (85 mm), radii (63 mm), ulnae (64 mm), radiale, ulnares, distal carpal I, distal carpal II, distal carpal III, distal carpal IV, distal carpal V, metacarpals I (14 mm), phalanges I-1 (14 mm), manual ungual I (14 mm), metacarpals II (20 mm), phalanx II-1 (12 mm), phalanx II-2 (12 mm), manual ungual II (14 mm), metacarpals III (21 mm), phalanges III-1 (10 mm), phalanx III-2 (9 mm), partial phalanx III-3 (8 mm), incomplete manual ungual III (~12 mm), metacarpals IV (16 mm), metacarpals V (10 mm), ilia (82 mm), pubes (121 mm), ischia (114 mm), femora (152 mm), tibiae (156 mm), fibulae (154 mm), astragalus (27 mm transversely), calcaneum, distal tarsal III, distal tarsal IV, metatarsals I (45 mm), phalanx I-1 (21 mm), pedal ungual I (15 mm), metatarsals II (72 mm), phalanx II-1 (25 mm), phalanx II-2 (19 mm), pedal ungual II (17 mm), metatarsals III (81 mm), phalanx III-1 (27 mm), phalanx III-2 (22 mm), partial phalanx III-3 (18 mm), metatarsals IV (74 mm), phalanx IV-1 (18 mm), phalanx IV-2 (14 mm), phalanx IV-3 (12 mm), phalanx IV-4 (11 mm), pedal ungual IV (16 mm), metatarsals V (35 mm; one incomplete)
Referred- (PVSJ 559) (~1.3 m adult) first dorsal vertebra, second dorsal vertebra, partial dorsal ribs, incomplete femur, tibia (170 mm), distal fibula, astragalus, calcaneum, metatarsal fragments (Sereno et al., 2013)
(PVSJ 745) (subadult) basisphenoid, basioccipital, several partial cervical vertebrae, several partial dorsal vertebrae, partial ilia, ischial fragment, partial femora, proximal tibiae, proximal fibula, two proximal metatarsals (Sereno et al., 2013)
?(PVSJ 852) (subadult) femur (Sereno et al., 2013)
?(PVSJ 855) (adult) femur (Sereno et al., 2013)
?(PVSJ 860) (adult) proximal femur, distal femora, proximal tibiae, distal tibia, proximal fibula (Sereno et al., 2013)
(PVSJ 862) (subadult) proximal humerus, distal femora, distal tibia, proximal fibula, astragalus (Sereno et al., 2013)
?(PVSJ 876) (adult) partial femur (Sereno et al., 2013)
Diagnosis- (after Sereno et al., 1993) leaf-shaped premaxillary and anterior maxillary teeth (also in derived silesaurids, sauropodomorphs and partly in ornithischians).
(after Langer and Benton, 2006) extremely transversely compressed proximal fibula.
(after Sereno et al., 2013) premaxilla posterolateral process with tongue-shaped distal expansion; nasal with transversely broad, horizontal shelf with convex lateral margin that overhangs antorbital fossa; pterygoid process on posterior palate margin articulates laterally in synovial socket in ectopterygoid (possibly also in Pampadromaeus); narrow premaxilla-maxilla diastema approximately one crown in width; maxillary crowns with a prominent lateral eminence; accessory articular process on medial aspect of mid cervical prezygapophyses; extreme hollowing of dorsal centra and neural arches.
Differs from Panphagia in- shallow neurovascular groove on lateral aspect of dentary; less pronounced ridge on lateral aspect of surangular; less expanded distal scapular blade (approximately twice neck width); more perpendicular distal border on scapular bade; longer pubic blades (more than four times distal blade width); cnemial crest and opposing proximal condyles more anteroposteriorly expanded; tibial distal end more transversely expanded; ascending process and posterior fossa on astragalus much broader transversely (approximately one-third the width of the astragalus).
Differs from Chromogisaurus in- more strongly inturned femoral head; markedly asymmetrical shape of fourth trochanter; tibial cnemial crest and opposing proximal condyles more anteroposteriorly expanded; tibial distal end more transversely expanded.
Other diagnoses- Sereno et al. (1993) listed the "slightly enlarged" external naris as a diagnostic character, to distinguish it from the enlarged nares of sauropodomorphs, but Eoraptor's naris is actually smaller than Silesaurus, Herrerasaurus, basal ornithischians and Tawa compared to skull height. The slender subnarial process of the premaxilla is present in all basal dinosauriforms except Herrerasaurus and ornithischians.
Rauhut (2003) listed "ventral process of the postorbital flexed sharply anteriorly in its lower part", but this is no different from Heterodontosaurus or Herrerasaurus.
Comments- Eoraptor's holotype was discovered in 1991 and given a preliminary description two years later (Sereno et al., 1993). Martinez (pers. comm. 2010) confirms the detailed description was complete in 1995, which has finally been published as Sereno et al. (2013).
While Sereno et al. (1993) stated the holotype is an adult, Tykoski (2005) noted that the poorly sutured cranial elements (including frontals, parietals, basisphenoid, basioccipital and supraoccipital), large orbits and short snout, and unfused scapulocoracoid, might suggest it is young. Most recently, Sereno et al. (2013) confirm all the presacral and caudal neural arches are partially or completely fused to their centra. The sacral neurocentral sutures are open however, and the combined evidence led the authors to consider it a young adult.
Tykoski (2005) proposes that Eoraptor may have four sacrals because there is enough room posterior to the third to attach a fourth to the postacetabular process, but Sereno (2007) reiterates that only three are present. Sereno (2007) also notes the presence of pterygoid teeth and an intermandibular joint.
Eoraptor was originally described as a theropod basal to herrerasaurids and avepods (Novas, 1993; Sereno et al., 1993; Sereno, 1999; Kischlat, 2000; Rauhut, 2003; Ezcurra and Novas, 2007). Two subsequent hypotheses are that it's outside the sauropodomorph-theropod clade (Eusaurischia) (Langer, 2004; Langer and Benton, 2006; Yates, 2007; Smith et al., 2007) or sister to avepods (Tykoski, 2005; Ezcurra, 2006, 2010; Nesbitt et al., 2009). Recently, Martinez et al. (2011) has placed it as a basal sauropodomorph. None of the alternatives has strong support, leaving Eoraptor's relationships within Saurischia controversial. The sections below note the characters authors have claimed support each position (with a few notes when another author has claimed the character to be inaccurate), but a full analysis will appear here in the future.
Eoraptor basal to Eusaurischia? The first authors to propose excluding Eoraptor from Theropoda and (Eu)Saurischia were Padian and May (1993), who merely stated it lacked most theropod and saurischian synapomorphies listed by Gauthier and other workers, so was not a theropod and perhaps not a dinosaur.
Holtz (1995) had a similar conclusion in an unpublished analysis, finding Eoraptor to be sister to Eusaurischia or Dinosauria.
Padian et al. (1999) listed a few characters supporting it-
1. centra of posterior cervical vertebrae (6-8) subequal in length to those of anterior dorsal vertebrae (refined by Langer, 2004). This is untrue in Eoraptor, as cervicals 6-8 are 18-23 mm and the anterior dorsals are 16-17 mm.
2. third finger of the manus longer than second finger. As this is true in Eodromaeus and Tawa, the opposite condition in known sauropodomorphs and avepods is probably convergent. Herrerasaurus also has III longer than II, while Guaibasaurus is like sauropodomorphs and avepods. Though Tianyulong has an avepod-like condition, Heterodontosaurus and derived ornithischians have III longest, so this was more likely the basal condition in Ornithischia.
3. metatarsal I contacts tarsus. Since this is also true in Sauropodomorpha, the authors had no reason to list it. It only excludes Eoraptor from Avepoda.
Langer (2004)
4. subnarial premaxillary process extends posteriorly to the external naris. This is also true in Daemonosaurus, making the basal condition for Theropoda ambiguous. As Herrerasaurus has this state as well, it's even worse if herrerasaurids are theropods.
5. radius >80% of humerus length. This is untrue in Eoraptor, which has a ratio of 74%.
6. manual ungual I shorter than metacarpal I. This is untrue in Eoraptor, which has a ratio of 100%. Note even if the ratio is actually barely in agreement, Eodromaeus and Tawa have short unguals I too, so it would be another character convergent in avepods and sauropodomorphs.
7. metacarpal III longer than metacarpal II. Another character also present in Eodromaeus and Tawa, again making sauropodomorphs and avepods convergent. As with the digit length comparison, Guaibasaurus has longer II while Herrerasaurus has longer III. Also while heterodontosaurids have longer II, other ornithischians and Saltopus have longer III, suggesting the former is convergence.
8. distal end of ischium unexpanded. This is untrue in Eoraptor.
9. medial margin of distal tibia not broader than lateral margin. This is untrue in Eoraptor.
Smith et al. (2007)
10. maxillary tooth count 12-18. Eoraptor has 17, while basal theropods have (9/10)-11 (Daemonosaurus, Eodromaeus) and basal sauropodomorphs have at least 20 (Pampadromaeus; as it has about 20 dentary teeth as well, and Saturnalia and Panphagia have 17 and 23 dentary teeth respectively, their maxillary counts were probably within the 12-18 range or higher too). Thus there is no shared derived state to differ from, since theropods have less teeth and sauropodomorphs have more teeth.
11. lateral surface of anterior end of nasal along the posterior margin of the external naris flat. Pantydraco and Daemonosaurus also lack this narial fossa, though Panphagia has it. This means Theropoda is basally ambiguous while Sauropodomorpha is barely basally derived in having the fossa, so the character is not an unambiguous eusaurischian synapomorphy. For what it's worth, Herrerasaurus also lacks the fossa.
12. posteroventral dentary process far posterior to posterodorsal process. This is true in basal sauropodomorphs (Panphagia, Pantydraco) and basal theropods (Eodromaeus, Tawa). It's not true in ornithischians (Tianyulong, Heterodontosaurus, Eocursor) though, making the condition in Saurischia's outgroup ambiguous (given Silesaurus having the opposite condition). Again for what it's worth, Herrerasaurus has the same condition as ornithischians.
13. foramen in the ventral part of the splenial absent. This is difficult to code as the anterior splenial is thin and often broken. In Sauropodomorpha, Panphagia has a foramen, Lamplughsaura is illustrated without one (though by Chatterjee, whose drawings are often idealized), Plateosaurus is polymorphic, and Lufengosaurus and Adeopapposaurus have it. In Theropoda, Liliensternus lacks one, Dilophosaurus is illustrated as lacking one but seems to be anteriorly incomplete, and Ceratosaurus has one. Thus the basal condition in either saurischian clade is unclear though more probably present in sauropodomorphs. Ornithischians lack the foramen, as does Staurikosaurus though the latter has a poorly preserved mandible.
14. iliac supraacetabular crest shelf-like and short, extending primarily laterally. This is also true in basal sauropodmorphs (Panphagia, Pampadromaeus) and basal theropods (Eodromaeus, Tawa).
15. ridge on lateral side of tibia for connection with fibula absent. This is untrue in Eoraptor.
Yates (2007)
16. relationship between posterolateral process of the premaxilla and the anteroventral process of the nasal a broad sutured contact. This is untrue in Eoraptor.
17. size and position of subnarial foramen small (no larger than adjacent maxillary neurovascular foramina) and positioned outside of narial fossa. Basal theropods (Tawa, coelophysids, Dilophosaurus) lack a subnarial foramen, as do outgroups (ornithischians, Silesaurus). Thus there is no obvious ancestral condition for the subnarial foramen, nor evidence theropods ancestrally had one. Herrerasaurus does have this condition.
18. pointed posterolateral process of the nasal overlapping the lacrimal absent. This is untrue in Eoraptor.
19. length of middle to posterior cervical centra (6-8) no more than the length of the axial centrum. This is unknown in Eoraptor, as the axis is fragmentary.
20. laminae bounding triangular infradiapophyseal fossae on dorsal neural arches absent. This is untrue in Eoraptor.
21. transverse width of the first distal carpal less than 120% of the transverse width of the second distal carpal. This is unknown or untrue in Eoraptor, as distal carpal I is either unpreserved or diagenetically fused to the radiale in the left carpus and ~192% the width of distal carpal II. Notably, the basal theropods Eodromaeus and Tawa and basal sauropodomorph Efraasia have a small distal carpal I though, so this is not a eusaurischian character, though Heterodontosaurus does have a large distal carpal I so that ornithischians have an ambiguous basal state. Herrerasaurus has a small distal carpal I.
Martinez and Alcober (2009)
22. no caudosacral. This is also true in basal theropods (Eodromaeus and Tawa) and ambiguous in sauropodomorphs (true in Pampadromaeus but not Efraasia and more derived taxa). Guaibasaurus, Sanjuansaurus and Herrerasaurus also lack a caudosacral, though Staurikosaurus may have one. Ornithischians have a caudosacral.
23. width of metacarpal I shaft less than 35% of length. This is untrue in Eoraptor.
Bittencourt Rodrigues (2010) also placed it basal to Eusaurischia, but this paper has yet to be translated.
Almost half (10-11) of the characters are not even being present in Eoraptor. The others are basically all also found in taxa agreed to be basal theropods (11) and/or sauropodomorphs (7). The best character is the absent anterior splenial foramen, which depends on illustration inaccuracy of a poorly preserved and seldomly exposed element.
Eoraptor a sauropodomorph? Martinez et al. (2011) used several characters to place Eoraptor as a non-plateosaurian sauropodomorph- external naris size large, expanded narial margin (incorrect- Sereno et al., 2013; also absent in Pampadromaeus); nasal posterolateral process present (true of all basal saurischians except Herrerasaurus); squamosal, ventral process a slender prong 3 or more times basal width (also in several basal ornithischians and avepods but not Tawa); dentary tooth 1 inset (also in Daemonosaurus, heterodontosaurids and silesaurids); maxillary and dentary crowns lanceolate (also in derived silesaurids and ornithischians); deltopectoral crest 45% or more of humeral length (incorrect- Sereno et al., 2013); manus phalanx I-1, rotation of axis through distal condyles 45 degrees ventromedially (incorrect- Sereno et al., 2013); preacetabular process subtriangular; M. iliotibialis 1 scar present on preacetabular process (also in herrerasaurids, Chindesaurus and basal dinosauriforms); ischial mid shaft cross-sectional shape subtriangular (also in Herrerasaurus and Heterodontosaurus, and absent in Panphagia); astragalus fibular facet primary orientation lateral (also in ornithischians and Herrerasaurus); astragalus anteromedial corner anteriorly projecting at least 25% width of the medial side of the astragalus. Additionally, Martinez et al. found Eoraptor to be closer to Panphagia and Saturnalia than plateosaurians based on- ventral acetabular flange of ilium present (primitive for dinosaurs); brevis fossa an arched ventrally-opening ovate or parallel-sided depression (a composite character also in present in Eodromaeus, Tawa and avepods, except some of the latter are further derived in having a posterior expansion); ischial antitrochanter, anteroposterior length less than adjacent length of the articular surface for the ilium (not present in Eoraptor [Sereno et al., 2013] or basal sauropodomorphs, though present in Heterodontosaurus).
Martinez et al. (2013)- length of quadrate occupied by pterygoid wing >70% (unknown- Sereno et al., 2013; though present in Eodromaeus); postparietal fenestra between supraoccipital and parietals (possible- Sereno et al., 2013); supraoccipital semilunate and wider than high (unknown- Sereno et al., 2013); serrations coarse and angled upwards at 45 degree angle (intermediate- Sereno et al., 2013, but similar to Panphagia and Pampadromaeus; absent in Saturnalia; also in derived silesaurids and ornithischians); postzygodiapophyseal lamina in cervical neural arches 4-8 absent (present but poorly developed in cervicals 7-8- Sereno et al., 2013; also in Eodromaeus and Heterodontosaurus); laminae of cervical neural arches 4-8 weakly developed low ridges or absent (includes previous character, and also the prezygodiapophyseal, anterior and posterior centrodiapophyseal laminae; also in Eodromaeus and Heterodontosaurus); minimum width of scapula <20% of its length (also in Eodromaeus, Tawa and heterodontosaurids); posterior end of fibular condyle on proximal tibia anterior to posterior margin of proximal articular surface (also in ornithischians; absent in Panphagia). They found it closer to plateosaurians than Panphagia based on- maxilla with strong inflection at base of ascending ramus (also in Eodromaeus and Tawa ; unknown in Panphagia); manual phalanx I-1 twisted ventrolaterally (more than the ~15 degrees of Herrerasaurus) (unknown in Panphagia); supraacetabular crest extended along entire pubic peduncle and contacts distal end as a well developed crest (incorrect- Sereno et al., 2013); distal outline of ischium triangular (also in Heterodontosaurus).
Eoraptor a theropod? Sereno et al. (1993)- vertebral centra extremely hollowed; cervical epipophyses prong-shaped (pointed, prominent, extend beyond postzygapophyses- Novas, 1993) (also in Heterodontosaurus); long bones extremely hollowed (also in Panphagia; Herrerasaurus has bones thicker than pterosaurs, Lagerpeton and Marasuchus- Novas, 1993; no significant difference from sauropodomorphs- Langer and Benton, 2006); manus >50% of humerus+radius length (only 41% in Eoraptor- Sereno et al., 2013; 57% in Heterodontosaurus); (asymmetrical- Sereno, 1999; and deep- Nesbitt et al., 2009) extensor pits on metacarpals I-III (also in heterodontosaurids and Plateosaurus- Langer, 2004; shallow in Eoraptor- Sereno et al., 2013); manual digit IV vestigial (mcIV <50% of mcII, less than two phalanges- Novas, 1993; much narrower than I-III- Nesbitt et al., 2009) (80% in Eoraptor- Sereno et al., 2013; also 78-84% in Eodromaeus; Tawa has two phalanges; Heterodontosaurus has a narrow mcIV); manual digit V vestigial (no phalanges- Rauhut, 2003) (absent in Eodromaeus).
Novas (1993)- trochanteric shelf (also in Chromogisaurus, Saturnalia and Pampadromaeus; absent in Tawa); humerus <50% of femoral length (56% in Eoraptor- Sereno et al., 2013; 53% in Eodromaeus; ~67% in Tawa).
Sereno (1999)- ectopterygoid fossa (absent in Eoraptor and coelophysoids- Nesbitt et al., 2011); intermandibular joint (present but of opposite convexity than herrerasaurids- Sereno et al., 2013); intermetacarpal articular facets on I-III (also in sauropodomorphs and heterodontosaurids); arched brevis fossa (also in sauropodomorphs; not in herrerasaurids- Langer, 2004); distal width of pubic blade <65% of proximal width; obturator flange present (also in sauropodomorphs- Langer, 2004).
Rauhut (2003)- manual phalanx II-2 longer than II-1 (absent- Sereno et al., 2013; though present in heterodontosaurids).
Tykoski (2005)- posterior jugal process divided (also in Pampadromaeus).
Ezcurra and Novas (2007)- rostral process of jugal excluded from the internal antorbital fenestra (absent in Daemonosaurus and Tawa; also in Heterodontosaurus); sublacrimal part of jugal squared rostrally with a very small dorsally directed prong, slightly overlapping the lacrimal (also in Heterodontosaurus; tapers in Eoraptor- Sereno et al., 2013; large dorsal expansion in Tawa); ventral process of the squamosal length less than half of the caudal border of the infratemporal fenestra (also in Heterodontosaurus); humerus length subequal or shorter than 60% length of femur (also in Tianyulong, Pampadromaeus and Saturnalia; absent in Tawa).
Nesbitt et al. (2009)- maxillary teeth, posterior edge of posterior maxillary teeth concave or straight (also in Pampadromaeus); parabasisphenoid recess present; articular, glenoid of the mandible located level with dorsal margin of the dentary (incorrect- Sereno et al., 2013); tooth crowns not mesiodistally expanded (incorrect- Sereno et al., 2013); metacarpals, proximal ends abut one another without overlapping (incorrect- Sereno et al., 2013; also absent in Eodromaeus; present in heterodontosaurids); ischium, cross-section of the distal portion rounded or elliptical (incorrect- Sereno et al., 2013; true in Panphagia); tibia, lateral condyle of proximal portion level with the medial condyle at its posterior border (incorrect- Sereno et al., 2013; also absent in Eodromaeus).
Nesbitt et al. (2010)- The matrix supposedly has maxillary interdental plates separate and anterior tympanic recess as additional theropod characters of Eoraptor, but both are unknown in the taxon and it seems some characters in the matrix were placed incorrectly.
Ezcurra and Brusatte (2011)- longitudinal ridge on jugal present and sharp (incorrect- Sereno et al., 2013; also in Pampadromaeus; absent in Tawa).
Langer et al. (2011)- Meckelian groove near dorsoventral center of dentary (unknown- Sereno et al., 2013); serrations small (intermediate- Sereno et al., 2013, as in Panphagia and Pampadromaeus; small in Saturnalia; polymorphic in heterodontosaurids); manus >40% of humerus+radius length (also in heterodontosaurids).
Eoraptor basal to Herrerasauridae+Avepoda? Sereno et al. (1993)- no intermandibular joint (incorrect- Sereno et al., 2013); distal caudal prezygapophyses not elongate (unknown- Sereno et al., 2013; though present in Eodromaeus and Tawa); scapula not strap-shaped (distal end width >33% of scapular length- Sereno, 1999) (not in coelophysoids or probably Staurikosaurus- Langer, 2004; exactly 3 times in Eoraptor- Sereno et al., 2013; strap-shaped in heterodontosaurids); short(er than proximal phalanges- Novas, 1993) penultimate phalanges on manual digits II and III (phalanges on II equal in Eoraptor- Sereno et al., 2013; untrue in heterodontosaurids); non-trenchant unguals (not enlarged, compressed, sharply pointed, recurved, with enlarged flexor tubercles- Novas, 1993; II and III slightly curved- Ezcurra and Novas, 2007) on manual digits I-III (untrue in heterodontosaurids); pubis less than six times longer than broad (untrue in ornithischians); pubic boot absent (folded in herrerasaurids, not larger than sauropodomorphs in Eodromaeus and coelophysoids); proximal end of fibula <75% proximal width of tibia.
Sereno (1999)- metacarpal IV >50% width of metacarpals II or III (only true of mcIII- Sereno et al., 2013; also true in Eodromaeus); metacarpal V >50% width of metacarpals II or III (untrue in Heterodontosaurus); anterior attachment depression absent on distal femur (also true in Eodromaeus).
Rauhut (2003)- sublacrimal part of jugal not expanded (incorrect- Sereno et al., 2013; though present in Daemonosaurus and untrue in heterodontosaurids); neural spines of posterior dorsals not significantly higher than long (also true in Eodromaeus and Tawa).
Ezcurra and Novas (2007)- height/length ratio of premaxilla below external naris 0.5-1.25; lacrimal antorbital fossa in lateral view only present in the ventral process of the bone (incorrect- Sereno et al., 2013; untrue in Heterodontosaurus); ventral process of squamosal tapering; dorsal ramus of the quadratojugal longer than the rostral ramus; mandibular joint approximately straight below the quadrate head (also true in Eodromaeus); constriction between tooth crown and root present in at least rostral maxillary teeth; caudoventral process of the coracoid rounded and not projected beyond glenoid fossa (projects beyond glenoid in Eoraptor- Sereno et al., 2013; untrue in Tianyulong); metacarpal IV with proximal portion set lateral to Mc III (untrue in Heterodontosaurus); ischial antitrochanter small (also in Tawa).
Eoraptor closer to Avepoda than Herrerasauridae? Tykoski (2005)- subnarial process of premaxilla narrow and rod-like (also in Pampadromaeus; absent in Daemonosaurus); maxilla anterodorsally concave (also in Pampadromaeus); nasal contacts antorbital fossa (absent in Daemonosaurus and Eodromaeus); lacrimal inverted L-shape (also in Pampadromaeus and Saturnalia); lacrimal at least equal to orbital height and reaches orbit's ventral rim (incorrect- Sereno et al., 2013; also absent in Daemonosaurus and Tawa); antorbital fossa exposure on lacrimal large, with triangular fossa on ventral process (also in Saturnalia and Pantydraco); vertebra 25 (dorsosacral) incorporated into sacrum (also in Heterodontosaurus); humerus twisted; metacarpal I distal condyles strongly asymmetrical (absent in Tawa; also in Heterodontosaurus); preacetabular process thin and blade-like (also in Chromogisaurus and Pantydraco); postacetabular process longer than acetabulum (also in Panphagia, Saturnalia, Chromogisaurus and Heterodontosaurus); pubic fenestra (incorrect- Sereno et al., 2013; absent in Eodromaeus and Tawa); distal ischium <3 times minimum width of shaft (also in Panphagia, Pantydraco and Heterodontosaurus); subrectangular distal tibia with posterolateral extension (incorrect- Sereno et al., 2013; though present in Panphagia, Saturnalia and ornithischians, and absent in Eodromaeus).
Ezcurra (2006)- lateral surface of premaxillary body pierced by a single neurovascular foramen above the second premaxillary tooth (a second foramen is present on one side- Sereno et al., 2013; also in Heterodontosaurus); maxillary antorbital fossa rostral to internal antorbital fenestra broad and rostrocaudally well extended (also in Pampadromaeus and Heterodontosaurus; absent in Daemonosaurus and Tawa); medial wall of the antorbital fossa extends through the entire ventral border of the internal antorbital fenestra as a very narrow lamina (also in Heterodontosaurus; absent in Tawa); lateral lamina of lacrimal with no interruption of the lacrimal antorbital fossa and restricted to the posterior margin of the ventral ramus along its dorsoventral extension (also in Heterodontosaurus); anterior process of jugal excluded from internal antorbital fenestra, bluntly squared anteriorly (tapered anteriorly- Sereno et al., 2013; also in Heterodontosaurus; contacts antorbital fenestra in Daemonosaurus and Tawa); ventral process of squamosal forms more than half posterior border of infratemporal fenestra (incorrect- Sereno et al., 2013; though present in Pampadromaeus and absent in Tawa); humerus length subequal or <60% length of femur (also in Pampadromaeus and Saturnalia; absent in Tawa); extensor pits on the dorsal surface of the distal end of metacarpals deep, well developed (shallow in Eoraptor- Sereno et al., 2013; also in Heterodontosaurus); shaft of metacarpal IV in relation to that of metacarpals I-III significantly narrower (also in Heterodontosaurus; untrue for IV vs. III in Eodromaeus); metacarpal IV and fourth digit, proximal portion set lateral to Mc III and with one or less phalanges (more ventrally set in Eodromaeus; more than one phalanx in Tawa); supraacetabular crest and lateroventral border of the postacetabular process (lateral brevis shelf) continuous as at least a weakly developed ridge (absent in Tawa); cnemial crest on proximal tibia moderately developed (also in Heterodontosaurus and Saturnalia; absent in Tawa?); posterior cleft between medial and lateral condyles on tibia (also in Heterodontosaurus, Panphagia, Saturnalia and Chromogisaurus).
Nesbitt et al. (2009)- premaxilla-maxilla, subnarial gap between the elements (incorrect- Sereno et al., 2013; also absent in Eodromaeus; present in Heterodontosaurus); nasal possesses posterolateral process that envelops part of lacrimal (true of all basal saurischians except Herrerasaurus); ilium, ischiadic peduncle orientation well expanded posteriorly to the anterior margin of the postacetabular embayment (also in Panphagia, Chromogisaurus and Pantydraco; absent in Tawa).
Ezcurra (2010)- posterior part of premaxillary alveolar margin edentulous, resulting in an interruption of the upper tooth row (incorrect, the diastema is on the maxilla- Sereno et al., 2013; also absent in Eodromaeus; present in Heterodontosaurus); anterior margin of maxillary antorbital fossa squared (also in Pampadromaeus; absent in Daemonosaurus, Eodromaeus and Tawa); dorsoventrally compressed ridge on lateral surface of maxilla, forming ventral border of antorbital fossa (also in Heterodontosaurus; absent in Daemonosaurus and Tawa); orientation of lacrimal orbital margin erect and close to vertical (incorrect- Sereno et al., 2013; also absent in Daemonosaurus and Tawa); supraacetabular crest of ilium flares lateroventrally to form hood-like overhang that hides anterodorsal half of acetabulum in lateral view (incorrect- Sereno et al., 2013; also absent in Eodromaeus and Tawa); well developed brevis fossa with sharp margins on the ventral surface of the postacetabular process of the ilium present, being directly ventrally facing (also in sauropodomorphs); well developed tibiofibular crest on distal femur (incorrect- Sereno et al., 2013; also absent in Eodromaeus).
Nesbitt et al. (2010)- As with the theropod diagnosis from this study, some characters in the matrix were seemingly placed incorrectly. Of these 'portion of maxilla anterior to anterior edge of antorbital fenestra equal or longer than posterior portion' is clearly absent in Eoraptor and meant to diagnose ornithischians as in the Tawa paper, 'lacrimal folds over posterior/posterodorsal part of antorbital fenestra' is a saurischian character meant to diagnose them as in the Tawa paper, (also in sauropodomorphs; absent in Daemonosaurus); and 'dorsal surface of frontal with longitudinal ridge along midline' is also not evident in Eoraptor and meant to diagnose Paracrocodyliformes.
Langer et al. (2011)- ilium, distal anteroposterior width of pubic peduncle (not considering the acetabular wall) more than half the total length of the peduncle (also in Pampadromaeus).
Nesbitt (2011)- trochanteric shelf absent (incorrect- Sereno et al., 2013; also wrong in Eodromaeus; true in Heterodontosaurus).
Sues et al. (2011)- external naris dorsal to a portion of maxilla (also in sauropodomorphs); posteroventral lacrimal process long and extending along dorsal edge of jugal (also in Pantydraco and some Heterodontosaurus; absent in Tawa); thin, anterolaterally expanded portion of postorbital that lies dorsal to orbit (also in Pampadromaeus and Pantydraco; absent in Tawa).
References- Novas, 1993. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina. Journal of Vertebrate Paleontology. 13, 400-423.
Padian and May, 1993. The earliest dinosaurs. Bulletin of the New Mexico Museum of Natural History and Science. 3, 379-381.
Sereno, Forster, Rogers and Monetta, 1993. Primitive dinosaur skeleton from Argentina and the early evolution of Dinosauria. Nature. 361, 64-66.
Holtz, 1995. A new phylogeny of the Theropoda. Journal of Vertebrate Paleontology. 15(3), 35A.
Gatesy and Middleton, 1997. Bipedalism, flight, and the evolution of theropod locomotor diversity. Journal of Vertebrate Paleontology. 17(2), 308-329.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences. 25, 435-489.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature of the major taxonomic categories of the carnivorous dinosaurs Dinosauria (Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Kischlat, 2000. Tecodoncios: A aurora dos Arcosaurios no Triassico. in Holz and De Rose (eds.). Paleontologia do Rio Grande do Sul. 273-316.
Middleton and Gatesy, 2000. Theropod forelimb design and evolution. Zoological Journal of the Linnean Society. 128, 149-187.
DigiMorph Staff, 2001, Eoraptor lunensis (On-line), Digital Morphology. Accessed October 21, 2010 at http://digimorph.org/specimens/Eoraptor_lunensis/.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Langer, 2004. Basal Saurischia. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Tykoski, 2005. Anatomy, ontogeny and phylogeny of coelophysoid theropods. PhD Dissertation. University of Texas at Austin. 553 pp.
Ezcurra, 2006. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas. 28(4),649-684.
Langer and Benton, 2006. Early dinosaurs: A phylogenetic study. Journal of Systematic Palaeontology. 4(4), 309-358.
Ezcurra and Novas, 2007. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.
Sereno, 2007. The phylogenetic relationships of early dinosaurs: A comparative report. Historical Biology. 19(1), 145-155.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Yates, 2007. Solving a dinosaurian puzzle: The identity of Aliwalia rex Galton. Historical Biology. 19(1), 93-123.
Martinez and Alcober, 2009. A basal sauropodomorph (Dinosauria: Saurischia) from the Ischigualasto Formation (Triassic, Carnian) and the early evolution of Sauropodomorpha. PLoS ONE. 4(2), e4397.
Nesbitt, Smith, Irmis, Turner, Downs and Norell, 2009. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science. 326, 1530-1533.
Bittencourt Rodrigues, 2010. Revisao filogenetica dos dinossauriformes basais: Implicacoes para a origem dod dinossauros. Unpublished Doctoral Thesis. Universidade de Sao Paulo. 288 pp.
Ezcurra, 2010. A new early dinosaur (Saurischia: Sauropodomorpha) from the Late Triassic of Argentina: A reassessment of dinosaur origin and phylogeny. Journal of Systematic Palaeontology. 8(3), 371-425.
Martinez, Sereno, Alcober, Columbi, Renne, Montanez and Currie, 2011. A basal dinosaur from the dawn of the dinosaur era in Southwestern Pangaea. Science. 331, 206-210.
Martinez, Apaldetti and Abelin, 2013. Basal sauropodomorphs from the Ischigualasto Formation. Journal of Vertebrate Paleontology. 32(Supplement to 6), 51-69.
Sereno, Martinez and Alcober, 2013. Osteology of Eoraptor lunensis (Dinosauria, Sauropodomorpha). Journal of Vertebrate Paleontology. 32(Supplement to 6), 83-179.

undescribed saurischian (Marsola, Bittencourt, Da-Rosa and Langer, 2015)
Carnian, Late Triassic
Alemoa Member of the Santa Maria Formation, Brazil

Material- (LPRP/USP 0651) partial posterior dorsal vertebrae, incomplete sacral vertebrae, caudal vertebrae, partial chevron, incomplete ilium, femur, partial tibiae, fibulae, metatarsals II, metatarsals IV, pedal phalanges, pedal unguals
Comments- Differs from Saturnalia in- strong ventral median keel on proximal caudal centra; posteriorly oriented ischiadic peduncle; concave ventral margin of iliac acetabulum; ilium proportionally smaller than femur; slender hind limb; poorly developed trochanteric shelf; distal tibia mediolaterally expanded; fibula longer than femur. Follows Eoraptor in analyses, so may be related.
Reference- Marsola, Bittencourt, Da-Rosa and Langer, 2015. A small-sized saurischian dinosaur from the Late Triassic Santa Maria Formation, southern Brazil. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 175.

Teyuwasu Kischlat, 1999 vide Kischlat, 2000
= "Teyuwasu" Kischlat, 1999
T. barberenai Kischlat, 1999 vide Kischlat, 2000
= "Teyuwasu barberenai" Kischlat, 1999
Late Carnian-Early Norian, Late Triassic
Alemoa Member of Santa Maria Formation, Brazil

Holotype- (BSPG AS XXV 53) femur (276 mm)
....(BSPG AS XXV 54) tibia (264 mm)
?...(BSPG AS XXV 56-59) dorsal centrum (40 mm), partial ilium, (?) distal ischium, femur
Diagnosis- (proposed) extremely robust femur and tibia (minimum transverse femoral width 19% of length); mound-like fourth trochanter.
Comments- The material was originally described by Huene (1938) as possibly belonging to his new taxon Hoplitosaurus raui, from slightly higher in the formation. Huene later (1942) renamed it Hoplitosuchus, as Hoplitosaurus was preoccupied by an ankylosaur. The taxon was based on two supposed osteoderms described as aetosaurian, but more recently these have been found to be unidentifiable bones by Kischlat (2000) and Desojo and Rauhut (2008). Kischlat (1999) reinterpreted the femur and tibia as being dinosaurian, though only listed features identifying it to the level of Dinosauriformes. He provided a very brief description and named the taxon Teyuwasu barberenai, but as the publication is a symposium abstract, it is invalid under the ICZN (Article 9.10). Kischlat's (2000) later article has similar information, credits the name to the 1999 paper, but is a valid publication. Ezcurra (2012) described the material in depth, finding it certainly belongs to the silesaurid+dinosaur clade, but that the only dinosaurian character is the inturned femoral head. As the bones have been heavily altered taphonomically, Ezcurra was uncertain if the femoral head orientation was artificial. He notes Kischlat's "two paralell ridges running proximodistally" are fractures formed when the medial femoral head was sheared distally. While Ezcurra declared Teyuwasu to be indeterminate, the robusticty itself is vastly different from other basal dinosauriforms, so should be enough to validate the taxon. If added to the Nesbitt archosaur supermatrix, Teyuwasu emerges as a saurischian based on femur longer or about the same length as the tibia, and medial articular facet of the proximal femur rounded. It's outside Eusaurischia based on the femoral head being unexpanded, symmetrical fourth trochanter (considered tentative by Ezcurra), cnemial crest not laterally curved, and posterior face of distal tibia without longitudinal ridge. Some of these characters vary within basal saurischians, which are incompletely sampled by Nesbitt, so could change polarity in the future.
Besides the femur and tibia, Huene referred additional material to this individual. A centrum identified by Kischlat (2000) as dorsal was stated by Huene to be possibly but not certainly referrable to this specimen. It is 40 mm long, 45 mm tall and wide, rounded in section and barely amphicoelous to amphiplatyan. A ventral ilium was considered very likely to belong to this individual. It has a supracetabular crest and 'strongly recessed' acetabulum, and narrows to 70 mm between the peduncles and blade. An element tentatively identified by Huene as a distal ischium is much too large to belong to this individual, with the distal end 90 mm deep and 60 mm wide. At the proximal break, these dimensions are 53 and 30 mm respectively. The distal end is triangular in section, which is a saurischian character. Finally, Kischlat (2000) mentioned an additional femur which was not noted by Huene. These were all noted by Kischlat as supplementary material for Teyuwasu, though Desojo and Rauhut stated referred Hoplitosuchus material belongs to Rauisuchia and Dinosauria. The centrum, ilium and/or ischium may comprise the 'rauisuchian' material, or this may refer to the two non-avemetatarsalian calcanea also referred to Hoplitosuchus by Huene.
References- Huene, 1938. Ein grosser Stagonolepid aus der jungeren Trias Ostafrikas. Neues Jahrbuch fur Mineralogie, Geologie und Palaontologie. 80(2), 264-278.
Huene, 1942. Die fossilen Reptilien des sudamerikanischen Gondwanalandes. Ergebnisse der Sauriergrabungen in Sudbrasilien 1928/29. Munich: Becksche Verlegbuchhandlung. 332 pp.
Kischlat, 1999. A new dinosaurian "rescued" from the Brazilian Triassic: Teyuwasu barbarenai, new taxon. Paleontologia em Destaque, Boletim Informativo da Sociedade Brasileira de Paleontologia. 14(26), 58.
Kischlat, 2000. Tecodoncios: A aurora dos Arcosaurios no Triassico. in Holz and De Rose (eds.). Paleontologia do Rio Grande do Sol. 273-316.
Desojo and Rauhut, 2008. New insights on "rauisuchian" taxa (Archosauria: Crurotarsi) from Brazil. SVPCA 2008 Programme and Abstracts. 18-19.
Ezcurra, 2012. Comments on the taxonomic diversity and paleobiogeography of the earliest known dinosaur assemblages (Late Carnian-Earliest Norian). Historia Natural. 2(1), 49-71.

undescribed possible saurischian (Reig, 1963)
Late Carnian-Early Norian, Late Triassic
Cancha de Bochas Member of the Ischigualasto Formation, San Juan, Argentina

Material- (PVL 2469) tibia (440 mm)
Comments- This was stated to perhaps belong to a new genus of saurischian by Reig (1963), as it did not match Herrerasaurus in morphology. As several additional taxa have since been named from that formation (e.g. Panphagia, Eoraptor, Sanjuansaurus), it may belong to one of those.
Reference- Reig, 1963. La presencia de dinosaurios saurisquios en los "Estratos de Ischigualasto" (Mesotriasico Superior) de las provincias de San Juan y La Rioja (República Argentina) [The presence of saurischian dinosaurs in the "Ischigualasto beds" (upper Middle Triassic) of San Juan and La Rioja Provinces (Argentine Republic)]. Ameghiniana. 3, 3-20.

unnamed saurischian (Galton, 1985)
Norian, Late Triassic
Lowenstein Formation, Germany
Material
- (SMNS 51958) proximal femur
Comments- Galton (1985) referred this specimen to the same unnamed family as Aliwalia, within Herrerasauria. This was followed by Paul (1988), who placed both in Herrerasauridae. However, Yates (2006) noted Aliwalia is a junior synonym of Eucnemesaurus, which is a sauropodomorph and differs from SMNS 51958 in several respects. Namely, the latter has a more proximally placed fourth trochanter which lacks a rounded profile and notched distal end. Also, the anterior trochanter does not form a proximodistally elongate ridge, unlike sauropodomorphs more derived than Saturnalia. The lack of a proximal notch separating the anterior trochanter and femoral shaft is unlike ornithischians, and most avepods except robust coelophysoids and robust ceratosaurs. The symmetrical fourth trochanter is unlike sauropodomorphs, Herrerasaurus and Eoraptor. The markedly inturned femoral head indicates SMNS 51958 is a dinosaur, more derived than Silesaurus or Lewisuchus.
References- Galton, 1985. The poposaurid thecodontian Teratosaurus suevicus v. Meyer, plus referred specimens mostly based on prosauropod dinosaurs, from the Middle Stubensandstein (Upper Triassic) of Nordwurttemberg. Stuttgart Beitrage zur Naturkunde (B). 116, 1-29.
Paul, 1988. Predatory dinosaurs of the world. Simon and Schuster, New York. A New York Academy of Sciences Book. 464 pp.
Yates, 2007. Solving a dinosaurian puzzle: the identity of Aliwalia rex Galton. Historical Biology. 19(1), 93-123.

undescribed saurischian (Murry and Long, 1989)
Early Norian, Late Triassic
Mesa Redondo Member of the Chinle Formation, Arizona, US

Material- (UCMP 177316; paratype of Chindesaurus bryansmalli) five dorsal centra
Comments- Murry and Long (1989) mention Chindesaurus remains from the Placerias quarry, which were specified as five dorsal centra in the UCMP by Long and Murry (1995) in their description of the taxon as a herrerasaurid. The latter publication figured them as UCMP A269, but this is the number of the Placerias quarry locality, not the specimen itself. A search through the UCMP collection database suggests they are UCMP 177316, five vertebrae now catalogued as Dinosauria and supposed to be figured. Hunt et al. (1998) stated they were not generically determinate, but retained their assignment to Herrerasauridae. As Chindesaurus now has a less certain position within Saurischia, perhaps being closer to avepods than herrerasaurids, these dorsals are provisionally assigned to Saurischia indet..
References- Murry and Long, 1989. Geology and paleontology of the Chinle Formation, Petrified Forest National Park and vicinity, Arizona and a discussion of vertebrate fossils of the southwestern Upper Triassic. in Lucas and Hunt (eds). Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. 29-64.
Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Natural History Science Bulletin. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the Western United States. Geobios. 31(4), 511-531.

undescribed possible Saurischia (Kirby, 1991)
Rhaetian, Late Triassic
Owl Rock Member of the Chinle Formation, Arizona, US
Material
- (MNA V7233) tooth
(MNA V7234) tooth
(MNA V7235) tooth
(MNA V7236) tooth
(MNA V7237) astragalus
(MNA V7238) ungual
(MNA V7239) ungual
(MNA V7241) centrum
Comments- Kirby (1991, 1993) assigned these specimens to Theropoda, and Hunt et al. (1998) agreed the astragalus was dinosaurian. None of the specimens are mentioned in Spielmann et al.'s (2007) latest review of Kirby's material. While the astragalus could prove to be distinctive to a subgroup of dinosaurs, theropod teeth, centra and unguals can often be indistinguishable from other archosaurs.
References- Kirby, 1991. A vertebrate fauna from the Upper Triassic Owl Rock Member of the Chinle Formation of northern Arizona. Unpublished Masters thesis. Northern Arizona University, Flagstaff. 496 pp.
Kirby, 1993. Relationships of Late Triassic basin evolution and faunal replacement in the southwestern United States: Perspectives from the upper part of the Chinle Formation in northern Arizona. In Lucas and Morales (eds.). The Nonmarine Triassic. New Mexico Museum of Natural History and Science Bulletin. 3, 233-242.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the Western United States. Geobios. 31(4), 511-531.
Spielmann, Lucas and Heckert, 2007. Tetrapod fauna of the Upper Triassic (Revueltian) Owl Rock Formation, Chinle Group, Arizona. In Lucas and Spielmann (eds.). The Global Triassic, New Mexico Museum of Natural History and Science Bulletin. 41, 371-383.

unnamed saurischian (Raath, Oesterlen and Kitching, 1992)
Early Carnian, Late Triassic
Pebbly Arkose Formation, Zimbabwe
Referred
- (253/3(7)) proximal femur
Comments- Raath et al. (1992) and Raath (1996) originally called this a prosauropod based on the crest-like fourth trochanter. Langer et al. (1999) considered it almost indistinguishable from Saturnalia and possibly congeneric. Langer (2001) elaborated, believing the trochanteric shelf and semi-pendant fourth trochanter to be found in only Saturnalia and Herrerasaurus, but referred it to the former based on size and the presence of a nutrient foramen. Langer nonsensically describes this foramen as "caudal to the fourth trochanter" in both, but Raath illustrates and describes the foramen to be opposite the fourth trochanter on the anterior surface in the Zimbabwean specimen, while Langer shows the foramen in Saturnalia to be just lateral to the distal edge of the fourth trochanter. Thus they are in different positions. Most recently, Ezcurra (2012) stated it could only be identified as an indeterminate saurischian, saying it not only fell into the morphological range of Saturnalia and Herrerasaurus, but also Staurikosaurus. However, the latter differs in lacking a trochanteric shelf. Since Langer (2001), other basal saurischians have been descfribed with the same combination of features- Sanjuansaurus, Pampadromaeus, Alwalkeria and Eoraptor. The latter three are all similar in size and at least Eoraptor seems to have the anteriorly placed nutrient foramen (PVSJ 559). Ezcurra's position is thus followed and the Zimbabwean femur is referred to Saurischia indet..
References- Raath, Oesterlen and Kitching, 1992. The first record of Triassic Rhynchosauria (Reptilia: Diapsida) from the Lower Zambezi Valley, Zimbabwe. Palaeontologia Africana. 29, 1-10.
Raath, 1996. Earliest evidence of dinosaurs from Central Gondwana. Memoirs of the Queensland Museum. 39, 703-709.
Langer, Abdala, Richter and Benton, 1999. A sauropodomorph dinosaur from the Upper Triassic (Carnian) of Southern Brazil. Comptes Rendus de l'Academie des Sciences, Paris, Sciences de la Terre et des Planetes. 329, 511-517.
Langer, 2001. Saturnalia tupiniquim and the early evolution of dinosaurs. PhD thesis, University of Bristol. 371 pp.
Ezcurra, 2012. Comments on the taxonomic diversity and paleobiogeography of the earliest known dinosaur assemblages (Late Carnian-Earliest Norian). Historia Natural. 2(1), 49-71.

Saurischia indet. (Long and Murray, 1995)
Late Carnian, Late Triassic
Mesa Redondo Formation of the Chinle Group, Arizona, US

Material- (UCMP 139662) proximal femur
Comments- Originally identified as Ceratosauria indet. by Long and Murray (1995), then made a paratype of Camposaurus by Hunt et al. (1998), this was relegated to Saurischia indet. by Nesbitt et al. (2007). It shows an offset head with a ventral sulcus, a moderately developed trochanteric shelf of the anterior trochanter and a trochanteric fossa.
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Nat. History Sci. Bull. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the western United States. Geobios, 31(4):511-531.
Nesbitt, Irmis and Parker, 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology. 5(2), 209–243.

undescribed Saurischia (Long and Murry, 1995)
Late Norian, Late Triassic
Petrified Forest Member of Chinle Formation, Arizona, US

Material- (PEFO 4849; paratype of Chindesaurus bryansmalli) dorsal centrum (Long and Murry, 1995)
caudal vertebra, unguals (Watts and Heckert, 2008)
Comments- PEFO 4849 was originally referred to Chindesaurus by Long and Murry (1995) in their description of the taxon as a herrerasaurid, but removed by Hunt et al. (1998) as Herrerasauridae indet.. As Chindesaurus now has a less certain position within Saurischia, perhaps being closer to avepods than herrerasaurids, these dorsals are provisionally assigned to Saurischia indet..
Watts and Heckert (2008) report theropod material from this member.
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Natural History Science Bulletin. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the Western United States. Geobios. 31(4), 511-531.
Watts and Heckert, 2008. Reevaluation of small vertebrate fossils from the Upper Triassic Chinle Group, Southeastern Utah. Journal of Vertebrate Paleontology. 28(3), 158A.

Saurischia indet. (Hunt, Lucas, Heckert, Sullivan and Lockley, 1998)
Norian, Late Triassic
Bull Canyon Formation, New Mexico, US
Material
- (NMMNH P-22494) (Hunt, Lucas, Heckert, Sullivan and Lockley, 1998)
Comments- NMMNH P-22494 was supposedly originally referred to Chindesaurus by Long and Murry (1995) in their description of the taxon as a herrerasaurid, but removed by Hunt et al. (1998) and assigned to Herrerasauridae indet.. However, this is not listed as a referred Chindesaurus specimen in the former paper. As Chindesaurus now has a less certain position within Saurischia, perhaps being closer to avepods than herrerasaurids, this material is provisionally assigned to Saurischia indet..
References- Long and Murry, 1995. Late Triassic (Carnian and Norian) tetrapods from the Southwestern Unites States. New Mexico Museum Natural History Science Bulletin. 4, 1-254.
Hunt, Lucas, Heckert, Sullivan and Lockley, 1998. Late Triassic dinosaurs from the Western United States. Geobios. 31(4), 511-531.

unnamed saurischian (Lehane, 2005)
Norian, Late Triassic
Bull Canyon Formation of the Dockum Group, Texas, US

Material- (TTUP 10072) several dorsal vertebral fragments, several mid and distal caudal vertebrae, chevron fragments, ilial fragments, proximal pubis, proximal femur, distal tibia, incomplete astragalus
Comments- While originally identified as Shuvosaurus (Lehane, 2005) or Coelophysis (Lehman and Chatterjee, 2008), this specimen was described as a new taxon of theropod closer to Avepoda than Herrerasaurus or Eoraptor by Nesbitt and Chatterjee (2008). As the latter placement was based on the highly miscoded Smith et al. matrix with very few non-avepod OTUs, I added it to an expanded version of Nesbitt's archosauriform matrix and found it to most parsimoniously be a saturnaliine sauropodomorph, but can move in a position equivalent to what Nesbitt and Chatterjee found (sister to Tawa) with one more step, and into Coelophysoidea with two more steps. Because of this, it is here placed as Saurischia incertae sedis pending further analysis.
References- Lehane, 2005. Anatomy and relationships of Shuvosaurus, a basal theropod from the Triassic of Texas. Masters thesis, Texas Tech University. 92 pp.
Lehman and Chatterjee, 2005. Depositional setting and vertebrate biostratigraphy of the Triassic Dockum Group of Texas. Journal of Earth System Science. 114(3), 325-351.
Nesbitt and Chatterjee, 2008. Late Triassic dinosauriforms from the Post Quarry and surrounding areas, west Texas, U.S.A. Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen. 249(2), 143-156.

unnamed saurischian (Ezcurra and Novas, 2007)
Late Norian-Rhaetian, Late Triassic
Upper Los Colorados Formation, La Rioja, Argentina

Material- (PULR coll.) partial ilium, proximal pubis
Comments- This was associated with the holotype of Zupaysaurus (Arcucci and Coria, 2003), but cannot be referred to that taxon due to several characters (ventrally projected articular facet of pubic peduncle; supraacetabular crest laterally projected and not extended over the ischial peduncle; supraacetabular crest well separated from the lateral magin of the brevis fossa; articular facet of the ischial peduncle reduced). Ezcurra and Novas (2007) considered it to be a non-neotheropod saurischian. The other material originally stated by Arcucci and Coria to be associated with but not referrable to Zupaysaurus (proximal scapulocoracoid, distal femora, proximal tibia) was later referred to that taxon by Ezcurra and Novas.
References- Arcucci and Coria, 2003. A new Triassic carnivorous dinosaur from Argentina. Ameghiniana. 40(2), 217-228.
Ezcurra and Novas, 2007. Phylogenetic relationships of the Triassic theropod Zupaysaurus rougieri from NW Argentina. Historical Biology. 19(1), 35-72.

Eusaurischia Padian, Hutchinson and Holtz, 1999
Definition- (Cetiosaurus medius + Passer domesticus) (modified from Langer, 2004)

Sauropodomorpha

Theropoda