Orionides Carrano, Benson and Sampson, 2012
Definition- (Megalosaurus bucklandii + Allosaurus fragilis + Passer domesticus) (Hendrickx, Hartman and Mateus, 2015; modified from Carrano, Benson and Sampson, 2012)
x= Carnosauria sensu Rauhut and Pol, 2019
Definition- (Megalosaurus bucklandii, Allosaurus fragilis <- Passer domesticus) (modified)
Comments- A clade containing megalosauroids and avetheropods to the exclusion of basal tetanurines such as Monolophosaurus, Chuandongocoelurus, Piatnitzkysaurus, Condorraptor, "Szechuanoraptor" (now Yangchuanosaurus zigongensis) and/or Xuanhanosaurus has been recognized by most recent analyses (e.g. Rauhut, 2003; Holtz et al., 2004; Smith et al., 2007; Carrano et al., 2012). Carrano et al. finally named it Orionides in 2012.
References- Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs. Scientific Reports. 9:18826.

= Bahariasauridae Huene, 1948
Bahariasaurus Stromer, 1934
B. ingens Stromer, 1934
Early Cenomanian, Late Cretaceous
Baharija Formation, Egypt
Syntypes- (IPHG 1922 X 47; destroyed) (~11.9 m; ~2.5 tons) posterior dorsal vertebra (200 mm), thirteenth dorsal vertebra (~180 mm), dorsal neural arch, dorsal rib fragment, first sacral vertebra (~135 mm), second sacral vertebra (~120 mm), third sacral vertebra (~120 mm), pubes (1.03 m), proximal ischium
?(IPHG 1922 X 48a; destroyed) incomplete posterior cervical vertebra (~125 mm)
(IPHG 1922 X 48b; destroyed) partial posterior dorsal vertebra (~170 mm)
(IPHG 1922 X 48c; mislabeled as 48d in plate 2 of Stromer, 1934; destroyed) incomplete posterior dorsal vertebra (180 mm)
(IPHG 1922 X 48 B; destroyed) pubis (~>760 mm)
Paratypes- ?(IPHG 1911 XII 23; destroyed) (juvenile?) proximal ischium
(IPHG 1912 VIII 62; destroyed) partial ilium, proximal pubis
(IPHG 1912 VIII 62a; destroyed) partial posterior dorsal centrum (220 mm)
....(IPHG 1912 VIII 62b; destroyed) incomplete posterior dorsal vertebra (225 mm)
....(IPHG 1912 VIII 62c; destroyed) posterior dorsal centrum (225 mm)
....(IPHG 1912 VIII 62d; destroyed) proximal caudal centrum (155 mm)
....(IPHG 1912 VIII 62e; destroyed) ?first dorsal neural arch
....(IPHG 1912 VIII 62f; destroyed) mid caudal vertebra (170 mm)
(IPHG 1912 VIII 74; destroyed) proximal ischium
(IPHG 1912 VIII 83a; destroyed) (juvenile?) mid caudal vertebra (73 mm)
....(IPHG 1912 VIII 83b; destroyed) mid caudal vertebra (70 mm)
Albian-Early Cenomanian, Early Cretaceous-Late Cretaceous
Tegama Group, Niger
Referred- ?(MNHN coll.; from Iguallala) (multiple individuals) proximal caudal centrum (65 mm), two mid caudal centra (60, 55 mm) (Lapparent, 1960)
?(MNHN coll.; from In Abangarit) (multiple individuals) three mid caudal centra (60, 55, 50 mm) (Lapparent, 1960)
Diagnosis- (proposed) distal gap between pubes longer than transverse width of pubis. Combination of elongate dorsal centra (posterior dorsal centra >150% longer than tall); pleurocoels in first through third sacral vertebrae; ventral median groove in first through third sacral centra; pubis lacks an obturator foramen; interpubic foramen short, about equal to pubic width; pubic boot short (~19% of pubic length); ischium has an obturator notch.
Comments- Stromer (1934) specifies not only IPHG 1922 X 47 as the type in his text, but also IPHG 1922 X 48. These then become syntypes for the species, though in the plate captions he only indicates 47 is the type. The syntypes were discovered in 1922, though the referred material is from 1911 and 1912. As with all of Stromer's Baharija material, the fossils themselves were destroyed in 1944. Rauhut (2003) claimed no diagnostic characters are apparent in Stromer's figures, so declared Bahariasaurus a nomen dubium. Yet it can be distinguished from all other taxa. The holotype possesses sacral pleurocoels, which are known in very few theropods. "Brontoraptor" (=Torvosaurus?) has them in sacral one, but differs in having no ventral sacral groove, a more robust pubis with obturator foramen, long interpubic foramen and little distal separation of the pubes, and no obturator notch in the ischium. Mapusaurus has pleurocoels in the first sacral as well, but the centra are tall, the ischium has a posterodorsal groove and the obturator process is low. The same can be said for Acrocanthosaurus, which also differs in its robust pubis with long boot and elongate interpubic foramen. Tyrannosaurids sometimes have anterior sacral pleurocoels, but differ in having tall centra, no ventral sacral groove, an elongate interpubic foramen, little distal separation of the pubes, a long pubic boot, and proximodorsal ischial process. Aerosteon has pneumatic sacral centra, but has tall centra, an elongate interpubic foramen, little distal separation of the pubes, and a long pubic boot. Other theropods with sacral pleurocoels are derived coelurosaurs with distally placed obturator processes. Chure (2000) claimed Bahariasaurus was a nomen nudum, but it was clearly established within the guidelines of the ICZN, so is valid.
Description- This refers only to the apparent holotype IPHG 1922 X 47. The two posterior dorsal centra are ~157% and 189% longer than tall and ~82% and ~95% as wide as tall, although both are crushed. Articular ends are very weakly opisthocoelous, so that Stromer described them as basically platycoelous. The dorsal rib fragment has a lenticular distal section and cresentric proximal section due to a developing medial ridge and is stated to be thinner than Spinosaurus'. Only the first three sacrals are preserved, which are fused and platycoelous with the fourth sacral unfused to them. Sacral centra are very elongate, with centra 1 and 3 being 193% and 200% longer than tall respectively, but about as wide as long with values of 108% and 89%. Carrano and Sampson (2008) said Stromer reported that both Bahariasaurus and Ceratosaurus "shared an unusual constriction of the vertebral centra", but Stromer actually wrote it was the elongation that was shared. Sacrals two and three have minimum central widths of 83% and 86% of sacral one. All preserved sacrals have a longitudinally elongate pleurocoel and a ventral median groove. The pubes are preserved lacking the proximal edges. The shaft is stated to have little curvature in lateral view and has a symphysis extending ~65% of its length. Distal to this there is an interpubic fenestra and the distal ends are separated as well. The pubic boot is stated to be small, and is posteriorly transversely narrow and more proximally placed than the distal pubic end. While the proximal ends are incomplete, enough of the posterior margin is preserved to show there were no obturator or pubic fenestrae. A laterally projected pubic tubercle is placed closer to the proximal end than to the symphysis, and the shaft proximal to the symphysis is anterolaterally-posteromedially compressed with the transverse axis 85% the diameter of the sagittal axis. Distally, the pubes show no obvious expansion or compression at the level of the boot. The ischium has no proximodorsal process but does have a low mid dorsal process slightly distal to the obturator process. While incomplete preserved, there is a proximally placed obturator process. Stromer states there is a deep pit in the ilial peduncle. The lateral surface dorsal to the obturator process is strongly concave, forming a fossa whose edge which continues distally as a strong ridge in the ventral half of the shaft.
Referred specimens- Stromer (1934) referred several specimens to Bahariasaurus. The posterior cervical IPHG 1922 X 48a is strongly opisthocoelous, lacks a ventral keel and has single pleurocoels which are not axially elongate. The parapophysis is placed directly ventral to the pleurocoel, and the anterior articular ball lacks the peripheral band of megalosauroids. The posterior dorsals IPHG 1922 X 48b and c seem too large to belong to the same individual as 48a, and have single pleurocoels and no obvious parapophyses. 48c has a neural spine without basal webbing or an I-beam section, and lacks accessory centrodiapophyseal laminae. There does seem to be a weak posterodorsal ridge lateral to the hyposphene. The pubis IPHG 1922 X 48 B is basically straight and similar to the holotype in lacking proximal fenestrae, though the ischial peduncle is slightly curved distally. The distal boot is broken posteriorly but has almost no anterior extension. Stromer feels these vertebrae "obviously" belong to Bahariasaurus and states the pubis is "very similar". Proximal ischium IPHG 1912 VIII 74 shows slight differences from the holotype which Stromer thought might be explained by sexual dimorphism but which are comparable to individual variation in other theropods. Stromer states posterior dorsals IPHG 1912 VIII 62a-c only differ from the holotype in being posteriorly broader and larger, which he attributes sensibly to positional and ontogenetic variation. Along with these, caudals IPHG 1912 VIII 62d-f were all found in the same layer and are of the right size to derive from one individual. Proximal caudal centrum IPHG 1912 VIII 62d is amphicoelous with a pair of pleurocoels and ventral groove. Stromer distinguishes it from Carcharodontosaurus based on the ventral groove, but this is known to vary within the tail of other theropods. The supposed proximal caudal neural arch IPHG 1912 VIII 62e is very unlike theropod caudals, but similar to the tenth cervical of e.g. Neovenator and Allosaurus in the broad diamond-shaped postzygapophyseal-spine region and broadly triangular diapophyses. The neural spine is very low. Caudal vertebra IPHG 1912 VIII 62f has central proportions similar to the 7th of Majungasaurus and 20th of Allosaurus, but has an extremely reduced neural spine compared to those taxa and no accessory anterior spine. The amphicoelous centrum lacks pleurocoels or a ventral groove. The partial ilium IPHG 1912 VIII 62 is not illustrated and the description lacks useful details. A proximal pubis is given the same number, though there is no mention of association. It's very similar to IPHG 1922 X 48 B and no doubt belonged to the same taxon. IPHG 1912 VIII 83a-b were referred to Bahariasaurus based on the similarity to IPHG 1912 VIII 62f, though considered to be juvenile. Small proximal ischium IPHG 1911 XII 23 was referred to juvenile Bahariasaurus as well, and is difficult to compare though its ilial peduncle is longer than the holotype or IPHG 1912 VIII 74. The pubis IPHG 1912 VIII 81 was referred to Bahariasaurus by Stromer and stated to be "generally very similar to it", but differs several ways. Bahariasaurus has a less conspicuous and more proximally placed lateral flaring (15% down the shaft, compared to 21%). The distal end is not flared laterally. There is an extensive separation of the pubic shafts distally, and the interpubic foramen is more distally placed (80% down the shaft, vs. 71%). Chure (2000) also questioned the referral of this pubis due to its sinuous curve in lateral view. There are several specimens Stromer questionably refers to the genus as well. IPHG 1912 VIII 60 (also 60a-h) is an associated skull fragment, eight caudal vertebrae, centrum and scapulocoracoid, of which the caudals may belong but the scapulocoracoid seems spinosaurid. IPHG 1912 VIII 69 is a large femur and IPHG 1912 VIII 70 is a large fibula that is avetheropodan or ceratosaurian. Neither can be compared to Bahariasaurus but are similar to Deltadromeus, the femur sharing a couple apomorphies with that genus.
Lapparent (1960) referred additional remains to this taxon. The centrum from In Abangarit seems to have a pleurocoel, but could equally belong to Carcharodontosaurus. Three caudal centra from Iguallala have no particular characters assigning them to any theropod clade.
Rauhut (1995) mentioned cf. Bahariasaurus vertebrae which were later described as Carcharodontosauridae indet. (Vb-607) by Rauhut (1999). This was compared to IPHG 1912 VIII 62d, which is referred to Bahariasaurus without good reason to not refer it to Carcharodontosaurus.
Relationships- Rauhut (1995) referred Bahariasaurus to the Carnosauria (his Allosauroidea) based on a few characters. The absence of a distal pubic symphysis in Bahariasaurus was conflated with the interpubic fenestra in Allosaurus and Sinraptor, whereas all tetanurines except some derived coelurosaurs have interpubic fenestrae and Bahariasaurus has an apomorphically long distal symphyseal gap. The absence of a pubic obturator foramen is found in allosaurian carnosaurs, but also in Marshosaurus, Eustreptospondylus, spinosaurids and most coelurosaurs among tetanurines. The form of the anterior trochanter is based on referred avetheropod specimen IPHG 1912 VIII 69 which may be a carnosaur, but is not necessarily Bahariasaurus. He placed the genus in Carcharodontosauridae based on the presence of caudal pleurocoels, but these are also known in Torvosaurus and megaraptorans among non-maniraptorans. Thus there is no evidence Bahariasaurus is a carnosaur.
Paul (1988) placed Bahariasaurus in his Allosauridae paraphyletic to Tyrannosauridae, as the genus closest to the latter family. This was based on the "classic allosaur" femur IPHG 1912 VIII 69 which is questionably referred, the referred fibula IPHG 1912 VIII 70 "of the tyrannosaur mold", and the supposedly "very narrow" pubes like tyrannosaurs. Yet the pubes of Monolophosaurus, Sinraptor and various coelurosaurs are equally narrow. A similar position was suggested by Kurzanov (1989), on the basis of a few characters. Non-opisthocoelous ('amphicoelous') dorsals are only known posteriorly in Bahariasaurus, as in most theropods. The supposedly centrally positioned neural processes are not placed differently than Allosaurus. The "drastic widening of the acromial region of the scapula" is based on referred probable spinosaurid specimen IPHG 1912 VIII 60, and is not evident from that specimen due to breakage. Molnar et al. (1990) repeated these, stating amphicoelous anterior dorsals were a character, but this region is unknown in Bahariasaurus. Similarly, the second character was restated as centrally (instead of distally) placed neural spines in distal caudals, when none are known from Bahariasaurus. The anteriorly oriented fibular tubercle is plesiomorphic for theropods. Chure (2000) repeated these characters, but also claimed Bahariasaurus had a triangular obturator process when the shape is unpreserved. Chure also proposed an extremely narrow scapular blade (based on referred probable spinosaurid specimen IPHG 1912 VIII 60) and a longitudinal ridge on the lateral ischium (in the type and IPHG 1912 VIII 74). This ridge is seemingly the one over the obturator region and at least somewhat more distally. Tyrannosaurus does have a version of this over the distal obturator area, which is also found in Guanlong, Juratyrant, Aviatyrannis and Yutyrannus. Yet this is also found in Torvosaurus, Neovenator, giganotosaurines, Huaxiagnathus and Ornitholestes so is not a tyrannosauroid character. This leads to no characters suggesting tyrannosauroid affinities.
Holtz et al. (2004) referred Bahariasaurus to Avetheropoda based on the obturator process, but this is also found in Eustreptospondylus and Afrovenator. The obturator notch in the pubis was said to be like allosaurians and coelurosaurs, but as noted above this is also present in Marshosaurus, Eustreptospondylus and spinosaurids. The interpubic foramen was also said to be like these clades, but this is present in all tetanurines except some derived coelurosaurs. Within Tetanurae, no characters of Bahariasaurus are unique to avetheropods.
Sereno et al. (1996) compared Bahariasaurus to their new taxon Deltadromeus, which has led to discussion of synonymy. The referred Bahariasaurus coracoid IPHG 1912 VIII 60, pubes IPHG 1912 VIII 81, femur IPHG 1912 VIII 69 and fibula IPHG 1912 VIII 70 were all referred to Deltadromeus by Sereno et al., which has also led to the comparison. Sereno et al. (1996) distinguished the genera using three characters, two of which were due to his misidentification of Deltadromeus' distal ischium as a pubis. The other is the narrower ilial peduncle of Bahariasaurus' ischium. The holotype of Tyrannosaurus has a narrower ilial peduncle on the ischium than referred specimens AMNH 5027 and RTMP 81.61, so this character falls within individual variation. While this has been viewed as possible evidence of synonymy, there is very little that can be compared between the genera. Besides undescribed pubic fragments of Deltadromeus, only the proximal ischium and mid caudals can be compared, which are schematically illustrated in Deltadromeus. Carrano and Sampson (2008) claimed Stromer said Bahariasaurus and neoceratosaurs "both shared an unusual constriction of the vertebral centra", but as noted above it is elongation that Stromer noted, which is also present in e.g. coelurosaurs. When entered into Tortosa et al.'s ceratosaur matrix, Bahariasaurus emerges sister to the only tetanurine Allosaurus, suggesting it is not a ceratosaur.
When entered into a modified version of Caranno et al's tetanurine matrix, Bahariasaurus emerges as an orionidan excluded from Piatnitzkysauridae, Megalosaurinae, Spinosauridae and Acrocanthosaurus+Carcharodontosaurinae. Further resolution depends on including more characters in analyses, especially around basal Coelurosauria.
References- Stromer, 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes Cenoman). 13. Dinosauria. Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung, Neue Folge. 22, 1-79.
Huene, 1948. Short review of the lower tetrapods. In Du Toit (ed.). Robert Broom commemorative volume. Royal Society of South Africa Special Publication. 65-106.
Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du Sahara central. Memoirs of the Geological Society of France. 88A, 1-57.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 464 pp.
Kurzanov, 1989. O proiskhozhdenii i evolyutsii infraotryada dinozavrov Carnosauria. Paleontologicheskiy Zhurnal. 1989(4), 3-14.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Osmólska and Dodson (eds.). The Dinosauria. University of California Press, Berkeley. 169-209.
Rauhut, 1995. Zur systematischen Stellung der afrikanischen Theropoden Carcharodontosaurus Stromer 1931 und Bahariasaurus Stromer 1934. Berliner Geowissenschaften Abhandlungen. 16, 357-375.
Sereno, Dutheil, Iarochene, Larsson, Lyon, Magwene, Sidor, Varricchio and Wilson, 1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation. Science. 272(5264), 986-991.
Rauhut, 1999. A dinosaur fauna from the Late Cretaceous (Cenomanian) of Northern Sudan. Palaeontologia Africana. 35, 61-84.
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.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria (second edition). University of California Press, Berkeley. 71-110.
Carrano and Sampson, 2008. The phylogeny of Ceratosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 6, 183-236.

Cruxicheiros Benson and Radley, 2010
= "Cruxicheiros" Benson and Radley, 2009 online
C. newmanorum Benson and Radley, 2010
= "Cruxicheiros newmanorum" Benson and Radley, 2009 online
Early Bathonian, Middle Jurassic
Chipping Norton Limestone Formation, England
Holotype- (WARMS G15770) partial femur
Paratype- ....(WARMS G15771) partial posterior cervical or anterior dorsal vertebra, incomplete dorsal neural arch, partial dorsal vertebra, several rib fragments, partial distal caudal vertebra, fragmentary scapulocoracoid, metacarpal (lost), partial ilium, proximal pubis, fibula (lost), fragments
Diagnosis- (after Benson and Radley, 2010) proximomedially inclined ridge within trochanteric fossa of femur.
Comments- The description of Cruxicheiros was realeased online in November 2009, though it was not officially published on paper until March 2010.
Benson and Radley (2010) found Cruxicheiros to be a tetanurine outside Megalosauria+Piatnitzkysauridae+Monolophosaurus+Chuandongocoelurus and Avetheropoda, though only one more step was needed to place it in Megalosauridae (as a eustreptospondyline). Additionally, only 3 more steps were needed to place it sister to Megalosaurus, showing numerous affinities are possible. Adding it to Carrano et al.'s (2012) matrix recovered Cruxicheiros as an orionidan outside Spinosauridae and Avetheropoda.
Reference- Benson and Radley, 2010. A new large-bodied theropod dinosaur from the Middle Jurassic of Warwickshire, United Kingdom. Acta Palaeontologica Polonica. 55(1), 35-42.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

Dandakosaurus Yadagiri, 1982
D. indicus Yadagiri, 1982
Hettangian-Pliensbachian, Early Jurassic
Kota Formation, India
Holotype- (GSI 1/54Y/76) proximal pubis
Paratypes- ?(GSI coll.) lateral tooth, dorsal vertebra (160 mm), proximal caudal vertebra (150 mm), proximal ischium
Diagnosis- open obturator notch in pubis; anteroproximal flaring and smoothly convex proximal surface of pubis; more proximally placed obturator notch than Patagonykus.
Description- The illustrated tooth is typical of most theropods in being laterally compressed, recurved and having fine serrations. The tooth seems more compressed (30% of FABL) than most theropods (eg. Liliensternus airelensis, Gojirasaurus, Dilophosaurus, Magnosaurus, Torvosaurus, Szechuanosaurus), though Liliensternus liliensterni is similar in this regard. Though interdental variation could be a factor here, as posterior teeth are known to be more laterally compressed in theropods, Dilophosaurus and Magnosaurus never reach a Dandakosaurus level of compression anywhere in the tooth row.
The dorsal vertebra is opisthocoelous and lacks a pleurocoel. Opisthocoelous dorsals are only known in non-tyrannoraptoran tetanurines (the exception is Parvicursorinae), suggesting Dandakosaurus is a member of this clade. The absence of a pleurocoel is of little use without positional data.
The caudal vertebra is amphicoelous, with "two lateral cavities on either side" and a keeled ventral surface. The first character is common in theropods. The second is only known in Spinostropheus, Carcharodontosaurus, megaraptorans, Patagonykus, caenagnathoids and Achillobator. Keeled proximal caudal centra are known in at least torvosaurs and carnosaurs.
The holotype proximal pubis has several odd characters. There is no obturator fenestra, just an open obturator notch, as in Elaphrosaurus, Eustreptospondylus, Suchomimus, Allosaurus, carcharodontosaurids and most coelurosaurs. The proximal border forms a smooth convex arch from the ilial contact to the ischia contact, with no distinct peduncles. This is approached in some coelophysoids (eg. Coelophysis longicollis referred specimen) and Archaeornithomimus, but is most similar to Patagonykus, Unenlagia and Achillobator. Proximally, the pubis flares sharply anteriorly to form an acute anteroproximal corner. This is similar to the situation in Coelurus, tyrannosaurids, Caudipteryx, Nanshiungosaurus and alvarezsaurids. It seems to only be developed in forms with somewhat mesopubic pelvic orientations. Additional evidence for mesopuby in Dandakosaurus may come from the possible pubic peduncle of the ischium, which fits the ischial surface of the pubis to form an angle of about 35 degrees between the bones.
Only the proximal portion of the ischium is preserved. There is a concave anterior margin which matches up with the ischial peduncular area of the pubis. No other area is appropriate for the pubic contact, though Yadagiri seems to have identified this as the area between the pubic peduncle and obturator process. If the concave area were the acetabulum, the pubis and ischium would be subparallel. Articulating the pubis more dorsoposteriorly on the ischium leaves nowhere for an acetabular surface or ischial peduncle. The acetabular surface is small and sharply concave, so may be broken and continuous with the ischial peduncle in life. The posterior surface is gently concave, while the anterior surface is broken.
Comments- This was excavated in 1958-1961 and first mentioned by Jain et al. (1962) as "a carnosaur ... less well represented than the sauropod in this first collection. Judging by its pelvis, it should prove to be of about the same size as the well-known Antrodemus valens ... that is, more than 30 ft. long." The phylogenetic relationships of this taxon are uncertain, especially considering how advanced it seems for its age. When entered into Carrano et al.'s (2012) matrix, it emerges as an orionidan excluded from Piatnitzkysauridae, Megalosaurinae, Spinosauridae, Yangchuanosaurus, derived metriacanthosaurines and Allosauria. The possibility elements have been misidentified or derive from more than one taxon (such as a sauropodomorph) should not be excluded. It is one of the earliest tetanurines. Previous suggestions regarding abelisauroid affinity (Aravind, DML 1997) were based on maxillary characters unknown in the specimen. Where Aravind got his data is unknown, but it can be considered irrelevant now that the known material has been determined.
References- Jain, Robinson and Roy Chowdhury, 1962. A new vertebrate fauna from the Early Jurassic of the Deccan, India. Nature. 194(4830), 755-757.
Yadagiri, 1982. Osteological studies of a carnosaurian dinosaur from the Lower Jurassic Kota Formation: Andhra Pradesh. Geological Survey of India (Progress Report for Field Season Programme 1981-1982), Regional Palaeontological Laboratories, Southern Region. 7 pp.
Aravind, DML 1997. https://web.archive.org/web/20191030094430/http://dml.cmnh.org/1997May/msg00840.html
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

"Megalosaurus" hungaricus Nopcsa, 1902
Coniacian-Santonian, Late Cretaceous
unnamed formation, Romania
Holotype- (MAFI ob. 3106, lost) tooth (~22 mm)
Comments- This species was discovered in an unnamed formation, not the later Sinpetru Formation as is often stated (Csiki and Grigorescu, 1998). Carrano et al. (2012) note the widely spaced distal serrations may be diagnostic, and the high DSDI (1.25) resembles basal tyrannosauroids and dromaeosaurids. This is also found in Marshosaurus, Acrocanthosaurus and Falcarius however, so the taxon is provisionally assigned to Orionides incertae sedis.
References- Nopcsa, 1902. Notizen uber Cretacische Dinosaurier. Pt. 2. Megalosaurus hungaricus nov. sp. ein Theropode der Siebenburgischen Kreide. Sitzungsberichte der Koniglichen Akademie Wissenschaften. 3, 104-107.
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.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

"Metriacanthosaurus" "reynoldsi" Welles, Powell and Pickering vide Pickering, 1995
Early Bathonian, Middle Jurassic
Chipping Norton Limestone Formation (= Charlbury Formation), England
Material- ?(SDM 44.16) proximal scapula (Reynolds, 1939)
(SDM 44.19; intended holotype) ilium (Reynolds, 1939)
? tooth (39 mm, FABL 23.5 mm) (Reynolds, 1939)
Diagnosis- (suggested) several pronounced vertical ridges above the supracetabular shelf extending halfway to the blade's dorsal edge.
Other diagnoses- Pickering (DML, 2002) listed several features as differing from Metriacanthosaurus parkeri. Of these, the straighter dorsal ilial margin and broadly exposed medial wall of the brevis shelf are seen in Eustreptospondylus, while the long ischial peduncle is also present in Yangchuanosaurus zigongensis. The lower ilial blade are seen in both. Contra Pickering, the pubic peduncle is not noticably longer nor the subpreacetabular notch more open than in Metriacanthosaurus.
Comments- Lydekker (1888) referred a metatarsal III (NHMUK R413) from the Chipping Norton Formation to Megalosaurus bucklandi. Gardiner (1937, 1938) and Reynolds (1938) reported large theropod remains from two quarries in the Chipping Norton Limestone Formation, which Reynolds (1939) described and referred to Megalosaurus. Pickering (1995) credited the name Metriacanthosaurus reynoldsi to Welles, Powell and Pickering in his 1995 unpublished bibliographic manuscript. It was later used in the comparative section of another unpublished manuscript (Welles and Pickering, 1999). This paper was largely extracted from the European megalosaur manuscript Welles and Powell worked on in the 1970s but never published, specifically the Megalosaurus redescription section. Pickering intends to publish an updated version of the megalosaur manuscript as Mutanda Dinosaurologica, and has posted small excerpts including the diagnosis of Metriacanthosaurus "reynoldsi" online (DML, 2002). In any case, the name is a nomen nudum as Pickering didn't follow ICZN Article 8.1.3- it must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies. The 1999 paper shows his new taxon to be based on all the Chipping Norton theropod material from both quarries, as well as NHMUK R413, scapula OUM J.29800 and a few other elements. Day and Barrett (2004) believed both their Megalosaurus femoral morphotypes A and B were present in the sample- SDM 44.23 as morphotype B and SDM 44.24 as morphotype A. Benson (2009, 2010) referred the New Park Quarry material to Megalosaurus bucklandii based on the close resemblence of maxilla SDM 44.1 to Taynton Limestone specimens and an M. bucklandii autapomorphy in sacrum SDM 44.4, while other elements were provisionally referred as there is no evidence of more than one taxon in the quarry. These additional elements are- maxilla NHMUK R8303, anterior dentary NHMUK R8304, partial dentary NHMUK R8305, proximal ischium NHMUK R9668, proximal caudal vertebrae NHMUK R9672-9673, partial anterior cervical vertebra NHMUK R9674, mid caudal vertebrae NHMUK R9675-9676, proximal caudal vertebra NHMUK R9677, mid caudal vertebra SDM 44.5, coracoids SDM 44.14-15, humerus SDM 44.18, femur SDM 44.24 and distal metatarsal IV 44.25. NHMUK R413 and OUM J.29800 were also referred to M. bucklandii based on autapomorphies. Additional New Park Quarry material referred by Welles and Pickering to "reynoldsi" but not mentioned by Benson is here provisionally referred to Megalosaurus- sacra NHMUK R9679 and R9680 and dorsal SDM 44.10. Finally, Benson (2010) referred two specimens from Oakham Quarry to M. bucklandii based on autapomorphies (ischium SDM 44.20 and metatarsal III NHMUK R9665), but refrained from referring additional elements from this quarry as he notes some "can be referred to a second, unnamed taxon (R. B. J. Benson, unpubl. data)." This near certainly refers to "reynoldsi", whose intended holotype is an ilium from Oakham Quarry (SDM 44.19). Further study by Benson and/or Pickering may clarify the identity of the other Oakham Quarry elements. Welles and Pickering also referred several elements from other quarries to "reynoldsi"- dorsal GSM 37523, scapula SDM 34.17 and humerus 44.22 from the Chipping Norton Formation, and proximal caudals OUM J13720 and J.29799 from the Sharp's Hill Formation. These are provisionally placed as Tetanurae incertae sedis here until their descriptions are published.
The identity of SDM 44.19 is difficult to determine from the published illustration, since many useful ilial characters would only be visible in medial or ventral views. However, it is a tetanurine based on the lack of a strong crest between the supracetabular crest and brevis fossa, and is not a coelurosaur based on the large ischial peduncle and widely exposed brevis fossa in lateral view. It differs from Megalosaurus and Metriacanthosaurus in many of the same ways- broad medial wall to the acetabulum, especially posteriorly; longer ischial peduncle; broad esposure of medial wall of the brevis fossa; low ilial blade. It is more similar to Metriacanthosaurus in the wide subpreacetabular notch, and anteroposteriorly wide ischial peduncle, but shares a series of short vertical ridges above the acetabulum with Megalosaurus (albeit more pronounced). Pickering's (DML, 2002) only ilial character for Metriacanthosaurus is "ilium + ischium fused", which isn't true of M. parkeri or "reynoldsi." Thus is seems whatever "reynoldsi" is, there's no reason to refer it to Metriacanthosaurus. A greater resemblence is seen to Eustreptospondylus, which shares the broad esposure of medial wall of the brevis fossa and low ilial blade. Another similar ilium is that of Yangchuanosaurus zigongensis, which shares the broad medial wall to the acetabulum, longer ischial peduncle, and low ilial blade. Which of these two taxa it is more closely related to will require study of the specimen itself and additional specimens from Oakham Quarry. Adding it to Carrano et al.'s (2012) matrix results in it being a non-carcharodontosaurid, non-coelurosaur orionidan.
The illustrated Oakham tooth (NHMUK or SDM colls) is short (height/FABL 1.66) and moderately recuved with perpendicular serrations on at least the apical half of the distal carina. Only light longitudinal striations are indicated on the enamel. At least one of the scapulae mentioned by Reynolds (SDM 44.16 or 44.17) is unfused to the coracoid, though it is unknown if this one was from Oakham Quarry.
References- Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum of Natural History, London. 309 pp.
Gardiner, 1937. Reptile-bearing oolite, Stow. Reports of the British Association for the Advancement of Science (Blackpool). 1936, 296.
Gardiner, 1938. Reptile-bearing oolite, Stow. Reports of the British Association for the Advancement of Science (Nottingham). 1937, 290.
Reynolds, 1938. A collection of reptilian bones from the Oölite near Stow-in-the-Wold, Glos. Reports of the British Association for the Advancement of Science. 1937, 356-357.
Reynolds, 1939. A collection of reptile bones from the Oolite near Stow-on-the-Wold, Gloucestershire. Geological Magazine. 76, 193-214.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry. A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola, California. 478 pp.
Welles and Pickering, 1999. Megalosaurus bucklandii. Private publication of Stephen Pickering, An extract from Archosauromorpha: Cladistics & Osteologies. A Fractal Scaling in Dinosaurology Project. 119 pp.
Day and Barrett, 2004. Material Referred to Megalosaurus (Dinosauria: Theropoda) from the Middle Jurassic of Stonesfield, Oxfordshire, England: One taxon or two? Proceedings of the Geologists' Association. 115, 359-366.
Benson, 2009. An assessment of variability in theropod dinosaur remains from the Bathonian (Middle Jurassic) of Stonesfield and New Park Quarry, UK and taxonomic implications for Megalosaurus bucklandii and Iliosuchus incognitus. Palaeontology. 52(4), 857-877.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Pickering, in prep. Mutanda Dinosaurologica.

"Yunyangosaurus" Dai and Xu, 2019
"Y. puanensis" Dai and Xu, 2019
Aalenian, Middle Jurassic
Laojun Village, Xintiagou Formation, Chongqing, China
Material- (CLGPR V00002) (subadult) (~4.7 m) axis (58 mm), third/fourth cervical vertebra (69 mm), partial fifth cervical vertebra (68 mm), partial sixth cervical vertebra (54 mm), seventh cervical vertebra (70 mm), ninth cervical vertebra (58 mm), tenth cervical vertebra (61 mm), two proximal cervical ribs, incomplete first dorsal neural arch (61 mm), second dorsal centrum (52 mm), third dorsal vertebra (60 mm), incomplete mid dorsal vertebra (63 mm), several partial dorsal ribs, three chevrons
Diagnosis- (after Dai et al., 2020) mediolateral width of axial centrum tapers ventrally, resulting in subtriangular outline in posterior view; axis with an accessory medial spinopostzygapophyseal lamina; ball-like structure at base of anterior margin of axial neural spine; ventral median ridge present in anterior cervicals (also in Marshosaurus); posterior pneumatic foramen or fossa present in some cervical centra (also in Piatnitzkysaurus and Condorraptor); prominent epipophyses present in posterior cervicals; cervical epipophyses not angled laterally; posterior cervical and anterior dorsal vertebrae with transversely bifurcated neural spines (also in anterior dorsals of Condorraptor, Marshosaurus and Piatnitzkysaurus); fossae immediately ventral to anterior dorsal parapophyses.
Comments- This was discovered in 2016 and first announced as a new theropod from Yunyang County (Global Times, 2019; the new ornithopod being Sanxiasaurus CLGPR V00003 and the plesiosaur Pliosauroidea indet. CLGPR V00004) and named by Dai and Xu (2019) in an SVP abstract, as a new taxon of megalosaurid. Unfortunately, ICZN Article 9.10 states "materials issued primarily to participants at meetings (e.g. symposia, colloquia, congresses, or workshops), including abstracts and texts of presentations or posters" do not count as published works, making "Yunyangosaurus puanensis" Dai and Xu, 2019 a nomen nudum. It was later described in detail by Dai et al. (2020) and named again as if new without mention of the SVP abstract, but this paper has no mention of ZooBank and as of January 21 2020 "Yunyangosaurus" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Yunyangosaurus puanensis" Dai et al., 2020 is also a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically.
Dai et al. added "Yunyangosaurus" to Carrano et al.'s tetanurine analysis and recovered it as a tetanurine closer to birds than Chuandongocoelurus but excluded from piatnitzkysaurids, spinosaurids, allosauroids and neocoelurosaurs. Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it as either a basal megalosauroid or sister to Allosauria.
References- Dai and Xu, 2019. A new megalosaurid theropod from the Middle Jurassic Xintiangou Formation of Chongqing, People's Republic of China and its implication for early tetanuran evolution. Journal of Vertebrate Paleontology. Program and Abstracts 2019, 86.
Global Times, 2019. New dinosaur species found in Chongqing, southwest China. Global Times. June 17. https://www.globaltimes.cn/page/201906/1154634.shtml
Dai, Benson, Hu, Ma, Tan, Li, Xiao, Hu, Zhou, Wei, Zhang, Jiang, Li, Peng, Yu and Xu, 2020. A new possible megalosauroid theropod from the Middle Jurassic Xintiangou Formation of Chongqing, People’s Republic of China and its implication for early tetanuran evolution. Scientific Reports. 10:139.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.

unnamed orionidan (Galton and Molnar, 2005)
Middle Bathonian, Middle Jurassic
Stonesfield Slate, England
Material- (OUM J28971) incomplete ilium
Comments- Galton and Molnar (2005) described OUM J28971 as a specimen of Iliosuchus. Benson (2009) believed OUM J28971 was from a different taxon because its lateral ilial ridge is more vertical, there are accessory ridges anterior and posterior to it, and the brevis fossa is visible laterally at the base of the postacetabular process. The latter difference is more accurately expressed by saying the brevis fossa of the holotype does not extend as close to the ischial peduncle, though even in OUM J28971 it is very shallow and laterally angled there. Additional differences are- the pubic peduncle of OUM J28971 is much wider compared to the length of its articular surface when compared to OUM J29780, and has a posteriorly concave edge on its articular surface; the pubic peduncle is less anteriorly oriented than the holotype. Carrano et al. (2012) agreed with Benson and also proposed larger size as a distinguishing character, but this could be ontogenetic as the age of all specimens is unknown.
Benson merely said this is Theropoda indet., but Carrano et al. specified it further as Tetanurae indet.. Indeed when added to Carrano et al.'s matrix it emerges as a non-piatnitzkysaurid, non-allosaurian orionidan. As most of the characters which differ from Iliosuchus are shared with Megalosaurus and both are found in the same formation, OUM J28971 may be a juvenile of that taxon.
References- Galton and Molnar, 2005. Tibiae of small theropod dinosaurs from Southern England. In Carpenter (Ed.). The Carnivorous Dinosaurs. 3-22.
Benson, 2009. An assessment of variability in theropod dinosaur remains from the Bathonian (Middle Jurassic) of Stonesfield and New Park Quarry, UK and taxonomic implications for Megalosaurus bucklandii and Iliosuchus incognitus. Palaeontology. 52(4), 857-877.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

unnamed orionidan (Royo-Torres, Cobos and Alcalá, 2009)
Tithonian-Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (CPT-1980) lateral tooth (98.2x43.1x19.9 mm)
Comments- Though originally referred to Allosauroidea by Royo-Torres et al. (2009), Gasco et al. (2012) reevaluated it as a megalosaurid or carcharodontosaurid.
References- Alcala, Royo-Torres, Cobos and Luque, 2009. Updating dinosaur record from Teruel (Aragon, Spain). Journal of Vertebrate Paleontology. 29(3), 53A.
Royo-Torres, Cobos and Alcalá, 2009. Diente de un gran dinosaurio terópodo (Allosauroidea) de la Formación Villar del Arzobispo (Titónico-Berriasiense) de Riodeva (España). Estudios Geológicos. 65(1), 91-99.
Gascó, Cobos, Royo-Torres, Mampel and Alcalá, 2012. Theropod teeth diversity from the Villar del Arzobispo Formation (Tithonian-Berriasian) at Riodeva (Teruel, Spain). Palaeobiodiversity and Palaeoenvironments. 92(2), 273-285.

unnamed probable orionidan (Brusatte and Clark, 2015)
Late Bajocian-Early Bathonian, Middle Jurassic
Vlatos Sandstone Formation, Scotland
Material- (GLAHM 152390a) lateral tooth (21.3x13.2x7.7 mm)
Comments- Brusatte and Clark (2015) found that this tooth is most similar to megalosaurids, tyrannosauroids and dromaeosaurids.
Reference- Brusatte and Clark, 2015. Theropod dinosaurs from the Middle Jurassic (Bajocian-Bathonian) of Skye, Scotland. Scottish Journal of Geology. 51(2), 157-164.

Megalosauroidea Huxley, 1869 vide Nopcsa, 1928
Definition- (Megalosaurus bucklandii <- Allosaurus fragilis, Passer domesticus) (Benson, 2010)
Other definitions- (Megalosaurus bucklandii <- Allosaurus fragilis, Tyrannosaurus rex) (Rauhut and Pol, 2019; modified from Carrano et al., 2012)
(Megalosaurus bucklandii <- Passer domesticus) (Hendrickx, Hartman and Mateus, 2015)
= Megalosauri Fitzinger, 1843
= Megalosauroides Gervais, 1852
= Spinosauroidea Stromer, 1915 sensu Olshevsky, 1991
Definition- (Spinosaurus aegyptiacus <- Passer domesticus) (Holtz et al., 2004)
Other definitions- (Spinosaurus aegyptiacus, Torvosaurus tanneri <- Allosaurus fragilis, Passer domesticus) (Allain et al., 2012)
= Spinosauria Olshevsky, 1991
= Torvosauroidea Jensen, 1985 vide Sereno et al., 1994
= Spinosauroidea sensu Allain et al., 2012
Definition- (Spinosaurus aegyptiacus, Torvosaurus tanneri <- Allosaurus fragilis, Passer domesticus)
= Megalosauroidea sensu Carrano et al., 2012
Definition- (Megalosaurus bucklandii <- Allosaurus fragilis, Tyrannosaurus rex)
= Megalosauroidea sensu Hendrickx, Hartman and Mateus, 2015
Definition- (Megalosaurus bucklandii <- Passer domesticus)
Comments- Spinosauroidea is the name most commonly given to the megalosaur-spinosaur clade from 1994 to 2010, based on analyses which usually excluded Megalosaurus during the period of time most workers restricted the name to the lectotype dentary and thus excluded it from most analyses. However, according to the ICZN, a superfamily containing Megalosaurus must be called Megalosauroidea. The issue is complicated by the fact Spinosauroidea was originally given a node-based definition, while Megalosauroidea has been given a more inclusive stem-based definition.
Sereno (in press) suggested using Spinosauria for a stem-based clade including Spinosauroidea but excluding avetheropods, in case taxa like eustreptospondylines or Poekilopleuron fall outside the Torvosaurus+Spinosaurus clade. This is similar to his prior use of Torvosauroidea (to contain Afrovenator plus what is now defined as Spinosauroidea) and to Holtz et al.'s and Allain et al.'s stem-based version of Spinosauroidea. However, Megalosauroidea has priority over all of those names and was defined this way by Benson (2010), with the Torvosaurus+Spinosaurus clade now named Megalosauria.
References- Carrano, Benson and Sampson, 2009. The phylogeny of megalosauroids (Dinosauria: Theropoda) with implications for the evolution of North African paleoecosystems. First International Congress on North African Vertebrate Palaeontology. 21-22.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften. 99(5), 369-377.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs. Scientific Reports. 9:18826.

unnamed megalosauroid (Molnar, Angriman and Gasparini, 1996)
Coniacian-Santonian, Late Cretaceous
Hidden Lake Formation, James Ross Island, Antarctica
Material- (MLP 89-XII-1-1) (~3 m) distal tibia
Comments- Molnar et al. (1996) believed this was from a megalosauroid or basal carnosaur (due to placing Piatnitzkysaurus in the clade). Carrano et al. (2012) agreed with the similarities, and suggested it was megalosauroid as they placed Piatnitzkysaurus in that clade.
References- Molnar, Angriman and Gasparini, 1996. An Antarctic Cretaceous theropod. Memoirs of the Queensland Museum. 39, 669-674.

Piatnitzkysauridae Carrano, Benson and Sampson, 2012
Definition- (Piatnitzkysaurus floresi <- Megalosaurus bucklandii, Spinosaurus aegyptiacus) (modified after Carrano, Benson and Sampson, 2012)
Other definition- (Piatnitzkysaurus floresi <- Megalosaurus bucklandii, Spinosaurus aegyptiacus, Allosaurus fragilis) (Rauhut and Pol, 2019)
= Piatnitzkysauridae sensu Rauhut and Pol, 2019
Definition- (Piatnitzkysaurus floresi <- Megalosaurus bucklandii, Spinosaurus aegyptiacus, Allosaurus fragilis)
Diagnosis- (after Carrano et al., 2012) short or absent anterior maxillary ramus; two parallel rows of nutrient foramina on lateral surface of maxilla; vertically striated or ridged paradental plates (also in Megalosaurus); no axial pleurocoels; moderate development of axial diapophyses; reduced axial parapophyses (also in Eustreptospondylus and Afrovenator); anteriorly inclined posterior dorsal neural spines; canted distal humeral condyles (also in Poekilopleuron).
Comments- Smith et al. (2007) recovered Piatnitzkysaurus and Condorraptor in a clade, while Benson (2008, 2010) found Marshosaurus to be related as well. Carrano et al. (2012) named the latter group Piatnitzkysauridae.
Currie and Zhao (1994) advanced the idea Piatnitzkysaurus was an abelisaurid, reminiscent of Bonaparte and Novas' (1985) remark their new family Abelisauridae has "diverse characters that unite it with Jurassic forms such as Piatnitzkysaurus and Ceratosaurus" (translated). Holtz (1995), Rauhut (2003) and Smith et al. (2007; with Condorraptor) found Piatnitzkysaurus to be a non-orionidan tetanurine. Benson (2008, 2010) and Carrano et al. (2012) found piatnitzkysaurids to be non-megalosaurian megalosauroids in their large analyses, which is provisionally accepted here. Both Bonaparte (1979) and Gao (1999) place Piatnitzkysaurus in Megalosauridae itself, and Holtz et al. (2004) found it to be a eustreptospondyline/afrovenatorine in his analysis. Paul (1988), Novas (1992), Perez-Moreno et al. (1993) and Holtz (2000) all find Piatnitzkysaurus to be a tetanurine closer to avetheropods than megalosauroids. Molnar et al. (1981) placed Piatnitzkysaurus in his Allosauridae, which is equivalent to modern Allosauroidea. Holtz (1992) found Piatnitzkysaurus to be a non-allosaurian carnosaur. While listed under Allosauridae in Kurzanov (1989) and Molnar et al. (1990), the text indicates they viewed Piatnitzkysaurus as a carnosaur outside Allosaurus+Tyrannosaurus. Bonaparte (1986) in his monograph of the genus placed it in an Allosauridae oddly expanded to include Dilophosaurus, but not Ceratosaurus or Torvosaurus.
Using Carrano et al.'s analysis to test the liklihood of alternate topologies, moving piatnitzkysaurids into Carnosauria is only 2 steps longer (megalosauroids follow), and in this case they are sister to allosauroids. It also takes 2 steps to move them sister to Avetheropoda. Enforcing a non-orionidan position leads to trees 3 steps longer, meaning any of these topologies is easily possible. Megalosaurid piatnitzkysaurids are 4 steps longer, and further enforcing them as afrovenatorines is 6 steps longer, so increasingly less likely but still possible. Placing them as closer to Allosauria than metriacanthosaurids as in Kurzanov and Molnar et al. is 18 steps longer though, and thus implausible. Making Piatnitzkysaurus an abelisaurid (only Condorraptor follows unlike previous tests where Marshosaurus does as well) is 56 steps more, so near certainly wrong.
References- Bonaparte, 1979. Dinosaurs: A Jurassic assembalge from Patagonia. Science. 205, 1377-1379.
Molnar, Flannery and Rich, 1981. An allosaurid theropod dinosaur from the Early Cretaceous of Victoria, Australia. Alcheringa. 5, 141-146.
Bonaparte and Novas, 1985. Abelisaurus comahuensis, n. g., n. sp., Carnosauria del Cretacico Tardio de Patagonia. Ameghiniana. 21(2-4), 259-265.
Bonaparte, 1986. Les dinosaures (Carnosaures, Allosauridés, Sauropodes, Cétosauridés) du Jurassique Moyen de Cerro Cóndor (Chubut, Argentina). Annales de Paléontologie (Vert.-Invert.). 72(3), 247-289.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 464 pp.
Kurzanov, 1989. O proiskhozhdenii i evolyutsii infraotryada dinozavrov Carnosauria. Paleontologicheskiy Zhurnal. 1989(4), 3-14.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Osmólska and Dodson (eds.). The Dinosauria. University of California Press, Berkeley. 169-209.
Holtz, 1992. An unusual structure of the metatarsus of Theropoda (Archosauria: Dinosauria: Saurischia) of the Cretaceous. PhD thesis. Yale University. 347 pp.
Novas, 1992. La evolucion de los dinosaurios carnivoros. In Sanz and Buscalioni (eds.). Los Dinosaurios y Su Entorno Biotico: Actas del Segundo Curso de Paleontologia in Cuenca. Instituto "Juan Valdez", Cuenca, Argentina. 126-163.
Perez-Moreno, Sanz, Sudre and Sige, 1993. A theropod dinosaur from the Lower Cretaceous of Southern France. Revue de Paleobiologie. 7, 173-188.
Currie and Zhao, 1994. A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences. 30(10), 2037-2081.
Holtz, 1995. A new phylogeny of the Theropoda. Journal of Vertebrate Paleontology. 15(3), 35A
Gao, 1999. [A complete carnosaur skeleton from Zigong, Sichuan: Yangchuanosaurus hepingensis]. Sichuan Science and Technology Press, Chengdu. 100 pp.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. GAIA. 15, 5-61.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Benson, 2008. A new theropod phylogeny focusing on basal tetanurans and its implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs. Scientific Reports. 9:18826.

Gasosaurus Dong and Tang, 1985
G. constructus Dong and Tang, 1985
Bajocian, Middle Jurassic
Dashanpu, Xiashaximiao Formation, Sichuan, China
Holotype- (IVPP V7264) four cervical centra, seven dorsal vertebrae, first sacral vertebra, second sacral vertebra, third sacral vertebra, fourth sacral vertebra, fifth sacral vertebra, seven caudal vertebrae, humerus, ilium, pubis (345 mm), ischium (305 mm), femur (425 mm), tibia (370 mm), fibula, astragalus, metatarsal II, metatarsal III, pedal ungual IV
Paratype- ?(IVPP V7265) three teeth
Referred- material (Holtz, 2000)
Comments- Traditionally associated with megalosauroids, Holtz (2000) found it to be a basal coelurosaur in his analyses. This was based on the upturned femoral head, anterior trochanter cleft from the head, and proximal fibula being >75% the proximal tibial width. However, he also indicated new undescribed specimens suggest Gasosaurus is a carnosaur, perhaps a sinraptorid (Currie pers. comm. 1998 to Holtz). Carrano et al. (2012) found that the proximal femur is broken, so that the seemingly high anterior trochanter is an illusion. They believed it most likely to be a non-coelurosaur tetanurine and stated the holotype is being restudied (Hone pers. comm. to Carrano et al.). Hartman et al. (2019) recovered it sister to Piatnitzkysaurus and Condorraptor, although non-maniraptoromorph characters were poorly sampled. It is provsionally placed there pending future analysis.
References- Dong and Tang, 1985. A new Mid-Jurassic theropod (Gasosaurus constructus gen. et sp. nov.) from Dashanpu, Zigong, Sichuan Province, China. Vertebrata PalAsiatica. 23(1), 77-82.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. Gaia. 15, 5-61.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Marshosaurus Madsen, 1976
M. bicentissimus Madsen, 1976
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of Morrison Formation, Colorado?, Utah, US
Holotype- (UMNH VP 6373; = UUVP 2826) ilium (375 mm)
Paratypes-....(UMNH VP 6379; = UUVP 2832) ischium (305 mm)
....(UMNH VP 380; = UUVP 2878) ischium
....(UMNH VP 6387; = UUVP 4736) pubis (393 mm)
?(UMNH VP 6364; = UUVP 40-555) dentary
?(UMNH VP 6367; = UUVP 3454) dentary
?(UMNH VP 6368; = UUVP 3502) dentary
(UMNH VP 6372; = UUVP 1182, UUVP 1845) ilia
(UMNH VP 6374; = UUVP 2742) ilium
?(UMNH VP 7820; = UUVP 3236) premaxilla
?(UMNH VP 7824; = UUVP 1846, UUVP 1864) maxillae
?(UMNH VP 7825; = UUVP 4695) maxilla
Referred- ?(BYUVP 5201) proximal caudal vertebra (53 mm) (Britt, 1991)
?(CMNH 21704; = DINO 16455b; = DMN 343; juvenile) posterior skull, posterior mandible, atlas, axis, cervical vertebrae, dorsal vertebrae, dorsal rib, scapula, humerus (Chure, Madsen and Britt, 1993)
(DMNH 3718) partial skull, vertebrae (Carrano et al., 2012)
(DMNH coll.) eight teeth (Kane, 2020)
(UMNH VP 6374; = UUVP 2742) ilium (Carrano et al., 2012)
(UMNH VP 6384; = UUVP 40-295) pubis (Carrano et al., 2012)
(UMNH VP 6386; = UUVP 1867) pubis (Carrano et al., 2012)
?(UUVP 99) caudal vertebra (Britt, 1991)
?(UUVP 441) caudal vertebra (Britt, 1991)
?(UUVP 5247) caudal vertebra (Britt, 1991)
?(UUVP 5780) caudal vertebra (Britt, 1991)
?(UUVP coll.) dorsal vertebrae (Chure, Britt and Madsen, 1997)
(YPM-PU 72-1) partial pelvis (Carrano et al., 2012)
Diagnosis- (after Carrano et al., 2012)
Comments- Marshosaurus was originally based on pelvic elements, with cranial elements tentatively referred, all from the Cleveland-Lloyd Quarry of Utah (Madsen, 1976). The holotype ilium was discovered in 1962, along with paratypes UMNH VP 380, 6367, 6368, 6374, 6379 and 7820. Britt (1991) described a caudal vertebra from the Dry Mesa Quarry of Colorado which resembled others from the Cleveland-Lloyd Quarry. He tentatively referred these to Marshosaurus, with another form (BYUVP 5073, 5103 and 8908) referred to Stokesosaurus. However, these identifications may be switched. Chure et al. (1993) noted a medium-sized partial skeleton found in the Brushy Basin Member of Dinosaur National Monument, stating the camerate vertebral pneumatization is primitive while the braincase pneumatization resembles tyrannosaurids more than Allosaurus or Ceratosaurus. This was described a bit more by Chure et al. (1997), and referred to Marshosaurus based on resemblence to undescribed and questionably referred dorsal neural spines from Cleveland-Lloyd.
Relationships- Madsen (1976) originally left Marshosaurus as Theropoda incertae sedis, but noted resemblences to several coelurosaurs (Coelurus, Microvenator, Velociraptor, Deinonychus). Benson (2008, 2010) and Carrano et al. (2012) have found it to be a non-megalosaurian megalosauroid, related to Piatnitzkysaurus in Piatnitzkysauridae, which is followed here. Russell (1984) included Marshosaurus in the Megalosauridae. Chure et al. (1993) called Marshosaurus a carnosaur, and in 1997 clarified it as a "primitive carnosaur", closer to Megalosaurus and Eustreptospondylus than derived carnosaurs. Paul (1988) placed it closer to Avetheropoda than megalosauroids. Kurzanov (1989) placed Marshosaurus as an allosaurid, though his family may have been paraphyletic to tyrannosaurids. Holtz et al. (2004) found Marshosaurus to be a basal coelurosaur, though with very low support. Interestingly, Paul, Kurzanov and Holtz et al. all believed Marshosaurus was more closely related to Allosaurus than Piatnitzkysaurus, though only the latter had the genera widely separated.
Enforcing Marshosaurus to be sister to Avetheropoda in Carrano et al.'s (2012) matrix is 2 steps longer, as is making it be a carnosaur, and making it be a megalosaurid is 4 steps longer, so all are easily possible. Forcing it to be a coelurosaur takes 12 more steps, and forcing it to be closer to Allosauria than metriacanthosaurids takes 18 more steps, so are both unlikely. All of these end up keeping it in Piatnitzkysauridae. Breaking this up by additionally forcing it to be closer to Allosaurus than Piatnitzkysaurus is unlikely regardless of the alternate placement- sister to Avetheropoda (Piatnitzkysaurus ends up just basal to both as in Paul, 1988) takes 10 more steps; coelurosaur (Piatnitzkysaurus ends up outside Avetheropoda) takes 16 more steps, and closer to Allosauria than metriacanthosaurids (Piatnizkysaurus is a non-allosauroid carnosaur) takes 23 more steps.
References- Madsen, 1976. A second new theropod dinosaur from the Late Jurassic of east central Utah. Utah Geology. 3, 51-60.
Russell, 1984. A check list of the families and genera of North American dinosaurs. Syllogeus. 53, 1-35.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Britt, 1991. Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri. Brigham Young University Geology Studies. 37, 1-72.
Chure, Britt and Madsen, 1993. New data on the theropod Marshosaurus from the Morrison Formation (Upper Jurassic: Kimmeridgian-Tithonian) of Dinosaur NM. In Santucci (ed.). National Park Service Paleontology Research Abstract Volume. Technical Report NPS/NRPEFO/NRTR 93/11:28
Chure, Madsen and Britt, 1993. New data on theropod dinosaurs from the Late Jurassic Morrison FM. (MF). Journal of Vertebrate Paleontology. 13(3), 30A.
Chure, Britt and Madsen, 1997. A new specimen of Marshosaurus bicentesimus (Theropoda) from the Morrison Formation (Late Jurassic) of Dinosaur National Monument. Journal of Vertebrate Paleontology. 17(3), 38A.
Benson, 2008. A new theropod phylogeny focusing on basal tetanurans and its implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Kane, 2020. Identifying Jurassic theropod genera using GIS maps of tooth serrations. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 197.
M? sp. (Naish, online 2001)
Late Jurassic?
Europe
Comments- Naish (DML, 2001) alludes to European Marshosaurus remains.
Reference- Naish, DML 2001. https://web.archive.org/web/20191030094445/http://dml.cmnh.org/2001Mar/msg00328.html

Condorraptor Rauhut, 2005
C. currumili Rauhut, 2005
Middle Toarcian, Early Jurassic
Cañadón Asfalto Formation, Chubut, Argentina
Holotype- (MPEF-PV 1672) tibia
Paratypes- ....(MPEF-PV 1673) ?fourth cervical vertebra (57 mm)
....(MPEF-PV 1674) ?seventh cervical vertebra (70 mm)
....(MPEF-PV 1675) tenth cervical vertebra (59 mm)
....(MPEF-PV 1676) first dorsal centrum (61 mm)
....(MPEF-PV 1677) mid dorsal centrum (~70 mm)
....(MPEF-PV 1678) mid dorsal centrum (~68 mm)
....(MPEF-PV 1679) partial anterior dorsal neural arch
....(MPEF-PV 1680) posterior dorsal vertebra (80 mm)
....(MPEF-PV 1681) second sacral vertebra (75 mm), third sacral vertebra (75 mm), fourth sacral vertebra (72 mm)
....(MPEF-PV 1682) mid caudal vertebra (77 mm)
....(MPEF-PV 1683) distal caudal vertebra (67 mm)
....(MPEF-PV 1684) dorsal rib fragment
....(MPEF-PV 1685) dorsal rib fragment
....(MPEF-PV 1686) ilial fragment
....(MPEF-PV 1687) ilial fragment
....(MPEF-PV 1688) pubic fragment
....(MPEF-PV 1689) partial ischium
....(MPEF-PV 1690) distal femur
....(MPEF-PV 1691) distal femur
....(MPEF-PV 1692) metatarsal IV (242 mm)
....(MPEF-PV 1693) proximal pedal ungual
....(MPEF-PV 1694) lateral tooth (FABL 11 mm)
....(MPEF-PV 1695) lateral tooth (FABL 10.5; ~26 mm)
....(MPEF-PV 1696) proximal pubis
....(MPEF-PV 1697) ?second dorsal vertebra (57.5 mm)
....(MPEF-PV 1700) posterior dorsal vertebra (76 mm)
.....(MPEF-PV 1701) first sacral vertebra (69 mm)
....(MPEF-PV 1702) anterior caudal vertebra (56 mm)
....(MPEF-PV 1703) partial chevron
....(MPEF-PV 1704) ilial fragment
....(MPEF-PV 1705) anterior dorsal vertebra (65 mm)
Diagnosis- (from Rauhut, 2005) pleurocoel in anterior cervical vertebrae placed behind the posteroventral corner of the parapophyses; large, shallow depression laterally on the base of the cnemial crest; metatarsal IV with a distinct step dorsally between shaft and distal articular facet.
Other diagnoses- Carrano et al. (2012) noted a large nutrient foramen on the lateral side of the ischial peduncle of the ilium is also present in other theropods like Piatnitzkysaurus and Megalosaurus. They also stated the proximal tibia is abraded, making the absence of a posterior notch uncertain.
Comments- Originally ambigiously said to be a basal tetanurine by Rauhut (2005), Smith et al. (2007), Benson (2008, 2010) and Carrano et al. (2012) have all since found it to be a piatnitzkysaurid.
References- Rauhut, 2002. Dinosaur evolution in the Jurassic: A South American perspective. Journal of Vertebrate Paleontology. 22(3), 89A.
Rauhut, 2005. Osteology and relationships of a new theropod dinosaur from the Middle Jurassic of Patagonia. Palaeontology. 48(1), 87-110.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Benson, 2008. A new theropod phylogeny focusing on basal tetanurans and its implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

Piatnitzkysaurus Bonaparte, 1979
P. floresi Bonaparte, 1979
Middle Toarcian, Early Jurassic
Cañadón Asfalto Formation, Chubut, Argentina
Holotype- (PVL 4073) (4.24 m, 275 kg (Mazzetta et al., 2000 estimated 450 kg) subadult) maxillae, frontal, braincase, anterior dentary, axis, incomplete fourth cervical vertebra, fifth cervical vertebra, partial seventh cervical vertebra, tenth cervical vertebra, incomplete first dorsal vertebra, third dorsal vertebra, incomplete fourth dorsal vertebra, sixth dorsal vertebra, partial seventh dorsal vertebra, ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal fragments, thirteenth dorsal neural arch, fourteenth dorsal vertebra, three dorsal rib fragments, four sacral vertebrae, second caudal vertebra, fourth caudal vertebra, incomplete scapulae (480 mm), partial coracoids, humerus (286 mm), ulna (218 mm), partial ilia, incomplete pubes (450 mm), ischium (~423 mm), femora (552 mm), tibiae (492 mm), fibulae (470 mm), metatarsal III (282 mm)
Referred- (MACN CH 895) two posterior dorsal vertebrae, two dorsal centra, four sacral vertebrae, humerus, pubis, partial ischia, tibia (515 mm), metatarsal II (253 mm), metatarsal III (290 mm), metatarsal IV (252 mm) (Bonaparte, 1986)
Diagnosis- (after Rauhut, 2004) parasphenoid recess; parasphenoid recesses communicate; basipterygoid recesses longer anteroposteriorly than high dorsoventrally; width of the articular surface of the basipterygoid processes is more than twice their length and the transverse span between the processes of the left and right side is less than the width of the basal tubera (also in Piveteausaurus?).
(after Carrano et al., 2012) strongly inflated base of maxillary ascending process; evenly rounded ventral surfaces of most sacral centra, except sacral 3 bears flat midline strip and sacral 5 is broad and flat.
Comments- This genus has had numerous suggested affinities, from abelisaurid to allosaurid (see Comments under Piatnitzkysauridae).
References- Bonaparte, 1979. Dinosaurs: A Jurassic assembalge from Patagonia. Science. 205, 1377-1379.
Bonaparte, 1986. Les dinosaures (Carnosaures, Allosauridés, Sauropodes, Cétosauridés) du Jurassique Moyen de Cerro Cóndor (Chubut, Argentina). Annales de Paléontologie (Vert.-Invert.). 72(3), 247-289.
Rauhut, 2004. Braincase structure of the Middle Jurassic theropod dinosaur Piatnitzkysaurus. Canadian Journal of Earth Science. 41(9), 1109-1122.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

Megalosauria Bonaparte, 1850
Definition- (Megalosaurus bucklandii + Spinosaurus aegyptiacus) (Hendrickx, Hartman and Mateus, 2015)
Other definitions- (Dubreuillosaurus valesdunensis, Eustreptospondylus oxoniensis <- Spinosaurus aegyptiacus, Allosaurus fragilis, Passer domesticus) (Allain et al., 2012)
= Spinosauroidea sensu Sereno, 1998
Definition- (Spinosaurus aegyptiacus + Torvosaurus tanneri) (modified)
= Spinosauroidea sensu Sereno, in press
Definition- (Spinosaurus aegyptiacus + Torvosaurus tanneri, - Allosaurus fragilis, Passer domesticus)
Comments- Though originally a term equivalent to Theropoda, this term has been used for two different definitions recently. Allain et al. (2012) defined it as all taxa closer to Dubreuillosaurus valesdunensis and Eustreptospondylus oxoniensis than to Spinosaurus aegyptiacus, Allosaurus fragilis or Passer domesticus. This is equivalent to Megalosauridae in some phylogenies where it is valid, and doesn't include Megalosaurus in other phylogenies (e.g. Holtz, 2000). A month later, Carrano et al. (2012) used this name for the megalosaurid+spinosaurid node. While they did not define it, (Megalosaurus bucklandii + Spinosaurus aegyptiacus) would be the obvious definition and was used by Hendrickx et al. (2015). As this is a more useful clade and always includes its eponymous genus, the latter definition is used here.
References- Bonaparte, 1850. Conspectus Systematum Herpetologiae et Amphibiologiae. Editio Altera Reformata [Survey of the systems of reptiles and amphibians. Second revised edition]. E. J. Brill, Leyden. [pp unknown]
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften. 99(5), 369-377.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.

Streptospondylidae Kurzanov, 1989
Streptospondylus Meyer, 1832
= "Streptospondylus" Meyer, 1830
S. altdorfensis Meyer, 1832
= Streptospondylus rostromajor Owen, 1842
= Laelaps gallicus Cope, 1867
= Poekilopleuron gallicum (Cope, 1867; as Poecilopleurum gallicum) Cope, 1869
= Dryptosaurus gallicus (Cope, 1867) Olshevsky, 1991
Late Callovian-Early Oxfordian, Middle Jurassic-Late Jurassic
Vaches Noires Cliffs, Calvados, France
Lectotype- (MNHN 8605) (subadult) distal pubis
....(MNHN 8606) distal fibula
....(MNHN 8607) distal tibia (140 mm wide)
....(MNHN 8608) astragalus (114 mm wide)
....(MNHN 8609) calcaneum
....(MNHN 8787) tenth cervical vertebra, first dorsal vertebra (76 mm), second dorsal vertebra (64 mm), proximal dorsal rib
....(MNHN 8788) posterior part of fifth sacral vertebra, first caudal vertebra
....(MNHN 8789) fifth or sixth dorsal vertebra (74 mm)
....(MNHN 8789) posterior dorsal vertebra (97 mm)
....(MNHN 8793) fourth or fifth dorsal vertebra (72 mm)
....(MNHN 8794) postzygopophysis of twelfth dorsal vertebra, posterior half of thirteenth dorsal vertebra, first sacral vertebra (98 mm), anterior part of second sacral vertebra
....(MNHN 8907) three posterior dorsal vertebrae (90, 95 mm)
Referred- ?...(MNHN 9645) distal femur (Gaudry, 1890)
Diagnosis- (from Allain, 2001) two hypapophyses on anterior dorsal vertebrae; anterior dorsal centra strongly opisthocoelous and strongly flattened; posterior dorsal vertebrae platycoelous; elongate mid and posterior dorsal centra; lateral extension of the medial buttress above the dorsomedial edge of the ascending process of the astragalus doesn’t reach the median part of the distal end of the tibia; large depression at the base of the ascending process of the astragalus; lack of posteromedial process on the astragalus.
Comments- Allain (2001) states "the theropod vertebrae are designed here as a lectotype of this taxon" but then lists all material except the femur as the lectotype, making it uncertain exactly what the lectotype is.
This taxon was recently suggested to be the sister taxon of Eustreptospondylus (Allain, 2001; Smith et al., 2007) based on the presence of carotid processes in their anterior dorsal vertebrae. Benson (2008, 2010) found it to be a member of either Megalosauroidea or Carnosauria, until further data (Benson et al., 2010) placed it in Sinraptoridae sister to Lourinhanosaurus. Their most recent study (Carrano et al., 2012) instead has it a megalosaurian outside Spinosauridae, and Megalosaurinae+Afrovenatorinae (or when properly ordered, Megalosaurinae OR Afrovenatorinae). It only takes 2 more streps to make Streptospondylus an afrovenatorine (though in that case Streptospondylinae has priority for the subfamily name).
References- Meyer, 1830. uber fossile Saurier. Isis. 1830, 517-519.
Meyer, 1832. Paleologica zur Geschichte der Erde und ihrer Gashopfe. Frankfurt am Main. 560 pp.
Owen, 1842. Report on British fossil reptiles. Report of the British Association for the Advancement of Science. 11, 60-204.
Cope, 1867. Account of extinct reptiles which approach birds. Proceedings of the Academy of Natural Sciences of Philadelphia. 1867, 234-235.
Cope, 1869. Synopsis of the Extinct Batrachia and Reptilia of North America. Part 1. Transactions of the American Philosophical Society, New Series. 14, 252 pp.
Gaudry, 1890. Les Enchaînements du monde animal dans les temps géologiques. Fossiles secondaires. Savy, Paris. 322 pp.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Allain, 2001. Redescription of Streptospondylus altdorfensis, Cuvier’s theropod dinosaur, from the Jurassic of Normandy. Geodiversitas. 23(3), 349-367.
Allain, 2002. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. 329 pp.
Smith, Makovicky, Hammer and Currie, 2007. Osteology of Cryolophosaurus ellioti (Dinosauria: Theropoda) from the Early Jurassic of Antarctica and implications for early theropod evolution. Zoological Journal of the Linnean Society. 151, 377-421.
Benson, 2008. A new theropod phylogeny focussing on basal tetanurans, and its implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Benson, Brusatte and Carrano, 2010. A new clade of large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften. 97, 71-78.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

unnamed megalosaurian (Janensch, 1925)
Late Tithonian, Late Jurassic
Upper Dinosaur Member of the Tendaguru Formation, Tanzania
Material- (MB R 1926) astragalus (212 mm wide), calcanear fragment
....(MB R 3627; = MW 1) incomplete fibula
Comments- MB R 3627 was originally a syntype of Ceratosaurus roechlingi (Janensch, 1925), though he did not mention the probably associated tarsus. Not only are they too large for the roechlingi type, Rauhut (2011) assigned them to Megalosauroidea based on the basal tetanurine grade astragalus and absent proximomedial fibular fossa. The latter has since been optimized as a megalosaurian character. Other tetanurine material from Tendaguru (e.g. ilium MB R 3628) may belong to the same taxon.
References- Janensch, 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten Deutsch-Ostafrikas. Palaeontographica. 1(supp. 7), 1-99.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania). Palaeontology. 86, 195-239.

unnamed possible megalosaurian (El-Zouki, 1980)
Hauterivian-Barremian, Early Cretaceous
Cabao Formation, Jannawan, Libya
Material- posterior dorsal centrum
Comments- El-Zouki (1980) referred a vertebra to ?Spinosaurus sp. as a caudal, but Le Loeuff et al. (2010) note "the published plate (a vertebral centrum in ventral view) does not allow any precise assessment." Carrano et al. (2012) reinterpret it as a posterior dorsal centrum, noting "The proportions match well with the posterior dorsals of Baryonyx and Suchomimus but are relatively shorter than those of Megalosaurus and carcharodontosaurids. The (presumed) ventral surface is only slightly narrower than the centrum, similar to the condition in Megalosaurus and Suchomimus." They "consider it likely that this specimen belongs to the clade Megalosauria."
References- El-Zouki, 1980. Stratigraphy and lithofacies of the continental clastics (Upper Jurassic and Lower Cretaceous) of Jabal Nafusah, NW Libya. In Salem and Busrewil (eds.). The Geology of Libya, Vol. II. Academic Press. 393-418.
Le Loeuff, Métais, Dutheil, Rubinos, Buffetaut, Ois Lafont, Cavin, Moreau, Tong, Blanpied and Sbeta, 2010. An Early Cretaceous vertebrate assemblage from the Cabao Formation of NW Libya. Geological Magazine. 147(5), 750-759.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

Megalosauridae Huxley, 1869
Definition- (Megalosaurus bucklandii <- Spinosaurus aegyptiacus, Allosaurus fragilis, Passer domesticus) (Holtz et al., 2004)
Other definitions- (Torvosaurus tanneri + Afrovenator abakensis + Dubreuillosaurus valesdunensis) (modified from Allain, 2002)
(Megalosaurus bucklandii <- Spinosaurus aegyptiacus, Sinraptor dongi, Allosaurus fragilis, Carcharodontosaurus saharicus) (Benson, 2010)
= Eustreptospondylidae Paul, 1988
= Torvosauridae Jensen, 1985 sensu Sereno, 1998
Definition- (Torvosaurus tanneri <- Spinosaurus aegyptiacus) (modified)
= Megalosauridae sensu Benson, 2010
Definition- (Megalosaurus bucklandii <- Spinosaurus aegyptiacus, Sinraptor dongi, Allosaurus fragilis, Carcharodontosaurus saharicus)
= Megalosauria sensu Allain et al., 2012
Definition- (Dubreuillosaurus valesdunensis, Eustreptospondylus oxoniensis <- Spinosaurus aegyptiacus, Allosaurus fragilis, Passer domesticus)
= Torvosauridae sensu Sereno, in press
Definition- (Torvosaurus tanneri <- Spinosaurus aegyptiacus, Allosaurus fragilis, Passer domesticus)
Comments- Traditionally a large paraphyletic family containing most basal tetanurines and carnosaurs, Megalosauridae has been limited since the late 1980's to include far fewer taxa.
References- Jensen, 1985. Uncompahgre dinosaur fauna: A preliminary report. Great Basin Naturalist. 45, 710-720.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.

Streptospondylus? cuvieri Owen, 1842
= Megalosaurus cuvieri (Owen, 1842) Huene, 1908
Early Bajocian, Middle Jurassic
Inferior Oolite Formation, England
Holotype- (Kingdon coll.; lost) tooth, partial anterior dorsal vertebra (~114 mm), partial dorsal neural spine, broad flat bone, long bone fragments
Comments- Eustreptospondylus oxoniensis was referred to Streptospondylus cuvieri before it was named. This taxon is not necessary the same as Streptospondylus altdorfensis, which lived later in France. Owen (1842) also referred an anterior dorsal from the Toarcian Jet Rock Formation (Ripley coll.), but this is indeterminate past Tetanurae.
The convex anterior articular surface suggests a tetanurine, while the camerate interior excludes it from Carcharodontosauridae and Megaraptora. The absence of a ventral keel is unlike Piatnitzkysaurus+Condorraptor, megalosaurines+afrovenatorines, spinosaurids and metriacanthosaurines. The large pleurocoel (49% of anterior articular surface height) is like Streptospondylus and megalosaurids, so it is tentatively referred to that clade here. It thus may actually belong to Streptospondylus.
References- Owen, 1842. Report on British fossil reptiles. Report of the British Association for the Advancement of Science. 11, 60-204.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse. Geologische und Palaeontologische Abhandlungen. Suppl. 1(1), 1-419.

undescribed Megalosauridae (Lubbe, Richter and Knötschke, 2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry, Germany
Material- ?(DFMMh/FV 658) tooth (24.5x8.8x4.8 mm) (Lubbe, Richter and Knötschke, 2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry and/or Hannover, Germany
(DFMMH/FV and/or NLMH coll.) teeth
Comments- While seven teeth assigned to Velociraptorinae by Lubbe et al. (2009), Gerke and Wings (2014) found four of these were Neotheropoda indet., megalosaurid and tyrannosauroid. Based on the information in Lubbe et al., FV 658 may be the megalosaurid (larger, elongate crown, more of mesial carina serrated).
Reference- Lubbe, Richter and Knötschke, 2009. Velociraptorine dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany. Acta Palaeontologica Polonica. 54(3), 401-408. Gerke and Wings, 2014. Characters versus morphometrics: A case study with isolated theropod teeth from the Late Jurassic of Lower Saxony, Germany, reveals an astonishing diversity of theropod taxa. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 137.

unpublished megalosaurid (Siegwarth et al., unpublished)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of Morrison Formation, Wyoming, US
Material- (TATE coll.) ilium
Comments- This was found by "Brontoraptor" but smaller and different in morphology.
Reference- Siegwarth, Lindbeck, Redman, Southwell and Bakker, unpublished. Megalosaurid dinosaurs from the Late Jurassic Morrison Formation of Eastern Wyoming. 27 pp.

Eustreptospondylinae Paul, 1988
Definition- (Eustreptospondylus oxoniensis <- Megalosaurus bucklandii) (Holtz et al., 2004)
Other definitions- (Eustreptospondylus oxoniensis <- Torvosaurus tanneri, Spinosaurus aegyptiacus, Allosaurus fragilis) (Sereno, in press)
Comments- Holtz et al.'s (2004) definition only includes Megalosaurus bucklandii as an external specifier, and I agree with Sereno (in press) that additional ones are useful in case eustreptospondylines are closer to allosaurids than to Megalosaurus (Paul, 1988; Kurzanov, 1989; Molnar et al., 1990; Holtz, 2000). This is especially true considering Megalosaurus' uncertain placement among theropods.

Eustreptospondylus Walker, 1964
E. oxoniensis Walker, 1964
= Magnosaurus oxoniensis (Walker, 1964) Rauhut, 2003
Late Callovian, Middle Jurassic
Middle Oxford Clay Formation, England
Holotype- (OUM J13558) (4.63 m, 218 kg; subadult) (partial skull ~483 mm) premaxillae, incomplete maxillae, lacrimal, postorbitals, squamosal, quadrate, frontals, parietal, partial braincase, dentaries, teeth (lost), axis (~44 mm), third cervical vertebra (45 mm), fourth cervical vertebra, fifth cervical centrum (55 mm), sixth cervical centrum (60 mm), seventh cervical centrum (60 mm), eighth cervical vertebra (60 mm), ninth cervical vertebra (60 mm), tenth cervical vertebra (60 mm), first dorsal vertebra (60 mm), second dorsal centrum (62 mm), third dorsal centrum (62 mm), fourth dorsal vertebra, fifth dorsal vertebra (63 mm), sixth dorsal vertebra, seventh dorsal vertebra (74 mm), eighth dorsal centrum (77 mm), ninth dorsal vertebra (78 mm), tenth dorsal vertebra (82 mm), eleventh dorsal centrum (88 mm), first sacral vertebra, third sacral vertebra, fourth sacral vertebra, fifth sacral centrum, first caudal vertebra (~68 mm), second caudal vertebra, third caudal centrum, fourth caudal vertebra, partial fifth caudal centrum, seventh caudal centrum, eighth caudal vertebra, tenth caudal centrum (~68 mm), twelfth caudal centrum (~61 mm), thirteenth caudal centrum (~60 mm), sixteenth caudal vertebra, twenty-first caudal vertebra, scapula (299 mm), humerus (231 mm), ilium (365 mm), pubes (451 mm), ischia (358 mm), femora (498 mm), tibiae (479 mm), fibulae (467 mm), astragali (86 mm wide), calcaneum (lost), metatarsal II (~193 mm), phalanx II-1 (64 mm), phalanx II-2 (~51 mm), metatarsal III (232 mm), phalanx III-1 (80, 85mm), phalanx III-2 (65, 63mm), phalanx III-3 (54 mm), pedal ungual III (54 mm), metatarsal IV (207 mm), phalanx IV-1 (76 mm), phalanx IV-2 (53 mm)
Diagnosis- (after Rauhut, 2000) differs from Magnosaurus in the proximal extent of the pubic symphysis.
(after Sadleir et al., 2008) shallow lacrimal fenestra that incorporates a second smaller foramen; fossa on posterolateral surface of ventral postorbital process; squamosal with a hypertrophied ventral flange that overhangs the laterotemporal fenestra in lateral view, partially obscuring its posterodorsal corner; ventral keels absent on presacral vertebrae; marked depression on anterior part of ventral surface of tenth cervical vertebra.
differs from Magnosaurus in- having interdental plates which are longer than tall; pubis with less transverse expansion at acetabular margin in proximal view; lateral margin of distal femur straight in posterior view; dorsoventrally extending ridge on lateral surface of cnemial crest absent.
(after Carrano et al., 2012) pubic peduncle of ilium as broad anteroposteriorly as mediolaterally; lateral wall of iliac brevis fossa nearly horizontal, exposing medial wall of fossa along entire length in lateral view.
Comments- This specimen was originally described by Phillips (1871) as combining Streptospondylus and Megalosaurus attributes, but left unnamed. Nopcsa (1905, 1906) referred the specimen to Streptospondylus cuvieri in his description, while Huene referred the specimen to either Streptospondylus cuvieri (1907-08, 1923, 1926) or Megalosaurus cuvieri (1926, 1932). Walker (1964) erected the new genus Eustreptospondylus for the specimen, as it not referrable to Megalosaurus or Streptospondylus. Streptospondylus? cuvieri is now restricted to the lost holotype partial dorsal vertebra, which is an indeterminate theropod.
The jugal mentioned by Huene (1926) is probably the squamosal. The supposed parts of the coronoid and angular figured by Nopcsa (1906) are probably parts of the dentaries. Huene's (1926) fourteenth dorsal vertebra does not exist and his account of 29 preserved caudal vertebrae is in error as well. Phillips (1871) and Huene (1926) describe and illustrate a distal metacarpal and manual phalanx, which are either lost or misidentified pedal elements.
Though found as a basal megalosaurid by Carrano et al. (2012), only three more steps are needed to place it in Afrovenatorinae (which would then be called Eustreptospondylinae) as in Allain (2002), so either possibility is likely. Five more steps are needed to place Eustreptospondylus closer to spinosaurids than Afrovenator as in Sereno et al. (1994) and 7 more steps are needed to place it closer to spinosaurids than Torvosaurus or Afrovenator are, as in Smith et al. (2007). These possibilities are possible though less likely.
References- Phillips, 1871. Geology of Oxford and the Valley of the Thames. 529 pp.
Nopcsa, 1905. Notes on British dinosaurs. Part III: Streptospondylus. Geological Magazine. 5, 289-293.
Nopcsa, 1906. Zur kenntnis des Genus Streptospondylus. Beiträge zur Paläontologie Österreich-Ungarns und des Orients. 19, 59-83.
Huene, 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse. Geologische und Palaeontologische Abhandlungen Suppl. 1(1), 1-419.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of the Geological Society of America. 34, 449-458.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London B. 248, 53-134.
Sereno, Wilson, Larsson, Dutheil and Sues, 1994. Early Cretaceous dinosaurs from the Sahara. Science. 266, 267-271.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. 329 pp.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Sadleir, Barrett and Powell, 2004. Anatomy and systematics of Eustreptospondylus oxoniensis (Dinosdauria: Theropoda): Evolutionary implications. Journal of Vertebrate Paleontology. 24(3), 24A.
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.
Sadleir, Barrett and Powell, 2008. The anatomy and systematics of Eustreptospondylus oxoniensis, a theropod dinosaur from the Middle Jurassic from Oxfordshire, England. Monograph of the Palaeontological Society. 1-82.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.

= Megalosaurinae sensu Holtz et al., 2004
Definition- (Megalosaurus bucklandii <- Eustreptospondylus oxoniensis)

= Megalosauridae sensu Allain, 2002
Definition- (Torvosaurus tanneri + Afrovenator abakensis + Dubreuillosaurus valesdunensis) (modified)

Megalosaurinae Huxley, 1869 sensu Nopcsa, 1928
Definition- (Megalosaurus bucklandii <- Afrovenator abakensis) (modified from Carrano et al., 2012)
Other definitions- (Poekilopleuron bucklandii <- Torvosaurus tanneri) (modified from Allain, 2002)
(Megalosaurus bucklandii <- Eustreptospondylus oxoniensis) (Holtz et al., 2004)
= Torvosaurinae Jensen, 1985 vide Allain, 2002
Definition- (Torvosaurus tanneri <- Poekilopleuron bucklandii, Afrovenator abakensis) (modified from Allain, 2002)

Duriavenator Benson, 2008a
= "Walkersaurus" Welles and Powell, 1995 vide Welles, Powell and Pickering, 1994
D. hesperis (Waldman, 1974) Benson, 2008a
= Megalosaurus hesperis Waldman, 1974
= "Walkersaurus" hesperis (Waldman, 1974) Welles and Powell, 1995 vide Welles, Powell and Pickering, 1994
Late Bajocian, Middle Jurassic
Upper Inferior Oolite, England
Holotype- (NHMUK R332) (~5 m) partial premaxillae, maxilla, vomer, partial dentaries, partial surangular, teeth, fragments
Diagnosis- (after Benson, 2008a) deep groove on dorsal surface of jugal process containing numerous pneumatic foramina; array of small foramina in ventral part of articular surface for premaxilla.
Comments- This specimen was originally described by Owen (1883) and referred to Megalosaurus bucklandi. Walker (1964) noted it was probably a distinct species, due to tooth count and a supposed lateral groove between the premaxilla and maxilla. Waldman (1974) officially named the species Megalosaurus hesperis. The taxon was studied by Welles and Powell in the 1970's as part of their redescription of European theropods, where they intended to rename it Walkersaurus hesperis. This was not made publically available until Pickering revised it and sent it to a few colleagues in 1994, though due to its low circulation most paleontologists only learned about the name in 1999 from his 1995 description of Dilophosaurus "breedorum" (often cited as being from 1999) and his bibliographic work "Jurassic Park: Unauthorized Jewish Fractals in Philopatry". Both of these papers face similar problems to the 1994 work in that they do not conform to IZCN Article 8.1.3- it must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies. Thus "Walkersaurus" is a nomen nudum, though Pickering does plan to include it in his book "Mutanda Dinosaurologica". Holtz (2000) found the species to have several possible positions as a tetanurine less derived than Afrovenator + Avetheropoda in his cladistic analysis. Most recently, Benson (2008a) has redescribed the material and placed it in a new genus- Duriavenator. Benson (2008b, 2010) found Duriavenator to emerge as a megalosaurid in his phylogenetic analysis. Carreano et al. (2012) further found it to be a megalosaurine, but as only one more step is necessary to make it an afrovenatorine, this is highly uncertain.
References- Owen, 1883. On the skull of Megalosaurus. Quarterly Journal of the Geological Society of London. 39, 334-347.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London B. 248, 53-134.
Waldman, 1974. Megalosaurids from the Bajocian (Middle Jurassic) of Dorset. Palaeontology. 17, 325-339.
Welles, Powell and Pickering, 1994. An extract from: Archosauromorpha: Cladistics and osteologies. A Fractal Scaling in Dinosaurology Project. 10 pp.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry. A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola, California. 478 pp.
Welles and Pickering, 1995. An extract from: Archosauromorpha: Cladistics and osteologies. A Fractal Scaling in Dinosaurology Project. 70 pp.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. Gaia. 15, 5-61.
Benson, 2008a. A redescription of 'Megalosaurus' hesperis (Dinosauria, Theropoda) from the Inferior Oolite (Bajocian, Middle Jurassic) of Dorset, United Kingdom. Zootaxa. 1931, 57-67.
Benson, 2008b. A new theropod phylogeny focusing on basal tetanurans and its implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.
Pickering, in prep. Mutanda Dinosaurologica.

Megalosaurus Buckland, 1824
= "Megalosaurus" Parkinson, 1822
M. bucklandii Mantell, 1827
= Megalosaurus "conybeari" Ritgen, 1826
= Megalosaurus bucklandi Meyer, 1832
= Metriacanthosaurus "brevis" Welles, Powell and Pickering vide Pickering 1995
Middle Bathonian, Middle Jurassic
Taynton Limestone Formation (=Stonesfield Slate), England
Lectotype- (OUM J13505) anterior dentary, teeth
Paralectotypes- (OUM J13561) incomplete femur (860 mm)
(OUM J13563) incomplete pubis
(OUM J13565) incomplete ischium (610 mm)
(OUM J13572) distal metatarsal II
(OUM J13576) incomplete sacrum (113, 108, 99, 108, 113 mm)
(OUM J13577) incomplete posterior dorsal vertebra (114 mm)
(OUM J13579) incomplete proximal caudal vertebra (99 mm)
(OUM J13580) posterior dorsal rib
(OUM J13585) posterior cervical rib
(OUM J29881) ilium
(OUM coll.; lost) tooth
Referred- (CAMSM Woodwardian collection D.11.35.c) partial scapula (Benson, 2009b)
?(Duke of Marlborough coll.) partial dentary (lost) (Owen, 1857)
(GSM 57809) metatarsal II (Benson, 2009b)
(GSM 113081) tooth (Benson, 2009b)
(MNHN 9630) incomplete femur (~745 mm) (Day and Barrett, 2004)
(NHMUK 2016) fragmentary humerus (Benson, 2009b)
(NHMUK 2331) tooth (Benson, 2009b)
(NHMUK 25581) partial mid caudal centrum (Lydekker, 1888)
?(NHMUK 25582) proximal ischium (Lydekker, 1888)
?(NHMUK 28301) fragmentary ?pubis (Lydekker, 1888)
?(NHMUK 28608) tooth (Lydekker, 1888)
(NHMUK 28957) two sacral vertebrae (Lydekker, 1888)
....(NHMUK R1098) partial sacrum (Owen, 1857)
(NHMUK 31804) incomplete femur (810.0 mm) (Lydekker, 1888)
(NHMUK 31805) tibia (Benson, 2010)
(NHMUK 31806) femur (805 mm) (Owen, 1857)
(NHMUK 31808) incomplete femur (820.0 mm) (Lydekker, 1888)
(NHMUK 31809) tibia (645 mm) (Owen, 1857)
(NHMUK 31810) coracoid (150 mm) (Lydekker, 1888)
(NHMUK 31811; intended holotype of Metriacanthosaurus "brevis") partial ilium (Lydekker, 1888)
(NHMUK 31812) partial mid caudal vertebra (Benson, 2009b)
(NHMUK 31813) posterior dorsal vertebra (Lydekker, 1888)
(NHMUK 31824) dorsal rib (Lydekker, 1888)
(NHMUK 31825) dorsal rib (Lydekker, 1888)
(NHMUK 31828) jugal (Welles and Pickering, 1999)
?(NHMUK 31834) tooth (Lydekker, 1888)
?(NHMUK 31932) pedal phalanx (Lydekker, 1888)
(NHMUK 33229) mid caudal vertebra (Benson, 2009b)
(NHMUK 36585) ulna (232 mm) (Lydekker, 1888)
(NHMUK 37303) dorsal rib (Lydekker, 1888)
(NHMUK 40127) tooth, tooth fragment, anterior tooth (Benson, 2009b)
(NHMUK 40128a; = 40125a in Lydekker, 1888?) metatarsal IV (Phillips, 1871)
(NHMUK 40131) partial coracoid (Lydekker, 1888)
(NHMUK 41305) partial tooth (Lydekker, 1888)
(NHMUK 42024) tooth (Lydekker, 1888)
(NHMUK 44097) dorsal rib (Lydekker, 1888)
(NHMUK 44097a) dorsal rib (Lydekker, 1888)
(NHMUK 47963) tooth, tooth tip(Lydekker, 1888)
(NHMUK R234) two teeth (Lydekker, 1888)
(NHMUK R283) partial ilium (Lydekker, 1888)
?(NHMUK R285) cervical vertebra (Lydekker, 1888)
(NHMUK R700) partial sacrum (Lydekker, 1888)
(NHMUK R1099) incomplete scapulocoracoid (830 mm; scapula 705 mm) (Lydekker, 1888)
(NHMUK R1100) ilium (770 mm) (Owen, 1857)
(NHMUK R1101) ilium (~832 mm) (Lydekker, 1888)
(NHMUK R1102) tibia (700.0 mm) (Lydekker, 1888)
(NHMUK R1103) distal tibia (Lydekker, 1888)
(NHMUK R12557) tibia (Benson, 2010)
(OUM J13506) maxilla (451 mm) (Huxley, 1869)
(OUM J13559) incomplete maxilla (Phillips, 1871)
(OUM J13560) ilium (~765 mm) (Walker, 1964)
(OUM J13562) tibia (660 mm) (Galton and Molnar, 2005)
(OUM J13567) incomplete pubis (Benson, 2009b)
(OUM J13568) incomplete tibia (722 mm) (Benson, 2009b)
(OUM J13569; cast as NHMUK R1104) metatarsal III (335 mm) (Phillips, 1871)
(OUM J13573) distal metatarsal II (Phillips, 1871)
(OUM J13574) scapulocoracoid (845 mm; scapula 704 mm) (Phillips, 1871)
(OUM J13575) humerus (388 mm) (Phillips, 1871)
(OUM J13578) proximal caudal vertebra (108 mm) (Phillips, 1871)
(OUM J13582) mid dorsal rib (Benson, 2010)
(OUM J13583) mid dorsal rib (Benson, 2010)
(OUM J13584) dorsal rib (Benson, 2010)
(OUM J29751) rib (Benson, 2010)
(OUM J29752) rib (Benson, 2010)
(OUM J29753a) proximal femur (Day and Barrett, 2004)
(OUM J29754) proximal femur (Benson, 2010)
(OUM J29758) distal tibia (Benson, 2010)
(OUM J29761) incomplete humerus (Phillips, 1871)
(OUM J29762) tooth (Benson, 2009b)
?(OUM 29764) long bone shaft (Benson, 2010)
(OUM J29766) sacral fragment (Benson, 2010)
(OUM J29767) sacral fragment (Benson, 2010)
(OUM J29768) proximal caudal centrum (Phillips, 1871)
(OUM J29777) basal tooth (Benson, 2009b)
(OUM J29779) pedal ungual (102 mm) (Phillips, 1871)
(OUM J29792) dorsal rib (Benson et al., 2008)
(OUM J29802) incomplete femur (690.0 mm) (Day and Barrett, 2004)
(OUM J29803) incomplete femur (740.0 mm) (Day and Barrett, 2004)
(OUM J29809) tooth (Benson, 2009b)
(OUM J29810) apical tooth (Benson, 2009b)
(OUM J29813) posterior mandible (Benson, 2010)
(OUM J29863) tooth (Benson, 2009b)
(OUM J29864) tooth fragment (Benson, 2009b)
(OUM J29869) distal humerus (Benson, 2010)
(OUM J29872) ischial shaft (Phillips, 1871)
(OUM J29873) proximal ischium (Benson, 2010)
(OUM J29874) scapula (Benson, 2010)
(OUM J29875; = OUM mount of Carrano, 1998?) tibia (693 mm?) (Benson, 2009b)
(OUM J29879) incomplete scapula (765 mm) (Day and Barrett, 2004)
(OUM J29880) femoral fragment (Benson, 2010)
(OUM J29882) ilium (Benson, 2009b)
(OUM J29883) partial ilium (Benson, 2010)
(OUM J29884) partial ilium (Benson, 2010)
(OUM J29885) partial ilium (Benson, 2010)
(OUM J29888) partial scapulocoracoid (795 mm) (Day and Barrett, 2004)
(OUM J29889) partial scapulocoracoid (Day and Barrett, 2004)
(OUM J29890) scapula (Benson, 2010)
(OUM J29891) scapulocoracoid (Benson, 2009b)
(OUM J29892) rib (Benson, 2010)
(OUM J29893) dorsal rib (Benson, 2010)
(OUM J29895) rib (Benson, 2010)
(OUM J48171) tooth (Benson, 2009b)
(OUM coll.) chevron (169 mm) (Huene, 1926)
Bathonian, Middle Jurassic
Great Oolite Group, England
(NHMUK R413) metatarsal III (338 mm) (Lydekker, 1888)
Early Bathonian, Middle Jurassic
Chipping Norton Limestone Formation, England
(NHMUK R8303) incomplete maxilla (Welles and Pickering, 1999)
(NHMUK R8304) partial dentary (Welles and Pickering, 1999)
(NHMUK R8305) partial dentary (Benson, 2009b)
(NHMUK R9665) metatarsal III (Welles and Pickering, 1999)
(NHMUK R9666) proximal metatarsal IV (Benson, 2009b)
(NHMUK R9668) proximal ischium (Welles and Pickering, 1999)
(NHMUK R9669) dorsal rib (Benson, 2010)
(NHMUK R9672) incomplete proximal caudal vertebra (124 mm) (Reynolds, 1939)
(NHMUK R9673) proximal caudal vertebra (118 mm) (Reynolds, 1939)
(NHMUK R9674) partial anterior cervical vertebra (Benson, 2010)
(NHMUK R9675) incomplete mid caudal vertebra (Reynolds, 1939)
(NHMUK R9676) mid caudal vertebra (Welles and Pickering, 1999)
(NHMUK R9677) incomplete proximal caudal vertebra (Welles and Pickering, 1999)
(NHMUK R9678) partial anterior dorsal vertebra (Benson, 2010)
?(NHMUK R9679) sacrum (Welles and Pickering, 1999)
?(NHMUK R9680) sacrum (Welles and Pickering, 1999)
?(NHMUK coll.) ribs, bone fragments (Benson, 2010)
(SDM 44.1) partial maxilla (Reynolds, 1939)
(SDM 44.4) partial sacrum (Reynolds, 1939)
(SDM 44.3) tooth (Benson, 2009b)
(SDM 44.5) incomplete mid caudal vertebra (82.5 mm) (Reynolds, 1939)
(SDM 44.6) posterior dorsal centrum (119.5 mm) (Reynolds, 1939)
(SDM 44.7) proximal caudal vertebra (Welles and Pickering, 1999)
?(SDM 44.10) dorsal vertebra (131 mm) (Reynolds, 1939)
(SDM 44.13) dorsal rib (Reynolds, 1939)
(SDM 44.14) coracoid (Reynolds, 1939)
(SDM 44.15) coracoid (Reynolds, 1939)
(SDM 44.18) incomplete humerus (Reynolds, 1939)
(SDM 44.20) incomplete ischium (Reynolds, 1939)
(SDM 44.21) distal ischium (Reynolds, 1939)
(SDM 44.23) femur (785 mm) (Day and Barrett, 2004)
(SDM 44.24) femur (800 mm) (Reynolds, 1939)
(SDM 44.25) distal metatarsal IV (Reynolds, 1939)
Early-Middle Bathonian, Middle Jurassic
Sharp's Hill Formation, England
(OUM J.29800) incomplete scapula (Welles and Pickering, 1999)
Jurassic?
England
?(Royal College of Surgeons coll.) tooth, tooth fragments, femur, partial femur, tibia, phalanx, bone fragment (Owen, 1954)
?(GPIT 18392?) proximal femur, fibula (Welles and Pickering, 1999)
?(GSM 3887) sacrum (Welles and Pickering, 1999)
?(OUM J12142) partial mandible, teeth (Delair, 1975)
?(OUM J13598) tooth (Benton and Spencer, 1995)
?(OUM J13882) tooth (Benton and Spencer, 1995)
?(OUM J29765) proximal scapula (Benton and Spencer, 1995)
?(OUM J29773) tooth (Benton and Spencer, 1995)
?(OUM J29797) partial pubis (Reid, 1985)
?(Phillips coll.) pedal ungual (lost) (Welles and Pickering, 1999)
?(RCS 72) femur (lost) (Welles and Pickering, 1999)
?(RCS 73) partial femur (lost) (Welles and Pickering, 1999)
?(RCS 74) distal tibia (lost) (Welles and Pickering, 1999)
? posterior cervical centrum (60 mm) (Phillips, 1871)
Diagnosis- (after Walker, 1964) dorsally directed flange around midheight on the scapular blade.
(after Benson, 2009b) lateral dentary groove broad (also in Torvosaurus); ventral surface of second sacral centrum bearing longitudinal, angular ridge (also in "Brontoraptor"); an array of posterodorsally inclined grooves on the lateral surface of the median iliac ridge (also in "Metriacanthosaurus" "reynoldsi"); anteroposteriorly thick ischial apron with an almost flat medial surface; a prominent, rugose distal ischial tubercle; and complementary groove and ridge structures on the articular surfaces between metatarsals II and III.
(after Carrano et al., 2012) dorsally directed flange at mid-height of scapular blade.
Other diagnoses- Benson et al. (2008) stated the longitudinal groove in the ventral part of the lateral surface of the dentary was an autapomorphy, but Benson (2010) found this to be due to damage. He also cited the slit-like foramen anterior to the termination of the Meckelian groove, but in 2010 reported this was prone to individual variation in Allosaurus fragilis. The presence of 13-14 dentary teeth (which is estimated from the 11 preserved in the lectotype) is also true in other megalosaurids- ~14(11+) in Magnosaurus, 13 in Dubreuillosaurus and Eustreptospondylus, and ~14-15(13+) in Duriavenator.
Benson (2009, 2010) reported a unique combination of dentary characters to distinguish the lectotype, but most of these are plesiomorphic. The lack of an enlarged third dentary alveolus is also found in Monolophosaurus and most non-megalosauroids. The dentary is straight in dorsal view in all megalosaurids and is not more transversely expanded anteriorly in Duriavenator, Eustreptospondylus or Dubreuillosaurus. Duriavenator and Magnosaurus also have tall dentary interdental plates. Those of all megalosaurids are unfused. All megalosaurids also have two Meckelian foramina and shallow Meckelian grooves. Benson also proposed postcranial autapomorphies. The sacral centra 1-3 and 5 of most non-maniraptoriform theropods are evenly rounded.
Comments- The lectotype dentary was collected in 1797. Parkinson (1822) used the name "Megalosaurus", but without description or indication of type material, making it a nomen nudum. The same can be said of Ritgen's (1826) species Megalosaurus "conybeari". Molnar et al. (1990) incorrectly attributed the species M. bucklandii to Ritgen, 1826. Meyer (1832) used the spelling bucklandi for the species, and has been followed by numerous authors, but this is incorrect. Molnar et al. designated OUM J13505 as the lectotype, making the additional elements described by Buckland (1824) paralectotypes.
Buckland (1824) described the paralectotype pubis as a fibula, and ischium as a clavicle (Phillips, 1871). Owen (1857) followed the ischial identification, described an Iguanodon ischium as the scapula, and the ilia as coracoids. Owen incorrectly stated the partial sacrum NHMUK R1098 is from the Wealden. Philips also reports Cuvier believed the metatarsal to be a humerus. He also described two elements as a scapula and ischium (= sacral rib?), but these are lost and may not belong to theropods (Benson, 2010). Phillips believed the proximal caudal vertebra was a posterior dorsal vertebra. The ulna (OUM J36585) was misidentified as a sacral rib by Lydekker (1888). Huene (1926) incorrectly listed two mandibles as being preserved in the type material, and listed the Kimmeridge Clay metatarsus (OUM J13586) and tibia (OUM J29886) as being from the Stonesfield Slate. The partial maxilla SDM 44.1 was misidentified by Reynolds (1939) as a dentary. Holtz (1994) lists OUM J13561 as being a femur and tibia, but the tibia is OUM J13562. While Welles and Pickering (1999) listed GPIT 18392 as a partial Megalosaurus hindlimb, this number has been associated in the literature with a Sellosaurus specimen. Benson et al. (2008) incorrectly stated rib OUM J29792 was part of the paralectotype.
Not Megalosaurus bucklandii- Owen (1857) figured several teeth as M. bucklandi (NHMUK 2828, 2332, 3222-3225), but these are from Cretaceous deposits and are probably indeterminate theropods. A large unfused scapulocoracoid mentioned by Phillips (1871) as being from a different species of megalosaur is possibly OMNH J29887, a proximal scapula which Day and Barrett (2004) and Benson (2009) reidentified as a sauropod. Phillips (1871) illustrated a metatarsus (OUM J13586) as Megalosaurus that was later illustrated by Huene (1926) with an associated tibia (OUM J29886) as M. bucklandii. Welles and Powell recognized these limb bones were distinct in the 1970's, and the nomen nudum Megalosaurus phillipsi was later assigned to the taxon by Pickering (1995). It is here suggested to be more closely related to Torvosaurus and described in its own entry. Owen (1883) described a partial skull as Megalosaurus bucklandi, but this was later named Megalosaurus hesperis by Waldman (1974) and placed it its own genus (Duriavenator) by Benson (2008). Lydekker (1888) referred several specimens to M. bucklandi- an incomplete sacrum (museum of the Geological Society collection) from the Coral Rag; two teeth (NHMUK 47152, R497) from the Inferior Oolite; a tooth (NHMUK 28608) from the Cotswold Slates; a dorsal centrum (NHMUK 47169) from the Cornbrash Formation; a tooth (NHMUK 39476) from the Forest Marble Formation; a dorsal vertebra (NHMUK 42028) from the Potton Sands; and a proximal tibia (NHMUK 32725) from the Vaches Noires Cliffs. These are all indeterminate theropods. He also claimed Owen (1857) referred the Cretaceous theropod manual ungual NHMUK R1105 and distal pedal ungual NHMUK R2482 to M. bucklandi, but Owen's plate caption merely says "Megalosaurus (?)". Huene (1906) described a partial braincase from the Taynton Limestone as Megalosaurus bucklandi, but this was reidentified as Cetiosaurus by Woodward (1910), a conclusion confirmed by Galton and Knoll (2006). Reynolds (1939) referred numerous specimens to Megalosaurus, of which ilium SDM 44.19 is a different taxon referred to as Metriacanthosaurus "reynoldsi" by Pickering (1995), while scapulae SDM 44.16 and 44.17, a tooth from Oakham Quarry and humerus SDM 44.22 are less diagnostic.
Megalosaurus morphotypes A and B- The disarticulated nature of Megalosaurus remains and lack of proper description until recently have led to suggestions that more than one taxon could be represented, and that only the lectotype dentary should be referred to the genus (e.g. Rauhut, 2000; Allain and Chure, 2002; Benson et al., 2008). Welles and Powell were among the first to suggest this, based on parts of their 1970's manuscript that have been subsequently released by Pickering (Welles and Pickering, 1999). They separated Chipping Norton remains as Metriacanthosaurus reynoldsi and some Taynton Limestone remains as Metriacanthosaurus brevis. These issues are discussed below. Allain and Chure (2002) noted the femur NHMUK 31806 differs from the paralectotype OUM J13561 in being straight with a medially directed head. This was expanded on by Day and Barrett (2004), who grouped Megalosaurus femora in two morphotypes. Morphotype A included NHMUK 31804 and 31806, MHHN 9630, OUM J29753a and J29802, and SDM 44.24. They are also distinguished by a distomedially oriented buttress supporting the anterior trochanter, a narrow ridge connecting the anterior trochanter to the femoral head, a shallow extensor groove, and subequally sized distal condyles. Morphotype B femora included NHMUK 31808, OUM J13561 and J29803, and SDM 44.23. Benson (2009) reviewed these supposed differences. He found the femoral head is always anteromedially oriented, though the more anterior angle in morphotype B femora (~40 degrees) is due to damage and deformation, seen in morphotype A femur NHMUK 31804 as well. The difference in curvature is attributed to distortion of originally straight shafts, with similar variation seen in Cleveland-Lloyd Allosaurus. The distomedial buttress is identified as the anterior intermuscular line and is found to not correspond to femoral types, with morphotype A femur NHMUK 31806 lacking it and morphotype B femora NHMUK 31808 and OUM J13561 having them. Individual variation is also noted for this feature in Dry Mesa Ceratosaurus. The ridge connecting the anterior trochanter and femoral head is present in all well preserved specimens, including morphotype B femur SDM 44.23. The extensor groove is present in all specimens as well, though shallower in OUM J13561 due to erosion of the anterior condyles. Finally, the distal condyles have similar proportions in most specimens, with a broader lateral condyle but longer medial condyle, with the exception of OUM J29803. The latter specimen is highly crushed transversely, explaining the difference. Benson concluded only one taxon was represented by the femora and most other Taynton Limestone and New Park Quarry cranial and postcranial material, which led to him referring all of it to M. bucklandii in his later (2010) osteology of the taxon.
A Stonesfield Slate sinraptorid? Bakker et al. (1992) thought maxilla OUM J13506 was from a sinraptorid due to its short anterior ramus, while Naish and Martill (2007) noted this was primitive and suggested it could be an abelisauroid instead. Note this would be consistant with Welles and Pickering' idea of Metriacanthosaurus "brevis" being in that formation, though they refer the maxilla to Megalosaurus. Benson (2007) noted it differed from sinraptorids in having a maxillary fossa instead of fenestra, and from ceratosaurs in having unfused interdental plates, but resembled other jaw elements of Megalosaurus.
Metriacanthosaurus brevis- Owen (1857) originally described Megalosaurus ilia as coracoids, noting three specimens were in the NHMUK and figuring one. Lydekker (1888) recognized these as ilia and incorrectly listed the one illustrated by Owen as NHMUK 31811 (as BMNH 31811), though Pickering (DML, 2002) notes it is actually NHMUK R1100 (as BMNH R1100). NHMUK 31811 remained as a specimen of Megalosaurus bucklandii until Pickering credited the name Metriacanthosaurus brevis to English theropod material in his 1995 unpublished bibliographic manuscript. It was later used in the comparative section of another unpublished manuscript (Welles and Pickering, 1999). This paper was largely extracted from the European megalosaur manuscript Welles and Powell worked on in the 1970s but never published, specifically the Megalosaurus redescription section. Pickering intends to publish an updated version of the megalosaur manuscript as Mutanda Dinosaurologica, and has posted small excerpts including the diagnosis of Metriacanthosaurus "brevis" online (DML, 2002). In any case, the name is a nomen nudum as Pickering didn't follow ICZN Article 8.1.3- it must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies. Pickering says of M. "brevis" and Metriacanthosaurus parkeri, "their great height and shortness -- diagnostic for Metriacanthosaurus -- separates them from ilia of Megalosaurus bucklandii and Allosaurus." Additionally, Allain and Chure (2002) stated NHMUK 31811 and R1100 (misidentified as OUM J13560) are "quite different from one another in shape in proportions." However, Day and Barrett (2004) found the shortness was an illusion caused by a broken postacetabular process and a plaster reconstructed preacetabular process. Benson (2009) agreed and also found it shared an autapomorphy with other M. bucklandii ilia- a series of posterodorsally oriented ridges that give the posterior part of the median ridge an undulating texture. Indeed, the characters Pickering lists as distinguishing M. "brevis" from M. parkeri are largely also those that distinguish M. bucklandii from M. parkeri (nearly straight upper margin; taller preacetabular process), or don't distinguish M. "brevis" from M. bucklandii (narrower notch between preacetabular process and pubic peduncle; longer, lower acetabulum; 250 mm long). So I see no reason to support M. "brevis" and synonymize it with M. bucklandii. Pickering also referred several other specimens to M. "brevis". OUM J29888 is a pectoral girdle which Day and Barrett and Benson couldn't distinguish from M. bucklandii. NHMUK 31806 is a femur which Allain and Chure stated differs from the paralectotype in being straight with a medially directed head. Day and Barrett (2004) used this as the basis for their morphotype A femora, but Benson (2009) found the head orientation to be individual variation and the curvature of type B femora to be taphonomic. Additional differences between morphotype A and B femora were also found to be preservational or individual variation (see below), and Benson (2009, 2010) referred both types to M. bucklandii. Ironically, M. parkeri's femur has a sigmoid shaft, though it does have a medially directed head. NHMUK 31809 and OUM J13562 are tibiae which Benson (2009, 2010) referred to M. bucklandii and which share a bulbous fibular crest with that taxon. NHMUK 31809 differs from M. parkeri in having a smaller cnemial crest which is not as laterally angled and a smaller and more laterally placed fibular crest. Thus all supposed Metriacanthosaurus "brevis" material is referrable to Megalosaurus bucklandii and does not bear particular resemblence to Metriacanthosaurus itself. Coincidentally, NHMUK 31806 and 31809 are the femur and tibia illustrated as examples of those elements in Benson's (2010) redescription of M. bucklandii, though the other tibia and the pectoral girdle remain unillustrated.
Chipping Norton remains- Lydekker (1888) referred a metatarsal III (NHMUK R413) from the Chipping Norton Formation to Megalosaurus bucklandi. Gardiner (1937, 1938) and Reynolds (1938) reported large theropod remains from two quarries in the Chipping Norton Limestone Formation, which Reynolds (1939) described and referred to Megalosaurus. Reynolds misidentified maxilla SDM 44.1 as a dentary, dorsal centrum SDM 44.6 as a caudal. Pickering (1995) credited the name Metriacanthosaurus reynoldsi to Welles, Powell and Pickering in his unpublished 1995 manuscript and later used it in the comparative section of Welles and Pickering, 1999. It is a nomen nudum in both though, as Pickering didn't follow ICZN Article 8.1.3- it must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies. The 1999 paper shows his new taxon to be based on all the Chipping Norton theropod material from both quarries, as well as NHMUK R413 and a few other elements. Day and Barrett (2004) believed both their femoral morphotypes A and B were present in the sample- SDM 44.23 as morphotype B and SDM 44.24 as morphotype A. Benson (2009, 2010) referred the New Park Quarry material to Megalosaurus bucklandii based on the close resemblence of maxilla SDM 44.1 to Taynton Limestone specimens and an M. bucklandii autapomorphy in sacrum SDM 44.4, while other elements were provisionally referred as there is no evidence of more than one taxon in the quarry. Additional New Park Quarry material referred by Welles and Pickering to "reynoldsi" but not mentioned by Benson is here provisionally referred to Megalosaurus- sacra NHMUK R9679 and R9680 and dorsal SDM 44.10. Benson (2010) also referred two specimens from Oakham Quarry to M. bucklandii based on autapomorphies (ischium SDM 44.20 and metatarsal III NHMUK R9665), but refrained from referring additional elements from this quarry as he notes some "can be referred to a second, unnamed taxon (R. B. J. Benson, unpubl. data)." This near certainly refers to "reynoldsi", whose intended holotype is an ilium from Oakham Quarry which resembles Eustreptospondylus and Yangchuanosaurus zigongensis most closely. Further study by Benson and/or Pickering may clarify the identity of the other Oakham Quarry elements.
References- Parkinson, 1822. Outlines of Oryctology. An introduction to the study of fossil organic remains, especially those found in the British Strata. London. 350pp.
Buckland, 1824. Notice on the Megalosaurus or great fossil lizard of Stonesfield. Transactions of the Geological Society of London. 21, 390-397.
Ritgen, 1826. Versuchte Herstellung einiger Becken urweltlichter Thiere. Nova Acta Caesareaa Leopold.-Carol. Ger. Nat. Curiosorum. 13, 331-358.
Mantell, 1827. Illustrations of the geology of Sussex. London: Lupton Relfe. 92 pp.
Meyer, 1832. Palaeologica zur Geschichte der Erde. Frankfurt am Main: S. Schmerber. 560 pp.
Owen, 1854. Descriptive catalogue of the fossil organic remains of Reptilia and Pisces contained in the Museum of The Royal College of Surgeons of England. Taylor and Francis, London. 184 pp.
Owen, 1857. Monograph on the Fossil Reptilia of the Wealden Formations. Part III. Dinosauria (Megalosaurus). Palaeontographical Society Monographs. 34, 1-26.
Huxley, 1869. On the upper jaw of Megalosaurus. Quarterly Journal of the Geological Society. 25, 311-314.
Huxley, 1870. Further evidence of the affinity between the dinosaurian reptiles and birds. Quarterly Journal of the Geological Society. 26, 12-31.
Phillips, 1871. Geology of Oxford and the Valley of the Thames. Oxford at the Clarendon Press. 523 pp.
Hulke, 1879. Note on Poikilopleuron bucklandi of Eudes Deslongchamps (père), identifying it with Megalosaurus bucklandi. Quarterly Journal of the Geological Society. 35(1-4), 233-238.
Owen, 1883. On the skull of Megalosaurus. Quarterly Journal of the Geological Society of London. 39, 334-347.
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.
Huene, 1906. Ueber das Hinterhaupt von Megalosaurus bucklandi aus Stonesfield. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie. 1906, 1-12.
Woodward, 1910. On a skull of Megalosaurus from the Great Oolite of Minchinhampton (Gloucestershire). Quarterly Journal of the Geological Society of London. 66, 111-15.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of the Geological Society of America. 34, 449-458.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Gardiner, 1937. Reptile-bearing oolite, Stow. Reports of the British Association for the Advancement of Science (Blackpool). 1936, 296.
Gardiner, 1938. Reptile-bearing oolite, Stow. Reports of the British Association for the Advancement of Science (Nottingham). 1937, 290.
Reynolds, 1938. A collection of reptilian bones from the Oölite near Stow-in-the-Wold, Glos. Reports of the British Association for the Advancement of Science. 1937, 356-357.
Reynolds, 1939. A collection of reptile bones from the Oolite near Stow-on-the-Wold, Gloucestershire. Geological Magazine. 76, 193-214.
Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London B. 248, 53-134.
Waldman, 1974. Megalosaurids from the Bajocian (Middle Jurassic) of Dorset. Palaeontology. 17, 325-339.
Delair and Sarjeant, 1975. The earliest discoveries of dinosaurs. Isis. 66(231), 5-25.
Reid, 1985. On supposed Haversian bone from the hadrosaur Anatosaurus, and the nature of compact bone in dinosaurs. Journal of Paleontology. 59, 140-148.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford. 169-209.
Bakker, Kralis, Siegwarth and Filla, 1992. Edmarka rex, a new, gigantic theropod dinosaur from the middle Morrison Formation, Late Jurassic of the Como Bluff outcrop, with comments on the evolution of the chest region and shoulder in theropods and birds and a discussion of the five cycles of origin and extinction among giant dinosaurian predators. Hunteria. 2(9), 1-24.
Benton and Spencer, 1995. Fossil Reptiles of Great Britain. Geological Conservation Review Series, Chapman & Hall. 386 pp.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry. A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola, California. 478 pp.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology, biomechanics, and modern analogs. PhD thesis, The University of Chicago. 424 pp.
Rayfield, 1998. Finite element analysis of the snout of Megalosaurus bucklandi. Journal of Vertebrate Paleontology. 18(3), 71A.
Welles and Pickering, 1999. Megalosaurus bucklandii. Private publication of Stephen Pickering, An extract from Archosauromorpha: Cladistics & Osteologies. A Fractal Scaling in Dinosaurology Project. 119 pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. 329 pp.
Allain and Chure, 2002. Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy. Palaeontology. 45, 1107-1121.
Pickering, DML 2002. https://web.archive.org/web/20191030094450/http://dml.cmnh.org/2002Mar/msg00553.html
Day and Barrett, 2004. Material referred to Megalosaurus (Dinosauria: Theropoda) from the Middle Jurassic of Stonesfield, Oxfordshire, England: One taxon or two? Proceedings of the Geologists' Association. 115, 359-366.
Galton and Molnar, 2005. Tibiae of small theropod dinosaurs from Southern England, from the Middle Jurassic of Stonesfield near Oxford and the Lower Cretaceous of the Isle of Wight. In Carpenter (ed.). The Carnivorous Dinosaurs. 3-22.
Benson, 2006. The taxonomy, systematics and evolution of the British theropod dinosaur, Megalosaurus. Journal of Vertebrate Paleontology. 27(3), 41A.
Galton and Knoll, 2006. A saurischian dinosaur braincase from the Middle Jurassic (Bathonian) near Oxford, England: From the theropod Megalosaurus or the sauropod Cetiosaurus? Geological Magazine. 143(6), 905-921.
Benson, 2008. A redescription of 'Megalosaurus' hesperis (Dinosauria, Theropoda) from the Inferior Oolite (Bajocian, Middle Jurassic) of Dorset, United Kingdom. Zootaxa. 1931, 57-67.
Benson, Barrett, Powell and Norman, 2008. The taxonomic status of Megalosaurus bucklandii (Dinosauria, Theropoda) from the Middle Jurassic of Oxfordshire, UK. Paleontology. 51, 419-424.
Benson, 2009a. The taxonomy, systematics and evolution of the British theropod dinosaur Megalosaurus. PhD thesis, University of Cambridge. [pp]
Benson, 2009b. An assessment of variability in theropod dinosaur remains from the Bathonian (Middle Jurassic) of Stonesfield and New Park Quarry, UK and taxonomic implications for Megalosaurus bucklandii and Iliosuchus incognitus. Palaeontology. 52(4), 857-877.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.
Pickering, in prep. Mutanda Dinosaurologica.
M? sp. indet. (Owen, 1857)
Early Jurassic
Coral Rag of Dry Sandford, England
Material- (GSL coll.; cast as NHMUK R1027) incomplete sacrum (first vertebra 127 mm) (Owen, 1857)
teeth (Huene, 1932)
Comments- Owen (1857) described the sacrum as Megalosaurus, and Lydekker (1888) referred it to M. bucklandi. This specimen is probably too early to be Megalosaurus, but should be checked for that genus' apomorphic keeled second centrum.
References- Owen, 1857. Monograph on the Fossil Reptilia of the Wealden Formations. Part III. Dinosauria (Megalosaurus). Palaeontographical Society Monographs. 34, 1-26.
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.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
M? sp. indet. (Owen, 1857)
Bajocian, Middle Jurassic
Inferior Oolite, England
Material- (NHMUK 47152) tooth (Lydekker, 1888)
(NHMUK R497) partial tooth (Owen, 1857)
Comments- Owen (1857) illustrates NHMUK R497 as Megalosaurus, while Lydekker (1888) refers it and 47152 to M. bucklandi. Huene (1932) thought these might belong to Magnosaurus instead of Megalosaurus. These specimens cannot be referred to Megalosaurus bucklandii because the teeth of that taxon lack described apomorphies.
References- Owen, 1857. Monograph on the fossil Reptilia of the Wealden and Purbeck Formations. Part III. Dinosauria (Megalosaurus). [Wealden]. The Palaeontographical Society, London. 1855, 1-26.
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.
M? sp. indet. (Lydekker, 1888)
Middle Bathonian, Middle Jurassic
Cotswold Slates, England
Material- (NHMUK 28608) tooth (Lydekker, 1888)
(NHMUK R2635) tooth (Benton and Spencer, 1995)
(GLCRM T.70-1, G.72-3, T.74-6) ribs, limb elements (Benton and Spencer, 1995)
Comments- These specimens cannot be referred to Megalosaurus bucklandii because the teeth of that taxon lack described apomorphies and the Cotsweld postcrania have yet to be described.
References- Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum of Natural History, London. 309 pp.
Benton and Spencer, 1995. Fossil Reptiles of Great Britain. Geological Conservation Review Series, Chapman & Hall. 386 pp.
M? sp. indet. (Naish, online 2022)
Middle Bathonian, Middle Jurassic
Enstone, Hampen Formation, England
Material- (OUMNH J29757) distal femur (~270 mm trans)
Comments- Naish (online 2022) wrote that Stewart had "noticed that what appeared to be the Plot bone ["Scrotum humanum"] was very much still in existence, and - furthermore - was on show in a museum; specifically, the Ashmolean Museum in Oxford ... where it’s accessioned as OUMNH PAL J29757." He noted several similarities, of which the longitudinal cracks in the flexor groove, elongate broken area covering the ectocondylar region and longitudinal striations proximal to that are valid and relatively unique, as well as both being found in Cornwell. As Naish notes, "It 'stands in' for the Plot bone" in the museum's display which identifies the latter as Megalosaurus, but is not explicitly said to be that specimen or Megalosaurus itself. However, Naish states that in 2021 Stewart and Ketchum weighed the specimen and determined it was about half the mass Plot reported for "Scrotum" so could not be the same specimen.
The Hampen Formation (used to be called Hampen Marly Formation or Beds) is only slightly above beds that diagnostic Megalosaurus is known from, so OUMNH J29757 could be that genus. However, Proceratosaurus, "Metriacanthosaurus" "reynoldsi" and Cruxicheiros are also known from around the same area and time, but the wider flexor groove is more like Megalosaurus than Cruxicheiros, and the shape is pretty dissimilar from proceratosaurid Guanlong and much larger than it or Proceratosaurus. "reynoldsi" is incomparable and of uncertain relations. The ectocondylar tuber does seem to be more medially placed than Megalosaurus (but similar to "Sctrotum"), but this may be due to compression and shearing.
Reference- Naish, 2022 online. Robert Plot’s Lost Dinosaur Bone. https://tetzoo.com/blog/2022/12/12/robert-plots-lost-dinosaur-bone
M? sp. indet. (Lydekker, 1888)
Late Bathonian, Middle Jurassic
Cornbrash Formation, England
Material- (NHMUK 47169) dorsal centrum
Comments- Lydekker (1888) provisionally referred this centrum to Megalosaurus bucklandi. This specimen cannot be referred to Megalosaurus bucklandii because the dorsal vertebrae of that taxon lack described apomorphies.
Reference- 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.
M? sp. indet. (Phillips, 1871)
Late Bathonian, Middle Jurassic
Forest Marble Formation, England
Material- (NHMUK 39476) tooth (Lydekker, 1888)
(OUM J29772) metatarsal IV (Phillips, 1871)
(Parker coll.) femur (Huene, 1926)
distal femur (~914 mm) (Phillip, 1871)
partial femur (~1.07 m) (Phillips, 1871)
teeth (Freeman, 1979)
Comments- Lydekker (1888) refers NHMUK 29476 to Megalosaurus bucklandi. Huene (1932) stated he could find no differences between Megalosaurus bucklandii and NHMUK 39476 and the Parker collection femur.
References- Phillips, 1871. Geology of Oxford and the Valley of the Thames: Oxford at the Clarendon Press. 523 pp.
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.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 1-167.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Delair, 1973. The dinosaurs of Wiltshire. Whiltshire Archaeology and Natural History Magazine. 68, 1-7.
Freeman, 1979. A Middle Jurassic mammal bed from Oxfordshire. Palaeontology. 22(1), 135-166.
M? sp. indet. (Owen, 1842)
Oxfordian, Late Jurassic
Corallian Oolite, England
Material- (Yorkshire Museum coll.) teeth
Reference- Owen, 1842. Report on British fossil reptiles, part II. Report of the British Association for the Advancement of Science. 11, 60-204.
M? sp. indet. (Lydekker, 1888)
Early Cretaceous
Potton Sands, England
Material- (NHMUK 42028) fragmentary posterior dorsal vertebra
Comments- Lydekker (1888) provisionally referred this vertebra to Megalosaurus bucklandi. This specimen is too late to be Megalosaurus, and cannot be referred to that taxon because the dorsal vertebrae of M. bucklandii lack described apomorphies.
Reference- 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.
M? sp. indet. (Owen, 1854)
Hauterivian-Barremian, Early Cretaceous
Weald Clay, England
Material- (Royal College of Surgeons coll.) posterior dorsal vertebra
(Royal College of Surgeons coll.) dorsal centrum, caudal vertebra, distal caudal vertebra
(Royal College of Surgeons coll.) partial neural arch
(Royal College of Surgeons coll.) ungual
Comments- These specimens are too late to be Megalosaurus, and cannot be referred to that taxon because the vertebrae and unguals of M. bucklandii lack described apomorphies.
Reference- Owen, 1854. Descriptive catalogue of the fossil organic remains of Reptilia and Pisces contained in the Museum of The Royal College of Surgeons of England. Taylor and Francis, London. 184 pp.
M? sp. indet. (Buckland, 1837)
Jurassic?
Besancon, France
Material- (University of Besancon coll.) (lost) jaw fragments, teeth, bones (Buckland, 1837)
(University of Besancon coll.) two sacral vertebrae (Buffetaut et al., 2007)
References- Buckland, 1837. Geology and Minerology Considered with Reference to Natural Theology. Philadelphia: Carey, Lea and Blanchard. 443 pp.
Buffetaut, Cuny and Pharisat, 2007. William Buckland's French Megalosaurus. Terra Nova. 6(3), 323-326.
M? sp. indet. (Sauvage, 1900)
Bathonian, Middle Jurassic
Indre, France
Material- tooth
Comments- This specimen cannot be referred to Megalosaurus bucklandii because the teeth of that taxon lack described apomorphies.
Reference- Sauvage, 1900. Note sur les poissons et les reptiles du Jurassique inferieur du department de l'Indre. Bulletin de la societie geologiques de France, 3rd series. 28, 500-504.
M? sp. indet. (Mercier, 1937)
Early Bathonian, Middle Jurassic
Ecouche Limestone, France
Material- tooth
Comments- This specimen cannot be referred to Megalosaurus bucklandii because the teeth of that taxon lack described apomorphies.
References- Mercier, 1937. Notes paleontologiques. Bulletin de la Société Linnéenne de Normandie, 8e série. 9, 33-36.
Buffetaut, Cuny and Le Loeuff, 1991. French Dinosaurs: The best record in Europe? Modern Geology. 16(1/2), 17-42.
M? sp. indet. (Lydekker, 1888)
Late Callovian-Early Oxfordian, Middle Jurassic-Late Jurassic
Vaches Noires Cliffs, Calvados, France
Material- (NHMUK 32725) proximal tibia (740 mm) (Lydekker, 1888)
(Tubingen Palaeontological University Museum coll.) distal femur, pedal phalanx III-1 (Huene, 1926)
Comments- Lydekker (1888) referred NHMUK 32725 to Megalosaurus bucklandi.
References- Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum of Natural History, London. 309 pp.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 1-167.
M? sp. indet. (Royo and Gomez, 1927)
Hauterivian-Barremian, Early Cretaceous
Weald Clay, Spain
Comments- These specimens are too late to be Megalosaurus.
Reference- Royo and Gomez, 1927. Sur le facies wealdien d'Espange. Bulletin de la societie geologiques de France, 4th series. 27, 125-128.
M? sp. indet. (Ubaghs, 1892)
Late Maastrichtian, Late Cretaceous
Maastricht Formation, Belgium
Material- fragmentary tooth
Comments- This specimen is too late to be Megalosaurus, and cannot be referred to that taxon because the teeth of M. bucklandii lack described apomorphies. Based on its age, this is more probably abelisaurid or dromaeosaurid.
Reference- Ubaghs, 1892. Le Megalosaurus dans la craie superieure du Limbourg. Bulletin de la Société belge de Géologie, de Paléontologie et d'Hydrologie, Bruxelles. 6, 26-29.

"Scrotum" Brookes, 1763
"S. humanum" Brookes, 1763
Aalenian-Bajocian?, Middle Jurassic
unknown quarry, Inferior Oolite?, Cornwell, England
Material- (lost) distal femur (~235 mm trans)
Diagnosis- (proposed) ectocondylar tuber limited to medial half of ectocondyle; ectocondyle subequal in size and shape to endocondyle.
Comments- Originally described and illustrated by Plot (1677) as the distal femur of a giant human, this is famous as being the first Mesozoic dinosaur bone to be published. Brookes (1793) later summarized Plot's description and opinions but labeled the specimen Sctrotum Humanum in his plate. Rieppel (2022) notes that "At the top of page 317, Brookes (1763) noted that 'other stones have been found exactly representing the private parts of a man; and others in the shape of kidneys . . . ', and continued further down on the same page" described the femur. As "the plates and the individual figures they contain are not numbered separately, but are identified by the pagination number of the page on which the respective specimens are mentioned or described. The conclusion seems to be that the illustrator took the femur fragment to be an example of those stones referred to on page 317 as 'exactly representing the private parts of a man', and erroneously labelled it accordingly." Phillips (1871) believed it was from the Inferior Oolite (Aalenian-Bajocian) and stated "It may have been the femur of a large megalosaurus or a small ceteosaurus" without evidence. As described by Delair and Sargeant (1975), Halstead (1970) "pointed out that because of its date of publication (post-Linnean, i.e. after 1758), this binomen can be considered a perfectly valid publication of the first generic and specific name ever applied to dinosaurian remains. It is perhaps fortunate that the name was not thereafter employed by any subsequent worker, and thus Scrotum humanum Brookes must be treated as a nomen oblitum and discarded." They (and Halstead) considered it more probable to be Megalosaurus than Cetiosaurus without evidence and stated "The specimen unfortunately is lost." Halstead and Sarjeant (1993; publication duplicated in 1995) noted that while Scrotum should be treated as a nomen oblitum under ICZN Article 23b (First Edition), "no application was made then, or has been made since, for the formal suppression of Brookes's binomen." The Third Edition of the ICZN came out in 1985 and eliminated the nomen oblitum clause, so the authors petitioned the ICZN in 1992 "(I) to use its plenary powers to suppress the generic name Scrotum Brookes, 1763 and the specific name S. humanum Brookes, 1763; (2) to retain on the Official List of Generic Names in Zoology the name Megalosaurus Buckland in Parkinson, 1822, type species by subsequent designation M. bucklandi Meyer. 1832. (3) to retain on the Official List of Specific Names in Biology the name bucklandi as published in the binomen Megalosaurus bucklandi (specific name of the type species of Megalosaurus Buckland in Parkinson, 1822, by designation in Meyer, 1832); (4) To place on the Official List of Rejected and Invalid Generic Names in Zoology the name Scrotum Brookes, 1763; (5) to place on the Official List of Rejected and Invalid Specific Names in Zoology the name humanum Brookes, 1763, as published in the binomen Scrotum humanum, and as suppressed in (1) above." as listed in 1993. As recalled by the authors, Tubbs (Executive Secretary to the ICZN) replied later that year that "The text on p. 301 of Brookes (1763) makes it quite clear that the two words "Scrotum humanum" on the plate were a description of a specimen, and that Brookes did not establish a genus Scrotum or a species humanum (any more than he did a species Kidney stone on the same plate!). The words just happened to be Latin." Furthermore, since "[the name Scrotum humanum] has never been used as a scientific name", it "is therefore unavailable under Article 11d of the Code" (Third Edition- "Names to be treated as valid when proposed. - Except as in (i) below, a name must be treated as valid for a taxon when proposed unless it was first published as a junior synonym and subsequently made available under the provisions of Section e of this article."). Finally, because "Plot's long-lost specimen was ... not certainly, a Megalosaurus bone", Tubbs wrote that "the Commission is willing to take action only when there is an appreciable and real, as opposed to hypothetical, threat to stability or nomenclature. This is not the case for Megalosaurus." Note that Tubbs was incorrect that Brookes ever specified Scrotum Humanum was a description instead of a name, with page 301 being an unrelated section on plant fossils, so his use of Article 11d was unwarranted although recently supported by Rieppel's logic. He was also wrong that it had never been used as a scientific name, as Molnar et al. (1990) listed Scrotum humanum as a carnosaur nomen dubium. Under the current ICZN, "Scrotum humanum" would be a nomen nudum based on Article 11.5- "To be available, a name must be used as valid for a taxon when proposed." Tubbs was right that the referral to Megalosaurus was merely hypothetical though, as it has never been supported by published evidence and seems unwarranted.
The femur is dissimilar from Cetiosaurus (both the lectotype OUMNH J13615 and the Rutland specimen LCM G468.1968) in that the sauropods are 45-86% larger, have a distally extended medial condyle, and a fibular groove placed at the lateral edge. Note the estimated transverse diameter is based on Plot's statement the narrowest shaft circumference was 15 inches (= 381 mm) and scaled from the figure. Compared to this, Megalosaurus femora are slightly smaller (shaft diameter 265-343 mm), and differ in having a more distomedially extended and pointed medial condyle, more laterally positioned ectocondylar tuber, and a straight lateral edge until the distal extent of the tuber. These same differences are also usually present in e.g. Cruxicheiros, piatnitzkysaurids, Eustreptospondylus, Erectopus, Allosaurus, and Juratyrant among large Jurassic theropods whose distal femora are undistorted and figured in posterior view. If "Scrotum" is from the Inferior Oolite it is also earlier than Megalosaurus, and differs from the contemporaneous Magnosaurus in the same ways when preserved (more laterally positioned ectocondylar tuber; straight lateral edge until the distal extent of the tuber), although it could derive from Duriavenator with which it cannot be compared. Sinraptor dongi is slightly more similar to "Scrotum" in having a convex lateral edge alongside the ectocondylar tuber, but it is "Brontoraptor" which is most similar in having that character, an evenly rounded medial condyle and a more medially placed ectocondylar tuber. Torvosaurus (ML 632) shows the last character at least but cannot be evaluated for the rest, so "Scrotum" might be best characterized as a torvosaur and may relate to "Megalosaurus" "phillipsi" from the Kimmeridgian of England that also has characters similar to "Brontoraptor" and Torvosaurus. "Brontoraptor" also has a similar circumference (376 mm) and internal cavity size based on Siegwarth et al.'s Figure 8E. Whether the remaining differences (ectocondylar tuber limited to medial half of ectocondyle; ectocondyle subequal in size and shape to endocondyle) are genuine or illustration error caused by Plot generalizing then unfamiliar megalosauroid anatomy is uncertain. Note "Scrotum" can be excluded from Ceratosauria based on the absence of a tall anteromedial crest, and from Coeluridae, Proceratosauridae and Maniraptoromorpha based on the deep extensor groove described by Plot and large size (with occasional exceptions, e.g. Yutyrannus).
References- Plot, 1677. The Natural History of Oxford-shire, being an essay towards the Natural History of England. Oxford. 358 pp.
Brookes, 1763. The Natural History of Waters, Earths, Stones, Fossils and Minerals, with their Virtues, Properties and Medicinal Uses: To which is added, the methods in which Linnaeus has treated these subjects. Vol. 5. J. Newberry. 364 pp.
Robinet, 1768. Vue philosophique de la gradation naturelle des formes de l'être, ou les essais de la nature qui apprend a faire l'homme. Harrevelt. 260 pp.
Phillips, 1871. Geology of Oxford and the Valley of the Thames. Oxford at the Clarendon Press. 523 pp.
Halstead, 1970. Scrotum humanum Brookes 1763 - the first named dinosaur. Journal of Insignificant Research. 5(7), 14-15.
Delair and Sargeant, 1975. The earliest discoveries of dinosaurs. Isis. 66, 5-25.
Buffetaut, 1979. A propos du reste de dinosaurien le plus anciennement décrit: l'interprétation de J.-B. Robinet (1768). Histoire et Nature. 14, 79-84.
Molnar, Kurzanov and Dong, 1990. Carnosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 169-209.
Halstead and Sarjeant, 1993. Scrotum humanum Brookes - the earliest name for a dinosaur? Modern Geology. 18, 221-224.
Halstead and Sarjeant, 1995. Scrotum humanum Brookes - the earliest name for a dinosaur? In Sarjeant (ed.), 1995. Vertebrate Fossils and the Evolution of Scientific Concepts; A tribute to L. Beverly Halstead. Gordon and Breach. 219-222.
Delair and Sargeant, 2002. The earliest discoveries of dinosaurs: The records re-examined. Proceedings of the Geologists' Association. 113, 185-197.
Rieppel, 2022 (online 2021). The first ever described dinosaur bone fragment in Robinet's philosophy of nature (1768). Historical Biology. 34(5), 940-946.

"Megalosaurus" "phillipsi" Welles, Powell and Pickering vide Pickering, 1995
Kimmeridgian, Late Jurassic
Kimmeridgian Clay, England
Material- (OUM J29886) tibia (740 mm) (Huene, 1926)
....(OUM J13586) metatarsal II (360 mm), incomplete metatarsal III, incomplete metatarsal IV (Phillips, 1871)
Diagnosis- (from Pickering, DML 2002) cnemial crest more laterally angled.
(suggested) metatarsal II transversely narrower in proximal view; a wide lateral flare to the ventral rim of the lateral ligament pit on metatarsal II.
Other diagnoses- Pickering has listed his intended diagnosis for this species online (DML, 2002). Of his listed characters, the tibia does not seem noticably straighter in "phillipsi" than in Megalosaurus or Torvosaurus. The fibular crest begins at a similar position proximally. I cannot confirm the presence of a pit in the astragalar facet in M. bucklandii, and the "concavity of the distal end is wider" in Torvosaurus and "Brontoraptor" as well. The latter is also true of the medial malleolus, which "does not extend so far below the medial end." The tibia is equally robust in "Brontoraptor", Torvosaurus tanneri and T? sp. ML 430 from Portugal, which also share the transversely expanded lateral condyle. While the fibular crest extends more distally than in M. bucklandii, it does not do so more than "Brontoraptor" or ML 430.
Comments- OUM J29886 is intended as the holotype, though Huene (1926) felt they belonged to the same individual.
Phillips (1871) illustrated the metatarsus as Megalosaurus in figure 68. Huene (1926) later illustrated both the tibia and metatarsus as Megalosaurus bucklandi, incorrectly stating they derive from the Stonesfield Slate (Pickering, DML 2002). Huene also lists the metatarsals again as Megalosaurus sp. under entry 54 of his list, which also get listed as ?Megalosaurus sp. by him in 1932. Pickering (1995) first mentioned the name Megalosaurus phillipsi in an unpublished bibliographic manuscript. It was later used in the comparative section of another unpublished manuscript (Welles and Pickering, 1999). This paper was largely extracted from the European megalosaur manuscript Welles and Powell worked on in the 1970s but never published, specifically the Megalosaurus redescription section. Pickering intends to publish an updated version of the megalosaur manuscript as Mutanda Dinosaurologica, and has posted small excerpts including the diagnosis of "phillipsi" online (DML, 2002). In any case, the name is a nomen nudum as Pickering didn't follow ICZN Article 8.1.3- it must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies. Curiously, Benson's (2010) recent redescription of Megalosaurus does not even mention the specimens, though they do lack the only character in his diagnosis of M. bucklandii for which it can be checked- complementary groove and ridge structures on the articular surfaces between metatarsals II and III.
When entered into my saurischian supermatrix, "phillipsi" emerges in a polytomy with non-spinosaurid megalosauroids. This specimen will be compared to Megalosaurus bucklandii, Torvosaurus and "Brontoraptor" (note Pickering and Welles synonymized the first two at least), as it differs markedly from Eustreptospondylus (used as the outgroup comparison), while Magnosaurus and Dubreuillosaurus are too incomplete to be compared usefully and Afrovenator remains largely undescribed. Poekilopleuron differs in having a proximal corner on its medial malleolus and an astragalar facet that extends far up the medial edge of the tibia. As noted above in the 'other diagnoses' section, "phillipsi" apomorphically resembles Morrison megalosaurs and ML 430 more than Megalosaurus in being robust and having a transversely expanded lateral condyle. It apomorphically resembles "Brontoraptor" and ML 430 more in having a distally extending fibular crest, but is primitively more like Megalosaurus than any of these taxa in having a low angled medial malleolus. The proximodistally wide medial side to the astragalar facet is most similar to "Brontoraptor" and Torvosaurus, and the fibular crest lacks the apomorphically bulbous shape of Megalosaurus'. In proximal view, the incisura tibialis is plesiomorphically shallower than Megalosaurus as in Torvosaurus and "Brontoraptor", but the shape is plesiomorphically more similar to Megalosaurus than "Brontoraptor" in having a narrower cnemial crest and more posteriorly extensive medial condyle. Metatarsal II is of equal robusticity to Megalosaurus, unlike Torvosaurus' apomorphically massive element. It apomorphically shares the medial flare to the ventral rim of the collateral ligament pit with Torvosaurus though. It differs from both in being transversely narrower proximally and having a wide lateral flare to the ventral rim of the opposite collateral ligament pit. In proximal view, Megalosaurus and "phillipsi" both have a posterior concavity on metatarsal II, and share a convex anterior edge, as in the outgroup. Distally, it is primitively deeper than in either Megalosaurus or Torvosaurus. Metatarsal III is primitively more medially bowed than either, and is again less robust than Torvosaurus'. Proximally, metatarsal III is closer to Torvosaurus in having an apomorphic highly concave lateral margin and lacking the apomorphic sinuous medial margin of Megalosaurus. It has a primitively blunter posteromedial corner than either of them. Distally, the lateral condyle is apomorphically narrower as in Torvosaurus. Metatarsal IV has an apomorphically concave lateral edge as in Megalosaurus, unlike the sinusoidal edge of Torvosaurus. The distal shaft width is intermediate, and the articular end plesiomorphically expands transversely less than in Megalosaurus. Ignoring the symplesiomorphies then, "phillipsi" shares five apomorphic states with Torvosaurus, and only one with Megalosaurus. It also shares these apomorphies with "Brontoraptor" and ML 430 where known, and shares the distally projecting fibular crest uniquely with those two specimens. I conclude Welles, Powell and Pickering were correct to refer "phillipsi" to their more inclusive version of Megalosaurus, but if Torvosaurus is separated as most modern workers do, it would be better referred to that genus. "Brontoraptor" has been recently assumed to be synonymous with Torvosaurus, and if so the features "phillipsi" shares with the former to the exclusion of the latter may be individual variation or incorrectly illustrated for "Brontoraptor". However, the fact Portuguese tibia ML 430 has the same characters suggests this may not be the case. Whether ML 430 should be referred to "phillipsi" is an interesting question, as they are both from Late Jurassic Europe. While the distally extensive fibular crest might suggest so, the broadly flared medial malleolus and laterally straight lateral malleolus of ML 430 resemble Torvosaurus more, while the medially narrow astragalar facet is more like Megalosaurus than any torvosaur. Both "phillipsi" and ML 430 are best referred to Megalosauridae.
References- Phillips, 1871. Geology of Oxford and the Valley of the Thames. Oxford at the Clarendon Press. 523 pp.
Huene, 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 35-167.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Pickering, 1995. Jurassic Park: Unauthorized Jewish Fractals in Philopatry. A Fractal Scaling in Dinosaurology Project, 2nd revised printing. Capitola, California. 478 pp.
Welles and Pickering, 1999. Megalosaurus bucklandii. Private publication of Stephen Pickering. An extract from Archosauromorpha: Cladistics & Osteologies. A Fractal Scaling in Dinosaurology Project. 119 pp.
Pickering, DML 2002. https://web.archive.org/web/20191030094450/http://dml.cmnh.org/2002Mar/msg00553.html
Pickering, in prep. Mutanda Dinosaurologica.

"Brontoraptor" Redman, 1995
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of Morrison Formation, Wyoming, US
Material- (TATE 0012; = CPS 1003; = TATE 1003) (adult) atlas (60 mm), axis (150 mm), sacrum (620 mm), first caudal vertebra (105 mm), second caudal vertebra (113 mm), third caudal vertebra (121 mm), fourth caudal vertebra (112 mm), fifth caudal vertebra (113 mm), seventh caudal vertebra (111 mm), eighth caudal vertebra (118 mm), ninth caudal vertebra (118 mm), tenth caudal vertebra (119 mm), eleventh caudal vertebra (118 mm), twentieth caudal vertebra (118 mm), six chevrons (305, 285, 265, 245, 210, 118 mm), scapula, coracoids, ilium (~930 mm), pubis (~610 mm), ischium (590 mm), femur (830 mm), tibia (700 mm), fibula (650 mm)
Diagnosis- (from Siegwarth et al., unpublished) differs from Torvosaurus tanneri in- ilium with straight dorsal edge; vertical pubic peduncle of ilium; brevis shelf oriented laterally; short pubic symphysis; reduced pelvic canal; ischium not strongly curved; obturator process present; no ischial symphysis.
differs from Edmarka rex in- scapula expanded distally; pubis and ischium more gracile with different distal expansions
Comments- Siegwarth et al. (unpublished, though written after 1994 based on the bibliography, and listed as 1997 by Carrano et al., 2012) wrote a manuscript that is oddly credited to volume 2 of the nonexistant journal 'Contributions from the Tate Museum Collections, Casper, Wyoming'. In it, they describe the sacrum, first caudal vertebra, ilium, pubis, femur, tibia and fibula of TATE 0012 as parts of Edmarka paratype TATE 1003. A later version of the manuscript included Bakker and described more material from the specimen, this time calling it TATE 0012, which Edmarka paratype CPS 1003 had apparently been integrated into as TATE 0012-11. All of this material was discovered in 1993-1996. Redman (1995) had used the name Brontoraptor for this material in a magazine article (without species name, making it a nomen nudum- ICZN 13.3). In the second unpublished manuscript, Seigwarth et al. state TATE 0012 is at least a different species than Torvosaurus tanneri and probably a different genus. They note earlier versions of the manuscript called the specimen Brontoraptor but that based on a recommendation by Carpenter, they will wait for further comparison to Torvosaurus to name a new genus. The final version of this manuscript was cited by Bakker (1996) as being published in volume 3 of Hunteria, but there's no evidence such a volume was produced. Mateus et al. (2006) and Carrano et al. have listed the material as Torvosaurus without justification. The latter merely state "Brontoraptor" and Edmarka "may represent species level variants of Torvosaurus, but we do not consider the observed differences as sufficient to justify a new taxon at this time."
References- Redman, 1995. [title unknown]. Paleo Horizons. Winter Issue, [pp unknown].
Bakker, 1996. The real Jurassic Park: Dinosaurs and habitats at Como Bluff, Wyoming. In Morales (ed.). Museum of Northern Arizona Bulletin. 60, 35-49.
Mateus, Walen and Antunes, 2006. The large theropod fauna of the Lourinha Formation (Portugal) and its similarity to the Morrison Formation, with a description of a new species of Allosaurus. In Foster and Lucas (eds.). Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin. 36, 123-129.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Siegwarth, Lindbeck, Redman, Southwell, unpublished. Giant carnivorous dinosaurs of the family Megalosauridae from the Late Jurassic Morrison Formation of eastern Wyoming. Contributions from the Tate Museum Collections, Casper, Wyoming. 2, 40 pp.
Siegwarth, Lindbeck, Redman, Southwell and Bakker, unpublished. Megalosaurid dinosaurs from the Late Jurassic Morrison Formation of Eastern Wyoming. 27 pp.
Siegwarth, Lindbeck, Bakker and Southwell, "1996" [unpublished]. Giant carnivorous dinosaurs of the family Megalosauridae. Hunteria. 3, 1-77.

Edmarka Bakker, Kralis, Siegwarth and Filla, 1992
E. rex Bakker, Kralis, Siegwarth and Filla, 1992
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of Morrison Formation, Wyoming, US
Syntypes- (CPS 1002) incomplete scapula (~950 mm), partial coracoid
(CPS 1004) dorsal ribs
(CPS 1005) jugal (444 mm)
(CPS 1006) first caudal vertebra (140 mm)
Paratype- (CPS 1001) partial third dorsal rib, fourth dorsal rib, ninth dorsal rib, tenth dorsal rib, partial eleventh dorsal rib, dorsal rib
Referred- ?(CPS 1010) pubis (866 mm) (Bakker et al., 1992)
?(TATE coll.) pubis (790 mm), proximal ischium (Siegwarth et al., unpublished)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Utah, US
?(CEUM 3301) pubes (Siegwarth et al., unpublished)
Comments- Bakker et al. (1992) confusingly label the proximal caudal vertebra as CPS 1005 in the material list, which is the same number as the jugal. In the discussion, it is called CPS 1006. Also, they label the ribs CPS 1001 in the discussion and figures, but as CPS 401 in the material list. Given the other specimen numbers, CPS 1001 is the more likely possibility. A coracoid (CPS 1003, though now apparently part of TATE 0012) was originally included as a paratype, but is part of a specimen Siegwarth et al. (unpublished) believe differs from Edmarka and which has been called "Brontoraptor".
Synonymized with Torvosaurus tanneri by Holtz et al. (2004) and Carrano et al. (2012) without detailed analysis.
References- Bakker, Kralis, Siegwarth and Filla, 1992. Edmarka rex, a new, gigantic theropod dinosaur from the Middle Morrison Formation, Late Jurassic of the Como Bluff outcrop, with comments on the evolution of the chest region and shoulder in theropods and birds and a discussion of the five cycles of originn and extinction among giant dinosaurian predators. Hunteria. 2(9), 1-24.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Siegwarth, Lindbeck, Redman, Southwell and Bakker, unpublished. Megalosaurid Dinosaurs from the Late Jurassic Morrison Formation of Eastern Wyoming. 27 pp.

Wiehenvenator Rauhut, Hübner and Lanser, 2016
W. albati Rauhut, Hübner and Lanser, 2016
Middle Callovian, Middle Jurassic
Ornatenton Formation, Germany
Holotype- (WMN P27454) (Das Monster von Minden) (>9 year old subadult) (~8.3 m) lateral tooth
....(WMN P27455) fused medial gastralia
....(WMN P27456) anterior tooth (~42 mm)
....(WMN P27457) lacrimal
....(WMN P27458) astragalus (193 mm trans)
....(WMN P27459) lateral tooth
....(WMN P27461-62) two dentary fragments
....(WMN P27464) dorsal rib fragment
....(WMN P27465) dorsal rib fragment
....(WMN P27466) dentary fragment
....(WMN P27467) lateral tooth
....(WMN P27470) anterior quadratojugal
....(WMN P27473) lateral tooth
....(WMN P27475) incomplete premaxilla
....(WMN P27476) dorsal rib fragment
....(WMN P27477) postorbital
....(WMN P27479) fibula (~730 mm)
....(WMN P27480) anterior dorsal rib (1.05 m)
....(WMN P27482) incomplete manual phalanx III-1 (~81 mm)
....(WMN P27483) lateral tooth (~73 mm)
....(WMN P27484) partial calcaneum
....(WMN P27499) mid caudal vertebra (114 mm)
....(WMN P27500) mid caudal vertebra (114 mm)
....(WMN P27501) incomplete proximal caudal vertebra (~110 mm)
....(WMN P27502) fibula
....(WMN P27403) posterior dorsal rib
....(WMN P27504) incomplete maxilla (skull ~870 mm)
?...(WMN P29131) partial caudal vertebra
?...(WMN P47417) partial caudal vertebra
Diagnosis- (after Rauhut et al., 2016) Anterior margin of nasal process of premaxilla offset from anterior margin of premaxillary body by slight concavity (also in Sciurumimus); medial premaxillary foramen placed over second alveolus, not the third; first premaxillary tooth considerably smaller than second (also in spinosaurids); maxillary antorbital fossa reduced; excavatio pneumatica on ascending process of maxilla (also in Ceratosaurus and some allosauroids); lacrimal with small oval depression in antorbital fossa anterior to lacrimal fenestra; very short and high anterior process of lacrimal, its length less than half the height of the bone; postorbital with transversely notably concave orbital facet in dorsal part of anterior side, with markedly raised lateral and medial margins; oblique ridge on medial side of fibula at about the level of iliofibularis tubercle; proximal part of astragalar ascending process deflected laterally, resulting in marked kink in lateral margin of process.
Comments- The holotype was discovered in 1998 and initially reported in 1999 as a giant (~15 m) allosaur (e.g. November National Geographic article), called Das Monster von Minden after the town it was discovered in. Nothing further was released until Rauhut et al.'s (2015) abstract, placing the taxon in Megalosauridae. It was officially described by Rauhut et al. (2016), who used a version of Carrano et al.'s tetanurine analysis to recover Wiehenvenator as a megalosaurid sister to Torvosaurus. Note the actual size turns out to be much smaller than the initial claims, with the authors' skeletal reconstruction being 8.3 meters long. Also, the manual phalanx was identified as a pedal phalanx in the 2015 abstract. Only two of the ribs are complete and only the left fibula is undistorted enough to measure, but the paper does not specify which specimen numbers correspond to these (Rauhut, pers. comm. 2016 provided the identifications for the materials list above)..
References- [author?], 1999. [title?] National Geographic German Edition. November, [pp].
Rauhut, Hübner and Lanser, 2015. A new theropod dinosaur from the late Middle Jurassic of Germany and theropod faunal turnover during the Jurassic. Libro de resúmenes del V Congreso Latinoamericano de Paleontología de Vertebrados. 62.
Rauhut, Hübner and Lanser, 2016. A new megalosaurid theropod dinosaur from the late Middle Jurassic (Callovian) of north-western Germany: Implications for theropod evolution and faunal turnover in the Jurassic. Palaeontologia Electronica. 19.2.26A, 1-65.

Torvosaurus Galton and Jensen, 1979
Diagnosis- (after Britt, 1991) interdental wall making up one-half the medial surface of the maxillary body.
(after Carrano et al., 2012) very shallow maxillary fossa; fused interdental plates.
(after Hendrickx and Mateus, 2014b) protuberant ridge below maxillary fossa, in ventral part of anterior corner of lateral antorbital fossa.
Other diagnoses- Hendrickx and Mateus (2014b) also listed expanded fossae in posterior dorsal and anterior caudal centra forming enlarged and deep pneumatic openings; highly ossified puboischiadic plate; and distal expansion of ischium with prominent lateral midline crest and oval outline in lateral view. However, these areas are unpreserved in T. gurneyi and thus only scorable in T. tanneri.
References- Galton and Jensen, 1979. A new large theropod dinosaur from the Upper Jurassic of Colorado. Brigham Young University Geology Studies. 26(2), 1-12.
Britt, 1991. Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri. Brigham Young University Geology Studies. 37, 1-72.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx and Mateus, 2014b. Torvosaurus gurneyi n. sp., the largest terrestrial predator from Europe, and a proposed terminology of the maxilla anatomy in nonavian theropods. PLoS ONE. 9(3), e88905.
T. tanneri Galton and Jensen, 1979
= Megalosaurus tanneri (Galton and Jensen, 1979) Paul, 1988
Tithonian, Late Jurassic
Brushy Basin Member of Morrison Formation, Colorado, Utah, Wyoming, US
Lectotype- (BYUVP 2002 in part) humerus (429)
Paralectotypes- (BYUVP 2002 in part) humerus (412 mm), radius (187 mm), ulnae (225, 229 mm)
(BYUVP 2003) (skull ~1.31 m) anterior dentary
(BYUVP 2004a-d) fragment of fifth cervical vertebra, sixth cervical vertebra (115 mm), seventh cervical vertebra (123 mm), eighth cervical vertebra (121 mm)
(BYUVP 2005) first dorsal vertebra (135 mm)
(BYUVP 2006) fifth dorsal vertebra (118 mm)
(BYUVP 2007) sixth dorsal vertebra (~107 mm)
(BYUVP 2008) twelfth dorsal vertebra
(BYUVP 2009) dorsal vertebra
(BYUVP 2010) metacarpal I (72.3 mm)
(BYUVP 2011) metacarpal II (117.5 mm)
(BYUVP 2012) metacarpal III (96.5 mm)
(BYUVP 2013) ilium
(BYUVP 2014) pubes (736 mm)
(BYUVP 2015) ischium (736 mm)
(BYUVP 2016) tibia (725 mm)
(BYUVP 2017) incomplete tibia
(BYUVP 2018) manual phalanx I-1 (~65 mm)
(BYUVP 5029) astragalus
(BYUVP 9620) fibula (688 mm)
(BYUVP 9622) calcaneum
Referred- (BYU 725/2003) tooth (Brusatte et al., 2007)
(BYUVP 2020) manual ungual I (Galton and Jensen, 1979)
(BYUVP 4860) third cervical vertebra (90 mm) (Britt, 1991)
(BYUVP 4881) distal ischium (Britt, 1991)
(BYUVP 4882) premaxilla (Jensen, 1985)
(BYUVP 4883) jugals (Jensen, 1985)
(BYUVP 4884) atlantal intercentrum (50 mm), neurapophysis (Jensen, 1985)
(BYUVP 4890) thirteenth dorsal vertebra (130 mm) (Britt, 1991)
(BYUVP 4976a-k) eighth caudal vertebra (115 mm), eighth chevron, ninth caudal vertebra (118 mm), ninth chevron, tenth caudal vertebra (119 mm), tenth chevron (232 mm), eleventh caudal vertebra (116 mm), eleventh chevron (220 mm), twelfth caudal vertebra (104 mm), twelfth chevron (190 mm), thirteenth caudal vertebra (109 mm), thirteenth chevron (185 mm), fourteenth caudal vertebra (100 mm), fourteenth chevron (~180 mm), fifteenth caudal vertebra, fifteenth chevron, sixteenth caudal vertebra (100 mm) (Jensen, 1985)
(BYUVP 4977) ilium (853 mm) (Britt, 1991)
(BYUVP 4998) fourth dorsal vertebra (124 mm) (Britt, 1991)
(BYUVP 5004) fourth caudal vertebra (127 mm) (Britt, 1991)
(BYUVP 5005) metatarsal III (320 mm) (Jensen, 1985)
(BYUVP 5009) metatarsal III (348 mm) (Jensen, 1985)
(BYUVP 5020) calcaneum (Britt, 1991)
(BYUVP 5077) metatarsal II (295 mm) (Jensen, 1985)
(BYUVP 5086) second caudal vertebra (116 mm) (Britt, 1991)
(BYUVP 5110) quadrate (Britt, 1991)
(BYUVP 5129) fibula (686 mm), astragalus (Britt, 1991)
(BYUVP 5136) fibula (Britt, 1991)
(BYUVP 5147) metatarsal II (303 mm) (Jensen, 1985)
(BYUVP 5241) metatarsal III (357 mm) (Jensen, 1985)
(BYUVP 5276) metatarsal II (288 mm) (Jensen, 1985)
(BYUVP 5277) metatarsal III (358 mm) (Jensen, 1985)
(BYUVP 5278) metatarsal IV (302 mm) (Jensen, 1985)
(BYUVP 5279) metatarsal IV (308 mm) (Jensen, 1985)
(BYUVP 5280) metatarsal III (365 mm) (Jensen, 1985)
(BYUVP 5281) metatarsal II (307 mm) (Jensen, 1985)
(BYUVP 5286) lacrimal (Jensen, 1985)
(BYUVP 8966) metatarsal IV (302 mm) (Jensen, 1985)
(BYUVP 9013) twelfth caudal vertebra (Britt, 1991)
(BYUVP 9090) tenth dorsal vertebra (~113 mm) (Britt, 1991)
(BYUVP 9120) seventh dorsal vertebra (112 mm) (Britt, 1991)
(BYUVP 9121) ninth dorsal vertebra (Britt, 1991)
(BYUVP 9122) maxilla (Jensen, 1985)
(BYUVP 9135) twenty-third caudal vertebra (86 mm) (Britt, 1991)
(BYUVP 9249) postorbital (Britt, 1991)
(BYUVP 9621) astragalus (Jensen, 1985)
(BYUVP coll.) teeth (Jensen, 1985)
(CM 1254) tooth (Britt, 1991)
(DMNH 2243) posterior dorsal vertebra (Britt, 1991)
(FMNH PR 3060) three gastralial fragments, metacarpal III (85 mm), phalanx III-2 (34 mm), pedal phalanx I-1 (80 mm), incomplete metatarsal II (~306 mm), incomplete metatarsal III (~356 mm), metatarsal IV (287 mm) (Hanson and Makovicky, 2014)
(MWC coll.) tooth (Kane, 2020)
?(YPM 58269) partial tibia (Dalman, 2014)
(YPM coll.) tooth (>100 mm) (Lull, 1927)
two teeth (Foster, 2005)
Kimmeridgian, Late Jurassic
Salt Wash Member of the Morrison Formation, Colorado, US
humerus (Carpenter, 1998)
Kimmeridgian-Tithonian, Late Jurassic
Morrison Formation, Colorado, US (Skull Creek quarry)
(CMC VP15575; Elvis) (~10 m) premaxillary teeth, maxilla, surangular, angular, articular, teeth, fifth to tenth cervical vertebrae, cervical ribs, first to thirteenth dorsal vertebrae, dorsal ribs, sacrum, first to second caudal vertebrae, partial furcula, ilia, pubes, ischia, femur, distal tibia, phalanx II-2, proximal phalanx III-2 (Galiano, 2015)
(DMNS coll.) tooth (Kane, 2020)
Diagnosis- (after Rauhut, 2000) opisthocoelous cervical vertebrae with a pronounced flat rim around the anterior ball (unknown in T. gurneyi); fenestra in neural arch of dorsal vertebrae in front of hyposphene (unknown in T. gurneyi).
(after Carrano et al., 2012) expanded pneumatic fossae in posterior dorsal and anterior caudal vertebrae centra forming enlarged, deep openings (unknown in T. gurneyi); highly ossified puboischiadic plate (unknown in T. gurneyi); distal expansion of ischium with prominent lateral midline crest and oval outline in lateral view (unknown in T. gurneyi).
(after Hendrickx and Mateus, 2014b) protuberant ridge on anterior part of medial shelf, posterior to anteromedial process; interdental wall falling short relative to lateral wall; interdental plates with broad V-shaped
ventral margin.
Other diagnoses- The shallow maxillary fossa and fused interdental plates listed by Carrano et al. (2012) are also present in T. gurneyi.
Comments- Britt (1991) noted the forelimb elements of BYUVP 2002 were found disarticulated and that at least two equally sized individuals were preserved in the quarry, so that restriction of the holotype would be preferrable. He chose the left humerus, but the ICZN doesn't allow a holotype to be designated after the fact. The humerus would be a lectotype, which is how it is interpreted by Hendrickx and Mateus (2014b). Also note the original BYU 2016 of Galton and Jensen (1979) was divided into several specimen numbers by Britt.
Allain et al. (2012) believed manual ungual I BYUVP 2020 could be from a spinosaurid because of its elongation, but as no other spinosaurid remains are known from the Morrison, this is questionable.
The most significant articulated specimen is nicknamed Elvis and was initially privately owned (Galiano, 2015) before being bought by the Cincinnati Museum Center as CMC VP15575 (Miller, pers. comm. 2021).
References- Lull, 1927. Organic Evolution, Macmillan, New York. 729 pp.
Galton and Jensen, 1979. A new large theropod dinosaur from the Upper Jurassic of Colorado. Brigham Young University Geology Studies. 26(2), 1-12.
Jensen, 1985. Uncompahgre dinosaur fauna: A preliminary report. Great Basin Naturalist. 45, 710-720.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 464 pp.
Britt, 1991. Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri. Brigham Young University Geology Studies. 37, 1-72.
Carpenter, 1998. Vertebrate biostratigraphy of the Morrison Formation near Cañon City, Colorado. Modern Geology. 23, 407-426.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Foster, 2005. Evidence of size-classes and scavenging in the theropod Allosaurus fragilis at the Mygatt-Moore Quarry (Late Jurassic), Rabbit Valley, Colorado: Journal of Vertebrate Paleontology. 25(3), 59A.
Brusatte, Benson, Carr, Williamson and Sereno, 2007. The systematic utility of theropod enamel wrinkles. Journal of Vertebrate Paleontology. 27(4), 1052-1056.
Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften. 99(5), 369-377.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Dalman, 2014. New data on small theropod dinosaurs from the Upper Jurassic Morrison Formation of Como Bluff, Wyoming, USA. Volumina Jurassica. 12(2), 181-196.
Hanson and Makovicky, 2014. A new specimen of Torvosaurus tanneri originally collected by Elmer Riggs. Historical Biology. 26(6), 775-784.
Hendrickx and Mateus, 2014b. Torvosaurus gurneyi n. sp., the largest terrestrial predator from Europe, and a proposed terminology of the maxilla anatomy in nonavian theropods. PLoS ONE. 9(3), e88905.
Galiano, 2015. Torvosaurus. Maxilla & Mandible Ltd. 11 pp.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.
Kane, 2020. Identifying Jurassic theropod genera using GIS maps of tooth serrations. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 197.
T. gurneyi Hendrickx and Mateus, 2014b
Late Kimmeridgian, Late Jurassic
Porto Novo-Amoreira Member of Lourinha Formation, Portugal
Holotype- (ML 1100) (~10 m, ~4-5 tons) (skull ~1.15 m) maxilla (612 mm), tooth (127 mm), partial proximal caudal centrum
Paratype- (ALT-SHN.116) maxillary fragment (Malafaia et al., 2008)
Late Kimmeridgian, Late Jurassic
Sobral Member of Lourinha Formation, Portugal
Referred- (FUB PB Ther 1) lateral tooth (Rauhut and Kriwet, 1994)
?(FUB PB Ther 2) partial tooth (Rauhut and Kriwet, 1994)
(ML 1188) (embryo) maxilla, dentary, dentary tooth, five teeth, three centra, bones, eggshell fragments (Araujo et al., 2012)
Early Tithonian, Late Jurassic
Bombaral Member of Lourinha Formation, Portugal
(ML 430) (~7 m, ~1.6-1.7 tons) tibia (820 mm) (Mateus and Antunes, 2000)
(ML 962) anterior tooth (85.8 x 31.2 x 20.2 mm) (Hendrickx and Mateus, 2014a)
Late Kimmeridgian-Early Tithonian, Late Jurassic
Lourinha Formation, Portugal
(ML 148) tooth (? x 35.2 x 17.7 mm) (Hendrickx, Mateus and Araújo, in press)
(ML 500) tooth (? x 41.3 x 19.9 mm) (Hendrickx, Mateus and Araújo, in press)
(ML 632) (~10 m; ~3-4 tons) distal femur (~1.11 m) (Mateus et al., 2006)
(ML 857) tooth (59.5 x 32.2 x 17.1 mm) (Hendrickx, Mateus and Araújo, in press)
(ML 1853) tooth (43.5 x 24.8 x 13.7 mm) (Hendrickx, Mateus and Araújo, in press)
Diagnosis- (after Hendrickx and Mateus, 2014b) than eleven maxillary teeth; interdental plates with straight ventral margin; interdental wall nearly coincidental with lateral wall of maxillary body; absence of protuberant ridge on anterior part of medial shelf, posterior to anteromedial process, with coincidental posterior extension of dorsal and medial ridges of anteromedial process.
Comments- The holotype was discovered in 2003 and first described by Mateus et al. (2006), then described in depth later by Hendrickx and Mateus (2014b). Rauhut and Kriwet (1994) first described FUB PB Ther 1 and 2 as "Megalosaurus" pombali specimens which they placed as Carnosauria indet., but the first was later referred to Torvosaurus gurneyi by Hendrickx and Mateus (2014b). The eggshells associated with Torvosaurus are dendroolithid.
References- Rauhut and Kriwet, 1994. Teeth of a big theropod dinosaur from Porto das Barcas (Portugal). Berliner Geowissenschaftliche Abhandlungen, E. 13, 179-185.
Mateus and Antunes, 2000. Torvosaurus sp. (Dinosauria: Theropoda) in the Late Jurassic of Portugal. Livro de Resumos do I Congresso Iberico de Paleontologia. 115-117.
Antunes and Mateus, 2003. Dinosaurs of Portugal. Comptes Rendus Palevol. 2, 77-95.
Mateus, Walen and Antunes, 2006. The large theropod fauna of the Lourinha Formation (Portugal) and its similarity to the Morrison Formation, with a description of a new species of Allosaurus. In Foster and Lucas (eds.). Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin. 36, 123-129.
Malafaia, Ortega, Silva and Escaso, 2008. Fragmento de un maxilar de teropodo de Praia da Corva (Jurasico Superior. Torres Vedras, Portugal). Palaeontologica Nova SEPAZ. 8, 273-279.
Araujo, Castanhinha, Mateus and Martins, 2012. Late Jurassic theropod embryos from Porto das Barcas, Lourinha Formation, Portugal. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 57.
Araujo, Castanhinha, Martins, Mateus, Hendrickx, Beckmann, Schell and Alves, 2013. Filling the gaps of dinosaur eggshell phylogeny: Late Jurassic theropod clutch with embryos from Portugal. Scientific Reports. 3, 1924.
Hendrickx and Mateus, 2014a. Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth. Zootaxa. 3759(1), 1-74.
Hendrickx and Mateus, 2014b. Torvosaurus gurneyi n. sp., the largest terrestrial predator from Europe, and a proposed terminology of the maxilla anatomy in nonavian theropods. PLoS ONE. 9(3), e88905.
Malafaia, Ortega and Silva, 2014. New cranial remains assigned to Megalosauridae (Dinosauria: Theropoda) from the Late Jurassic of Lusitanian Basin (Portugal). Journal of Vertebrate Paleontology. Program and Abstracts 2014, 175.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.
T? sp. (Gerke and Wings, 2014)
Early Tithonian, Late Jurassic
Thueste, Germany
Material- (NLMH coll.) tooth tip
Reference- Gerke and Wings, 2014. Characters versus morphometrics: A case study with isolated theropod teeth from the Late Jurassic of Lower Saxony, Germany, reveals an astonishing diversity of theropod taxa. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 137.
T? sp. (Lü and Hu, 1998)
Late Jurassic
Datong, Shanxi, China
Material- eighth dorsal centrum (~140 mm), partial eleventh dorsal vertebra, partial thirteenth dorsal centrum (~134 mm), fused partial second, third and partial fourth sacral centra
Comments- Lü and Hu (1998) referred these to Allosaurus sp., but Carrano et al. (2012) felt the "overall morphology and cavernous lateral depressions" were comparable to Torvosaurus and considered them Tetanurae indet. cf. Torvosaurus.
References- Lü and Hu, 1998. Dinosaur remains from Datong Suburb, Shanxi Province. Vertebrata PalAsiatica. 36(3), 252-256.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

Afrovenatorinae Carrano, Benson and Sampson, 2012
Definition- (Afrovenator abakensis <- Megalosaurus bucklandii)
= Megalosaurinae sensu Allain, 2002
Definition- (Poekilopleuron bucklandii <- Torvosaurus tanneri) (modified)
Diagnosis- (after Carrano et al., 2012) squared anterior margin of maxillary antorbital fenestra (also in Irritator); puboischiadic plate broadly open along midline.
References-Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

unnamed possible afrovenatorine (Liu and Yeh, 1957)
Aalenian?, Middle(?) Jurassic
Shuangyukou, Qianfoya Formation?, Sichuan, China
Material- (IVPP V893) (~1.4 m) (juvenile?) incomplete manual ungual I (~24 mm)
Comments- This was discovered in 1955 associated with the holotype of Ceratodus minor (IVPP V892) (Liu and Yeh, 1957). The locality is listed as "Shung-ya-kou, Hai-t'ang-pu, Kiangyou, Szechuan", which is now called Shuangyukou, Haitangpu, Jiangyou, Sichuan. The authors list the age as "? M. Jurassic", which should stand for Middle(?) Jurassic, as the Jiangyou area is all Jurassic (e.g. fig. 3 of Luo et al., 2014). This would make IVPP V893 from either the Shangshaximiao, Xiashaximiao or Qianfoya Formations, which in the absence of geological description will probably only be resolved via more C. minor fossils or detailed stratigraphic studies of Jiangyou. Chang and Jin (1996) listed the horizon as "?Qianfoyan Series". Liu and Yeh call this "Carnosauria indet." under the Huenian system of carnosaurs being large theropods, though they correctly say it "represents a small carnivorous dinosaur." They say only that "For comparison with the Megalosaurian phalangeals it is shorter and stouter than the latter", which might be based on comparison to Allosaurus given the limited number of 'megalosaurian' manual unguals described at the time, unless it was being compared to pedal unguals where such a statement would also always be true. Comparing proximal height (excluding flexor tubercles), IVPP V893 is ~21% of the size of Afrovenator's holotype, which may mean this is a juvenile if correctly assigned to Megalosauroidea.
Comparison- Compared to ceratosaurs (Saltriovenator, Eoabelisaurus; elaphrosaurs have reduced unguals so are not relevant), IVPP V893 has a less prominent flexor tubercle with no proximal notch and a more concave artcular surface. Carnosaurs have more prominent flexor tubercles (except Mapusaurus) and are transversely thicker (at least in Allosaurus and Sinraptor). This includes the possibly contemporaneous Yangchuanosaurus shangyouensis (CV 00214). Of roughly contemporaneous Chinese taxa with preserved manual unguals, Xuanhanosaurus' ungual I has only been figured in a mostly dorsal view while Kaijiangosaurus' figured ungual is almost flat proximally and has a sharp angle ventrally like ungual II of Asfaltovenator. Most Jurassic coelurosaurs also have more prominent flexor tubercles (Bicentenaria, Zuolong, Tanycolagreus, Juravenator, Guanlong, Ornitholestes, archaeopterygids, scansoriopterygids), and of those that don't- Compsognathus (digit I), Shishugounykus (digit I) and Sciurumimus (digit I) have a proximally projected dorsal lip and shallower claw base, with the first having a proximodistally longer tubercle and the last a shorter tubercle; Aorun (digit II) has almost no tubercle and less recurvature; and Shishugounykus (digit II) has a proximally projected dorsal lip and proximodistally shorter tubercle. The most similar unguals to IVPP V893 are Tugulusaurus (digit I), Dubreuillosaurus (digit I or II), Leshanosaurus (digit I) and Afrovenator (digit I), of which the latter is closest but has a slightly deeper tubercle and more distally placed and ventrally angled (proximally) vascular groove.
References- Liu and Yeh, 1957. Two new species of Ceratodus from Szechuan, China. Vertebrata PalAsiatica. 1(4), 305-311.
Chang and Jin, 1996. Mesozoic fish faunas of China. In Arratia and Viohl (eds.). Mesozoic Fishes - Systematics and Paleoecology. Friedrich Pfeil. 461-478.
Luo, Qi, Zhang, Wang and Hen, 2014. Detrital zircon U-Pb ages of Late Triassic-Late Jurassic deposits in the western and northern Sichuan Basin margin: constraints on the foreland basin provenance and tectonic implications. International Journal of Earth Sciences. 103, 1553-1568.

Afrovenator Sereno, Wilson, Larsson, Duthell and Sues, 1994
A. abakensis Sereno, Wilson, Larsson, Duthell and Sues, 1994
Bathonian-Oxfordian, Middle-Late Jurassic
In Gall, Tiourarén Formation of the Irhazer Group, Niger
Holotype- (UC UBA 1) (6.76 m) skull (~840 mm) (lacking premaxilla, nasal, frontal, parietal and quadratojugal), prearticular, axis, third cervical cervical vertebra, fourth cervical vertebra, eighth cervical vertebra, cervical rib, first dorsal vertebra, fourth dorsal vertebra, seventh dorsal centrum, eighth dorsal centrum, eleventh dorsal vertebra, twelfth dorsal vertebra, two dorsal ribs, proximal caudal vertebra, two mid caudal vertebrae, ten distal caudal vertebrae, fifteen chevrons, humerus (~400 mm), distal radius, distal ulna, semilunate carpal, metacarpal I (62 mm), phalanx I-1 (112 mm), manual ungual I (~135 mm), metacarpal II (135 mm), proximal phalanx II-1, manual ungual II (~118 mm), phalanx III-3 (53 mm), manual ungual III (~66 mm), incomplete ilium (567 mm), incomplete pubis (644 mm), ischium (533 mm), femur (760 mm), tibia (~687 mm), fibula, astragalus lacking ascending pr., calcaneum, metatarsal I (103 mm), phalanx II-1 (122 mm), pedal ungual II (76 mm), (metatarsal III ~344 mm), metatarsal IV (321 mm), phalanx IV-1 (90 mm), phalanx IV-2 (106 mm), phalanx IV-3 (87 mm)
Referred- ?(MUPE HB-118) tooth (37.5x18.6x9.9 mm) (Serrano-Martínez, Vidal, Scisio, Ortega and Knoll, 2016)
?(MUPE HB-125) tooth (~20.3x8.7x5.3 mm) (Serrano-Martínez, Vidal, Scisio, Ortega and Knoll, 2016)
?(MUPE HB-142) tooth (~31x14.1x9.2 mm) (Serrano-Martínez, Vidal, Scisio, Ortega and Knoll, 2016)
?(TP4-5) tooth (42.5x19.9x12.7 mm) (Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015)
?(TP4-8) tooth (~62x26.6x13.8 mm) (Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015)
?(TP4-10) tooth (~37x9.3x20.7 mm) (Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015)
?(TP4-12) maxillary fragment (Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015)
Diagnosis- (after Sereno et al., 1994) maxilla with lobe-shaped anterior margin of antorbital fossa; third cervical with a low, rectangular neural spine; very flat semilunate carpal; metacarpal I with a broad flange for articulation
against metacarpal II.
(after Carrano et al., 2012) posterior extension of distal pubic expansion inset from lateral surfaces of anterior extension, forming pair of conjoined, posterior flanges.
Comments- Though originally identified as Hauterivian-Barremian and called the Tiouraren beds by Sereno et al. (1994), the Tiouraren Formation has been reinterpreted as Bathonian-Oxfordian (Rauhut and Lopez-Arbarello, 2009). Note Sereno et al. measured manual ungual length as "measured perpendicular to the alticular end", so maximum length from proximodorsal edge to distal tip is estimated from their figure here.
Sereno et al. placed Afrovenator as a non-megalosaurian megalosauroid, which takes five more steps in Carrano et al.'s matrix (whose phylogeny is followed here). Rauhut (2003) found it to be closer to allosauroids than megalosauroids, which takes 3 more steps when constrained. Each possibility is quite likely.
References- Sereno, Wilson, Larsson, Dutheil and Sues, 1994. Early Cretaceous dinosaurs from the Sahara. Science. 266, 267-271.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Rauhut and Lopez-Arbarello, 2009. Considerations on the age of the Tiouaren Formation (Iullemmeden Basin, Niger, Africa): Implications for Gondwanan Mesozoic terrestrial vertebrate faunas. Palaeogeography, Palaeoclimatology, Palaeoecology. 271, 259-267.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015. New theropod remains from the Tiourarén Formation (?Middle Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans. Proceedings of the Geologists' Association. 126(1), 107-118.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.
Serrano-Martínez, Vidal, Scisio, Ortega and Knoll, 2016 (online 2015). Isolated theropod teeth from the Middle Jurassic of Niger and the early dental evolution of Spinosauridae. Acta Palaeontologica Polonica. 61(2), 403-415.

Dubreuillosaurus Allain, 2005
= "Calvadosaurus" Holtz, 2007
D. valesdunensis (Allain, 2002) Allain, 2005
= Poekilopleuron valesdunensis Allain, 2002
Middle Bathonian, Middle Jurassic
Calcaire de Caen Formation, France
Holotype- (MNHN 1998-13) partial skull (~490 mm), dentary splenials, surangular impression, angular, incomplete cervical centrum (45 mm excluding anterior ball), incomplete posterior cervical vertebra (50 mm excluding anterior ball), axial rib, proximal cervical ribs, two partial anterior dorsal centra, three partial anterior dorsal neural arches, two posterior dorsal centra (70 mm), dorsal ribs, two gastralia, partial second sacral vertebra, partial third sacral vertebra (85 mm), partial fifth sacral vertebra, fifth sacral rib, proximal caudal neural arch, seven mid caudal vertebrae (60-65 mm), two distal caudal vertebrae (38, 40 mm), two chevrons (110, 90 mm), partial scapula, manual ungual I or II (83 mm on curve), incomplete femur (~450 mm), proximal tibia, incomplete fibula, phalanx III-1 (65 mm), metatarsal V (58 mm)
Diagnosis- (after Allain, 2005) very low skull, at least three times longer than high; double curvature of the anterodorsal margin of the maxillary nasal ramus; postorbital ventral process with U-shaped cross section; parietals not visible in lateral view; straight medial margin of the upper temporal fenestra; absence of paraquadrate fenestra; well developed ventral process on the jugal ramus of the ectopterygoid; deeply grooved posterior margin of the ectopterygoid ahead of the subtemporal fenestra; large external mandibular fenestra; mylohyoid foramen largely opened anteroventrally; femoral head medioventrally directed; convex anterior surface of distal end of femur.
(after Carrano et al., 2012) ventral notch in posterior outline of braincase between exoccipitalopisthotics and basioccipital.
References- Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. Museum National d'Histoire Naturelle Laboratoire de Paleontologie. 329 pp.
Allain, 2002b. Discovery of a megalosaur (dinosauria, Theropoda) in the Middle Bathonian of Normandy (France) and its implications for the phylogeny of basal Tetanurae. Journal of Vertebrate Paleontology. 22(3), 548-563.
Allain, 2005. The postcranial anatomy of the megalosaur Dubreuillosaurus valesdunensis (Dinosauria Theropoda) from the Middle Jurassic of Normandy, France. Journal of Vertebrate Paleontology. 25(4), 850–858.
Holtz, 2007. Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages. Random House. 432 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.

Leshansaurus Li, Peng, Ye, Jiang and Huang, 2009
L. qianweiensis Li, Peng, Ye, Jiang and Huang, 2009
Kimmeridgian?, Late Jurassic
Xiaogu, Shangshaximiao Formation, Sichuan, China
Holotype- (QW200701) (~6.2 m) incomplete skull, posterior mandible, three teeth, hyoid (95 mm), atlas, axis (78 mm), third cervical vertebra (80 mm), fourth cervical vertebra (76 mm), fifth cervical vertebra (80 mm), ninth cervical vertebra (70 mm), tenth cervical vertebra (60 mm), first dorsal vertebra (70 mm), second dorsal vertebra (80 mm), third dorsal vertebra (87 mm), fourth dorsal vertebra (98 mm), fifth dorsal vertebra (99 mm), sixth dorsal vertebra (103 mm), seventh dorsal vertebra (105 mm), eighth dorsal vertebra (107 mm), ninth dorsal vertebra (102 mm), tenth dorsal vertebra (97 mm), twelfth dorsal vertebra (102 mm), fourteenth dorsal vertebra (82 mm), most dorsal ribs (to 600 mm), sacrum (100, 95, 90, 90, 95 mm), two mid caudal vertebrae, chevron, metacarpal I, phalanx I-1, manual ungual I, phalanges II-1, metacarpal III (70 mm), phalanx III-3, manual ungual III, incomplete ilium, pubis (585 mm), proximal ischial fragment, femur (600 mm), tibia (580 mm), incomplete fibula, metatarsal II (275 mm), phalanx II-1, phalanx II-2, pedal ungual II, phalanx III-1, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-4, pedal ungual IV
Paratype- (QW200702) (juvenile) femur (220 mm)
Diagnosis- (after Li et al., 2009) supraoccipital with sharp midline ridge; frontal 2.4 times longer than wide; slender basipterygoid processes; atlantal intercentrum horseshoe-shaped; slender dorsal transverse processes; thin dorsal and sacral neural spines; distinct ventral sacral keel; conspicuous medial ridge along acetabular edge of ilium.
Comments- Li et al. (2009) assigned this taxon to Sinraptoridae, but Cau (online, 2009) referred it to Megalosauridae instead, as the sister taxon to Megalosaurus. As he noted, the elongate anterior maxillary ramus, lack of medial fenestrae in the antorbital fossa, long frontal, only slightly decurved paroccipital processes, and long dorsal centra are all more similar to megalosauroids than sinraptorids. In addition, I note a ventral keel is present in the sacra of Megalosaurus and "Brontoraptor", and the medial acetabular ridge is a megalosauroid synapomorphy in Benson's analysis. Carrano et al. (2012) found it to be an afrovenatorine particularly similar to Piveteausaurus. Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it as either a basal spinosaurid or a eustreptospondyline.
References- Cau, online 2009. http://theropoda.blogspot.com/2009/12/leshansaurus-li-et-al-2009-sinraptoride.html
Li, Peng, Ye, Jiang and Huang, 2009. A new carnosaur from the Late Jurassic of Qianwei, Sichuan, China. Acta Geologica Sinica. 83(9), 1203-1213.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.

Magnosaurus Huene, 1923
M. nethercombensis (Huene, 1923) Huene, 1932
= Megalosaurus nethercombensis Huene, 1923
= Streptospondylus nethercombensis (Huene, 1923) Paul, 2010
Early Bajocian, Middle Jurassic
Inferior Oolite Formation, England
Holotype- (OUM J12143) (~4.6 m; adult) incomplete dentaries, partial posterior dorsal vertebra, rib impressions, partial mid caudal vertebra (~66 mm), ilial fragment, pubes (one proximal; one incomplete), distal femora, tibiae (one proximal; one incomplete; 490 mm), distal fibula, fragments
Diagnosis- (after Rauhut, 2000) differs from Eustreptospondylus in the proximal extent of the pubic symphysis.
(after Sadleir et al., 2008) differs from Eustreptospondylus in having interdental plates which are taller than long; pubis with greater transverse expansion at acetabular margin in proximal view; lateral margin of distal femur concave in posterior view; dorsoventrally extending ridge on lateral surface of cnemial crest.
(after Benson, 2010) anteroposteriorly elongate foramina located ventrally on the lateral surface of the dentary.
Comments- In Carrano et al.'s (2012) matrix, it only takes a single step to move Magnosaurus from Afrovenatorinae and into Eustreptospondylinae as in Rauhut (2000). Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it as either a basal spinosaurid, eustreptospondyline or sister to Allosauria.
References- Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of the Geological Society of America. 34, 449-458.
Huene, 1926a. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata. 29, 167 pp.
Huene, 1926b. On several known and unknown reptiles of the order Saurischia from England and France. Annal and Magazine of Natural History, ser. 9. 17, 473-489.
Huene, 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. Monographien zur Geologie und Palaeontologie. 4(1), viii + 361 pp.
Waldman, 1974. Megalosaurids from the Bajocian (Middle Jurassic) of Dorset. Palaeontology 17, 325-339.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. Museum National d'Histoire Naturelle Laboratoire de Paleontologie. 329 pp.
Sadleir, Barrett and Powell, 2008. The anatomy and systematics of Eustreptospondylus oxoniensis, a theropod dinosaur from the Middle Jurassic from Oxfordshire, England. Monograph of the Palaeontological Society. 82 pp.
Benson, 2010. The osteology of Magnosaurus nethercombensis (Dinosauria, Theropoda) from the Bajocian (Middle Jurassic) of the United Kingdom and a re-examination of the oldest records of tetanurans. Journal of Systematic Palaeontology. 8(1), 131-146.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Hendrickx, Mateus and Araújo, 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627-642.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.

Piveteausaurus Taquet and Welles, 1977
P. divesensis (Walker, 1964) Taquet and Welles, 1977
= Eustreptospondylus divesensis Walker, 1964
= Proceratosaurus divesensis (Walker, 1964) Paul, 1988
Late Callovian, Middle Jurassic
Marnes de Dives, France
Holotype- (MNHN 1920-7) (adult) frontals, parietals, braincase
Comments- Carrano et al. (2012) note that because the braincase is so similar to Leshansaurus, and other known megalosaurid braincases (Eustreptospondylus and Dubreuillosaurus) are from juveniles, the validity of this species is uncertain.
This taxon is generally placed in Megalosauridae (Taquet and Welles, 1977; Kurzanov, 1989), which was recently found in Benson's (2008, 2010) and Carrano et al.'s (2012) phylogenetic analyses. Exceptions are Allain (2002) and Paul (1988), who found it to be carnosaurian. Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it as either a basal spinosaurid or carcharodontosaurid outside Carcharodontosaurinae.
References- Walker, 1964. Triassic reptiles from the Elgin area: Ornithosuchus and the origin of carnosaurs. Philosophical Transactions of the Royal Society of London B. 248, 53-134.
Taquet and Welles, 1977. Redescription du crâne de dinosaure théropode de dives (Normandie). Annales de Paléontologie (Vertébrés). 63(2), 191-206.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York. 464 pp.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. Museum National d'Histoire Naturelle Laboratoire de Paleontologie. 329 pp.
Benson, 2008. A new theropod phylogeny focussing on basal tetanurans, and its implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.

Poekilopleuron Eudes-Deslongchamps, 1837
= "Poecilopleuron" Eudes-Deslongchamps vide Bronn, 1937
P. bucklandii Eudes-Deslongchamps, 1837
= "Poecilopleuron bucklandi" Eudes-Deslongchamps vide Bronn, 1937
= Liopleurodon bucklandi (Eudes-Deslongchamps, 1837) Sauvage, 1873
= Megalosaurus poikilopleuron Huene, 1923
Middle Bathonian, Middle Jurassic
Calcaire de Caen Formation, France
Holotype- (destroyed; plastoholotype MNHN 1897-2) cervical ribs (lost), dorsal ribs (lost), gastralia, twenty-one caudal vertebrae (lost), six chevrons (lost), scapula (lost), humerus (~306 mm), radius (170 mm), ulna (182 mm), metacarpal I (58 mm), phalanx I-1 (66 mm; lost), femur (lost), distal tibia (lost), partial fibula (lost), astragalus (lost), phalanx I-1, pedal ungual I, phalanx II-1, phalanx II-2, pedal ungual; II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-2, phalanx IV-3, pedal ungual IV, ten gastroliths
Diagnosis- (after Allain and Chure, 2002) mid caudal neural spines as long as their corresponding centrum length; deltopectoral crest extending to midlength of humerus; ulna lacking olecranon process; distal end of radius as wide as proximal end; strong ulnar process at midlength of posteromedial edge of radius; convex lateral margin of ascending process of astragalus.
(after Rauhut, 2000) metacarpal I with small lateral flange behind distal articular facet.
Comments- YPM 4839 (2427) is listed in Carrano (1998) as Poekilopleuron? valens with metatarsal (180 mm) and humeral (323 mm) measurements, but the online catalogue indicates YPM 4839 is a cast of the Poekilopleuron bucklandii holotype perhaps measuring the ulna and humerus. YPM 2427 is a Pteranodon proximal ulna, so the inclusion of this number by Carrano is unexplained.
Traditionally considered a megalosaurid (Allain, 2002; Holtz et al., 2004), Benson (2008, 2010) found it to be a carnosaur and more recently (Benson et al., 2010) a sinraptorid. Carrano et al. (2012) recovered it as an afrovenatorine, though this is highly uncertain as only one more step is needed for it to be a piatnitzkysaurid, allosaur or any other kind of megalosaurid. Moving it to Sinraptoridae takes 4 more steps so is still quite possible. Most recently, Rauhut et al. (2024) used the Mesozoic Tetrapod Group Theropod Matrix to recover it as carcharodontosaurid sister to Concavenator.
References- Bronn, 1837. Lethaea Geognostica oder Abbildungen und Beschreibungen der für die Gebirge-Formationen bezeichnendsten Versteinerungen. 1, 1-544.
Eudes-Deslongchamps. 1837. Mémoire sur le Poekilopleuron bucklandi, grande saurien fossile, intermédiare entre les crocodiles et les lézards, découvert dans les carrières de la Maladrerie, près Caen, au mois de juillet 1835. Mémoires de la Société Linnéenne Normandie. 6, 1-114.
Sauvage, 1873. Notes sur les reptiles fossiles. Bulletin de la Société Géologique de France, série 3. 1, 365-386.
Huene, 1923. Carnivorous Saurischia in Europe since the Triassic. Bulletin of the Geological Society of America. 34, 449-458.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology, biomechanics, and modern analogs. PhD Thesis, The University of Chicago. 424 pp.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis. University of Bristol. 440 pp.
Allain, 2002a. Les Megalosauridae (Dinosauria, Theropoda). Nouvelle découverte et révision systématique: Implications phylogénétiques et paléobiogéographiques. PhD thesis. Museum National d'Histoire Naturelle Laboratoire de Paleontologie. 329 pp.
Allain, 2002b. Discovery of a megalosaur (dinosauria, Theropoda) in the Middle Bathonian of Normandy (France) and its implications for the phylogeny of basal Tetanurae. Journal of Vertebrate Paleontology. 22(3), 548-563.
Allain and Chure, 2002. Poekilopleuron bucklandi, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy. Paleontology. 45(6), 1107-1121.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Benson, 2008. A new theropod phylogeny focussing on basal tetanurans, and its implications for European 'megalosaurs' and Middle Jurassic dinosaur endemism. Journal of Vertebrate Paleontology. 28(3), 51A.
Benson, 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. 158(4), 882-935.
Benson, Brusatte and Carrano, 2010. A new clade of large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften. 97, 71-78.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Rauhut, Bakirov, Wings, Fernandes and Hübner, 2024. A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan. Zoological Journal of the Linnean Society. 201(4), DOI: 10.1093/zoolinnean/zlae090.
P? sp. indet. (Eudes-Deslongchamps, 1837)
Middle Jurassic
Grande Oolithe, France
Material- (destroyed) tooth
Comments- This cannot be compared to Poekilopleuron, so cannot be referred to the genus.
Reference- Eudes-Deslongchamps. 1837. Mémoire sur le Poekilopleuron bucklandi, grande saurien fossile, intermédiare entre les crocodiles et les lézards, découvert dans les carrières de la Maladrerie, près Caen, au mois de juillet 1835. Mémoires de la Société Linnéenne Normandie. 6, 1-114.
P? sp. indet. (Owen, 1854)
Hauterivian-Barremian, Early Cretaceous
Weald Clay, England
Material- (Royal College of Surgeons coll.) partial dorsal centrum
Comments- Based on its age, this material is unlikely to be Poekilopleuron, and cannot be compared to the genus in any case.
Reference- Owen, 1854. Descriptive catalogue of the fossil organic remains of reptilia and pisces contained in the Museum of The Royal College of Surgeons of England. 184 pp.

Spinosauridae Stromer, 1915
Definition- (Spinosaurus aegyptiacus <- Torvosaurus tanneri) (modified from Sereno, 1998)
Other definitions- (Spinosaurus aegyptiacus <- Megalosaurus bucklandii, Allosaurus fragilis, Passer domesticus) (Holtz et al., 2004)
(Spinosaurus aegyptiacus <- Torvosaurus tanneri, Allosaurus fragilis, Passer domesticus) (Allain et al., 2012)
(Spinosaurus aegyptiacus <- Megalosaurus bucklandii, Allosaurus fragilis) (Rauhut and Pol, 2019)
= Baryonychidae Charig and Milner, 1986
= Irritatoridae Martill et al., 1996
= Sigilmassasauridae Russell, 1996
= Spinosauridae sensu Holtz et al., 2004
Definition- (Spinosaurus aegyptiacus <- Megalosaurus bucklandii, Allosaurus fragilis, Passer domesticus)
= Spinosauridae sensu Allain et al., 2012
Definition- (Spinosaurus aegyptiacus <- Torvosaurus tanneri, Allosaurus fragilis, Passer domesticus)
= Spinosauridae sensu Rauhut and Pol, 2019
Definition- (Spinosaurus aegyptiacus <- Megalosaurus bucklandii, Allosaurus fragilis)
Diagnosis- (after Rauhut, 2000) Dentary with strongly developed anterior expansion; anterior dentary teeth much larger than the relatively small and closely spaced posterior teeth; medial alveolar border is as high as the lateral border and formed by a sheet of bone of the dentary, rather than by separately ossified interdental plates; teeth almost round in basal cross section and only slightly recurved; very long premaxillae, forming a rostral rosette; seven premaxillary teeth; ventral margin of premaxilla strongly concave; anterior ramus of maxilla strongly elongated; angle between anterior and ventral ramus of the lacrimal less than 45°; dorsal vertebrae with several small vertical laminae connecting the transverse process with the neural spine dorsally; humerus extremely robust, with strongly expanded internal tuberosity and distal condyles; ulna with a broad and very strongly developed olecranon process; ischium with a long and low obturator flange.
Comments- Sereno's (in press) definition is the same as Holtz et al.'s (2004) except for the substitution of Torvosaurus for Megalosaurus. Sereno's has an advantage that Torvosaurus is more often included in analyses than Megalosaurus (Sereno, 1999; Rauhut, 2000; Allain, 2002), and has a more certain phylogenetic position. Incidentally, I believe a node-based Spinosauridae may be a better idea, for this stem-based one could include Chilantaisaurus (Rauhut, 2000, 2003), coelophysoids (Paul, 1988), tyrannosaurids (Walker, 1964), carcharodontosaurids (Bonaparte et al., 1990) and other taxa proposed to be more closely related to Spinosaurus than megalosaurids, allosaurids or birds.
Ex-spinosaurids- Candeiro et al. (2004) referred two teeth from the Adamantina Formation of Brazil (UFRJ-DG 354-Rd and UFRJ-DG 372-Rd) to Spinosauridae, but Machado et al. (2008) considered them more like to be crocodyliforms. Canudo et al. (2004) and Salgado et al. (2009) referred a tooth (Endemas-PV 6) from the Cerro Lisandro Formation of Argentina to Spinosauridae and cf. Spinosauridae respectively, but Hasegawa et al. (2010) referred it to Hamadasuchus. Buffetaut (2008) considered teeth from the Tendaguru Formation of Tanzania (MB R 1084 and possibly 1091) previously referred to Labrosaurus stechowi to be spinosaurid, but Rauhut (2011) noted they were more similar to other stechowi teeth and thus possibly ceratosaurid. Barrett et al. (2010) described cervical NMV P221081 from the Eumeralla Formation of Australia as a spinosaurid, but it was later referred to Megaraptoridae by Poropat et al. (2019). Hone et al. (2010) described a tooth (XMDFEC V0010) from the Majiacun Formation of China as a baryonychine, but Kubota et al. (2017) reassigned it to a more generic theropod.
References- Candeiro, Abranches, Abrantes, Avilla, Martins, Moreira, Torres and Bergqvist, 2004. Dinosaur remains from western Sao Paulo State, Brazil (Bauru Basin, Adamantina Formation, Late Cretaceous). Journal of South American Earth Sciences. 18, 1-10.
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.
Rayfield and Milner, 2005. Cranial functional morphology of spinosaurid "crocodile-mimic" dinosaurs. Journal of Vertebrate Paleontology. 25(3), 102A.
Rayfield and Milner, 2006. The evolution of piscivory in theropod dinosaurs. Journal of Vertebrate Paleontology. 26(3), 114A.
Buffetaut, 2008. Spinosaurid teeth from the Late Jurassic of Tengaduru, Tanzania, with remarks on the evolutionary and biogeographical history of the Spinosauridae. In Mazin, Pouch, Hantzpergue and Lacombe (eds.). Mid-Mesozoic life and environments. Documents des Laboratoires de Geologie Lyon. 164, 26-28.
Machado, Campos and Kellner, 2008. On a theropod scapula (Upper Cretaceous) from the Marília Formation, Bauru Group, Brazil. Paläontologische Zeitschrift. 82(3), 308-313.
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.
Barrett, Benson, Rich and Vickers-Rich, 2010. A definitive spinosaurid theropod from the Lower Cretaceous of Australia and its implications for Gondwanan paleobiogeography. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 57A.
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.
Hone, Xu and Wang. 2010. A probable baryonychine (Theropoda: Spinosauridae) tooth from the Upper Cretaceous of Henan Province, China. Vertebrata PalAsiatica. 48, 19-26.
Rauhut, 2011. Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania). Palaeontology. 86, 195-239.
Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften. 99(5), 369-377.
Anduza and Fowler, 2014. How might spinosaurids have caught fish? Testing behavioral inferences through comparisons ith modern fish-eating tetrapods. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 80.
Kubota, Takakuwa and Hasegawa, 2017. Second discovery of a spinosaurid tooth from the Sebayashi Formation (Lower Cretaceous), Kanna Town, Gunma Prefecture, Japan. Bulletin of Gunma Museum of Natural History. 21, 1-6.
Poropat, White, Vickers-Rich and Rich, 2019. New megaraptorid (Dinosauria: Theropoda) remains from the Lower Cretaceous Eumeralla Formation of Cape Otway, Victoria, Australia. Journal of Vertebrate Paleontology. 39, e1666273.
Rauhut and Pol, 2019. Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs. Scientific Reports. 9:18826.

"Vectispinus" Naish, online 2022
Late Barremian, Early Cretaceous
Compton Chine, Cowleaze Chine Member, Vectis Formation, England
Material- (IWCMS 2018.30; White Rock spinosaurid) (~13 m) partial anterior dorsal vertebra (69.4 mm without condyle), two ?dorsal rib fragments, two synsacral centra (~156, ~142 mm), sacral or caudal central fragment, partial proximal caudal vertebra, two ilial fragments, two long bone fragments, four fragments
Comments- Discovered over several months in 2018, this was first mentioned by Austen and Batten (2018) as "theropod bones from Compton Bay, Isle of Wight (Fig. 46.1) (J.A.F. Lockwood, pers. comm. 2018)" with one element figured as 46.1 "Probable theropod pelvic fragment showing curved boring b (Vectis Formation at junction with the Wessex Formation, Compton Bay. IWCMS 2018.30)." Barker et al. (2022) described the remains as Spinosauridae indet., "with enormous dimensions similar to those of the Spinosaurus holotype and exceeding those of the largest European theropods previously reported." While no total length estimate was provided, scaling the proximal caudal centrum height from Baryonyx in their table 5 (assuming a 9.1 m Baryonyx) leads to 13.2 m which is identical to this site's estimate for the Spinosaurus type which has equal sacral centrum length to IWCMS 2018.30. Hence the 13 m estimate given here. Naish (online 2022) stated "Dinosaur species tend to be unique to specific geological formations, and ... there’s no indication from anatomy that these remains belong to any of the previously known Wealden spinosaurids ... For these reasons, we can be mildly confident that this is a new animal, and we did at times consider the possibility of giving it a name (cough 'Vectispinus')." This is of course an intentional nomen nudum for reasons such as ICZN article 11.5 ("a name must be used as valid for a taxon when proposed") and 16.1 ("Every new name published after 1999, including new replacement names (nomina nova), must be explicitly indicated as intentionally new").
Barker et al. (2022) used Cau's Wessex baryonychine matrix to recover it very weakly supported as a spinosaurine basal to Ichthyovenator and Spinosaurus. They found "Three characters were shared between the White Rock spinosaurid and other spinosaurines, all from the anterior caudal series: the presence of centrodiapophyseal laminae (Ch. 358:1), the presence of prezygodiapophyseal laminae (Ch. 626:1), and the presence of a deep prezygocentrodiapophyseal fossa (Ch. 1605:1)", but that Ichthyovenator and Spinosaurus "appear to show that fossae and laminae become less prominent in the more posterior parts of the axial skeleton" and that the caudal of IWCMS 2018.30.3 [is considered to be] to be more anteriorly placed than any of the known caudal elements of Riparovenator or Vallibonavenatrix" that were scored for baryonychine proximal caudal morphology. Thus the characters may be present in the currently unknown proximalmost baryonychine caudals as well.
References- Austen and Batten, 2018. English Wealden fossils: An update. Proceedings of the Geologists' Association. 129, 171-201.
Barker, Lockwood, Naish, Brown, Hart, Tulloch and Gostling, 2022. A European giant: A large spinosaurid (Dinosauria: Theropoda) from the Vectis Formation (Wealden Group, Early Cretaceous), UK. PeerJ. 10:e13543.
Naish, online 2022. https://tetzoo.com/blog/2022/6/8/giant-spinosaurid-dinosaur-from-the-isle-of-wight

undescribed Spinosauridae (Isasmendi, Sáez-Benito, Torices and Pereda-Suberbiola, 2019)
Late Barremian-Early Aptian, Early Cretaceous
Enciso Group, Virgen del Villar-2, Peña Cárcena, Igea, La Rioja, Spain
Material- (CPI coll.; Villar) incomplete skeleton (University of the Basque Country, 2024 online)
Late Barremian-Early Aptian, Early Cretaceous
Enciso Group, Marino 2, Peña Cárcena, Igea, La Rioja, Spain
(CPI coll.; Garras) (medium-sized) palatal elements, (?)dorsal vertebrae, two partial dorsal ribs, caudal vertebrae, ilium, pubis, femur, tibia, incomplete fibula, three metatarsals (~281 mm), phalanges, three manual and pedal unguals (pedal ~74 mm) (Isasmendi, Sáez-Benito, Torices and Pereda-Suberbiola, 2019)
Comments- Isasmendi et al. (2019) mentioned "The “Garras” excavation carried out in the summer of 2018 in Peña Cárcena" (translated) and stated that partial skeleton was partially articulated. They further say "its affinity with other theropods cannot be determined, but it would correspond to a medium-sized animal." Isasmendi et al. (2021) specified "partially articulated skeletons belonging to an indeterminate theropod and to other mid- to large-sized dinosaur have been recovered from Marino 2 and Colladillos 1 sites, respectively", with the former being Garras that is also photographed in their Figure 2A as "remains of a partially articulated indeterminate theropod." The latter figure shows two dorsal ribs, a fibula, three metatarsals and a pedal ungual. University of the Basque Country's 2024 press release for Riojavenatrix mentions "other spinosaurid fossils found at Igea, such as the partial skeleton called “Garras” at Peña Cárcena, and the one excavated in recent years at Virgen del Villar 2 (very close to Virgen del Villar 1), which has been given the informal name Villar. The latter is “one of the most complete skeletons known in the European and world record”. The “Garras” material is in the final stages of study before publication and the Villar material is being prepared by the lab team members." Whether these skeletons belong to Riojavenatrix or the Enciso spinosaurine, or new taxa, remains to be determined although they should both be comparable to the former genus.
References- Isasmendi, Sáez-Benito, Torices and Pereda-Suberbiola, 2019. Restos
oseos de dinosaurios ter
opodos del Cret
acico Inferior de Igea (La Rioja). Zubía. 31, 181-186.
Isasmendi, Navarro-Lorbés, López-Horgue, Sáez-Benito, Viera, Torices and Pereda-Suberbiola, 2021. Updating the geological and stratigraphic occurrences of La Rioja dinosaur sites. Palaeobiodiversity of the Lower Cretaceous Enciso Group at Peña Cárcena (Igea, Spain). Comunicações Geológicas. 108(Especial I), 91-96.
University of the Basque Country, 2024 online. UPV/EHU researchers discover a new species of carnivorous dinosaur in La Rioja. Campusa. 3-12-2024.
Isasmendi, in prep.. [title]. PhD thesis. University of the Basque Country. [pp].

unnamed possible spinosaurid (Bittencourt and Kellner, 2003)
Albian, Early Cretaceous
Romualdo Formation of Santana Group, Brazil
Material- (MN 4743-V) (subadult) incomplete third and fourth sacral vertebrae, fifth sacral vertebra, six proximal caudal vertebrae, three proximal chevrons
Comments- Bittencourt and Kellner (2003) excluded MN 4743-V from Ceratosauria based on the lack of a synsacrum and a ventral midline groove on the proximal caudal centra. They excluded it from Coelurosauria based on the transversely rounded ventral sacral margin and anterior chevrons with a roblike proximal section. They suggested spinosaurid affinities based on the infradiapophyseal laminae on the proximal caudals, as well as undisclosed similarities to an undescribed specimen (MN 4819-V) tentatively referred to that family.
However, a ventral midline groove on proximal caudal centra is present in nearly every theropod, even Shuvuuia (only the first two caudals are keeled). It's absence is only reported for Elaphrosaurus, Carnotaurus, Eustreptospondylus, Cristatusaurus, Shaochilong and Ornithomimus? sedens. Additionally, synsacra are highly variable in theropods (seemingly due to age and individual variation). "Ceratosaurs" (sensu lato) that lack complete sacral fusion include Liliensternus, some Coelophysis rhodesiensis specimens, Dilophosaurus and abelisaurids. Tyrannosauroids lack ventrally flattened sacral centra, a condition that has also been reported for SMNK 2349 PAL and Ornitholestes. Thus, the flat venter may be a maniraptoriform character, not a coelurosaurian one. I have not observed any difference in the anteroposterior elongation of the proximal end of anterior chevrons between coelurosaurs and non-coelurosaurs. The expansion seen in most coelurosaurs (but not all- eg. Gallimimus) is also found in many other theropods, such as Dilophosaurus, Torvosaurus, Acrocanthosaurus and Allosaurus. This is mostly caused by paired anterior processes at the proximal end. Baryonyx lacks these however, adding further proof to their possible spinosaurid identification.
Reference- Bittencourt and Kellner, 2003. New theropod remains from the Romualdo Member, Santana Formation (Aptian-Albian), Northeastern Brazil. III Simpósio Brasileiro de Paleovertebrados, Resumos. 18.
Bittencourt and Kellner, 2004. On a sequence of sacrocaudal theropod dinosaur vertebrae from the Lower Cretaceous Santana Formation, northeastern Brazil. Arquivos do Museu Nacional Rio de Janeiro. 62(3), 309-320.

undescribed spinosaurid (Kellner, 2001)
Albian, Early Cretaceous
Romualdo Member of Santana Formation, Brazil
Material- (MN 4819-V) dorsal vertebrae, sacrum, caudal vertebrae, incomplete forelimbs including carpal, metacarpals, phalanges and manual ungual I, ilium (553 mm), pubis, ischium, incomplete hindlimbs
Comments- Both Kellner (2001) and Kellner and Bittencourt (2004) incorrectly referred to this specimen as MN 4802-V instead of 4819-V (Machado and Kellner, 2005). Kellner stated the manual ungual I is large and the ischium has an obturator notch. Machado and Kellner stated it has tall neural spines as well. Bittencourt and Kellner stated it was indeterminate but also that it differed from MN 4743-V.
Reference- Kellner, 2001. New information on the theropod dinosaurs from the Santana Formation (Aptian-Albian), Araripe Basin, Northeastern Brazil. Journal of Vertebrate Paleontology. 21(3), 67A.
Bittencourt and Kellner, 2004. On a sequence of sacrocaudal theropod dinosaur vertebrae from the Lower Cretaceous Santana Formation, northeastern Brazil. Arquivos do Museu Nacional Rio de Janeiro. 62(3), 309-320.
Machado and Kellner, 2005. Notes on Spinosauridae (Theropoda, Dinosauria). Anuário do Instituto de Geociências - UFRJ. 28(1), 158-173.
Machado, Kellner and Campos, 2005. Preliminary information on a dinosaur (Theropoda, Spinosauridae) pelvis from the Cretaceous Santana Formation (Romualdo Member) Brazil. II Congresso Latino-Americano de Paleontologia de Vertebrados. Rio de Janeiro, 2005. Boletim de resumos. 161-162.
Machado and Kellner, 2008. An overview of the Spinosauridae (Dinosauria, Theropoda) with comments on the Brazilian material. Journal of Vertebrate Paleontology. 28(3), 109A.

undescribed possible spinosaurid (Sayao, Saraiva, Silva and Kellner, 2011)
Albian, Early Cretaceous
Romualdo Member of Santana Formation, Brazil
Material- (MPSC R-2089) distal femur, proximal tibia, fibula, pedal digit I, metatarsal II, phalanx II-?, metatarsal III, pedal ungual III (35 mm), metatarsal IV, pedal ungual IV (31 mm)
Reference- Sayao, Saraiva, Silva and Kellner, 2011. A new theropod dinosaur from the Romualdo Lagerstatte (Aptian-Albian), Araripe Basin, Brazil. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 187.

unnamed possible spinosaurid (Leonardi and Borgomanero, 1981)
Albian, Early Cretaceous
Romualdo Member of Santana Formation, Brazil
Material- (MN 7021-V; = CB-PVF-089) partial dorsal rib
Comments- This was described as an ornithischian ischium by Leonardi and Borgomanero (1981), but Kellner (1996) felt it needed further preparation to identify which element and dinosaurian clade it represents. This preparation was completed after Borgomanero's death which transferred the specimen from his private collection to the MN. Machado and Kellner (2001a, b) then were able to identify it as a large dinosaur's rib, which they provisionally referred to Spinosauridae as that is the only large dinosaur material currently known from that formation.
References- Leonardi and Borgomanero, 1981. Sobre uma possível ocorrência de Ornitischia na Formação Santana, Chapada do Araripe (Ceará). Revista Brasileira de Geociências. 11, 1-4.
Kellner, 1996. Remarks on Brazilian dinosaurs. Memoirs of the Queensland Museum. 39(3), 611-626.
Machado and Kellner, 2007a. From an ischium to a rib: Review of the osteological evidence of the supposed ornithischian dinosaur from the Araripe Basin, Brazil. 190A.
Machado and Kellner, 2007b. On a supposed ornithischian dinosaur from the Santana Formation, Araripe basin, Brazil. Paleontologia: Cenários de Vida. 1, 291-299.

undescribed Spinosauridae (Mueller, Bussert, David, Klein and Salih, 2011)
Cenomanian, Late Cretaceous
Shendi or Wadi Milk Formation, Sudan
Material- jaw fragments, teeth
Reference- Mueller, Bussert, David, Klein and Salih, 2011. New discoveries and investigations on the Late Cretaceous vertebrate fauna of Northern Sudan. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 163-164.

undescribed possible Spinosauridae (Harris and Russell, 1986 unpublished)
Campanian?, Late Cretaceous
lower member of Lapurr Sandstone (= Turkana Grits), Kenya
Material- (KNM coll.)
Comments- Discovered in a 1985 KNM expedition (Williamson and Savage, 1986; Owusu Agyemang et al., 2019), Harris and Russell (1986) reported a possible spinosaurid from what was then called the Turkana Grits in Kenya. They used this to support a "middle" Cretaceous age. Unfortunately, this document is only an unpublished manuscript which has had low distribution. The second-hand source of Weishampel et al. (1990) listed "cf. Spinosaurus sp." and "?Late Cretaceous ("middle" Cretaceous; Harris and Ruissell MS)." The second edition of the book keeps the same taxonomic list but Weishampel et al. (2004) now states the age as "Turonian-Santonian (Westcott et al. 1993)." The latter has made this a potentially important find because spinosaurids are unknown globally after the Cenomanian. However, Westcott et al. (1993) reports nothing more precise than Cretaceous for these sediments. O'Connor et al. (2011) state "an estimate of Upper Cretaceous (Turonian-early Campanian) is based on comparisons with subsurface geological information and overall faunal composition" while Owusu Agymang et al. report "palynological evidence supports a Campanian–Maastrichtian age" without supporting evidence. Thus both taxonomic identity and stratigraphic age are unevidenced assertions at the moment.
References- Harris and Russell, 1986 unpublished. Preliminary notes on the occurrence of dinosaurs in the Turkana Grits of northern Kenya. Unpublished manuscript. 11 pp.
Williamson and Savage, 1986. Early rift sedimentation in the Turkana Basin, northern Kenya. In Frostick, Renaut, Reid and Tiercelin (eds.). Sedimentation in the African Rifts. Geological Society of London, Special Publication 25, 267-283.
Weishampel, 1990. Dinosaurian distribution. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 63-139.
Wescott, Morley and Karanja, 1993. Geology of the "Turkana grits" in the Lariu Range and Mt. Porr areas, southern Lake Turkana, northwestern Kenya. Journal of African Earth Sciences (and the Middle East). 16, 425-435.
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004. Dinosaur Distribution. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria: Second Edition. University of California Press. 517-606.
O'Connor, Sertich and Manthi, 2011. A pterodactyloid pterosaur from the Upper Cretaceous Lapurr Sandstone, west Turkana, Kenya. Anais da Academia Brasileira de Ciências. 83, 309-315.
Owusu Agyemang, Roberts, Downie and Sertich, 2019 (online 2018). Sedimentary provenance and maximum depositional age analysis of the Cretaceous? Lapur and Muruanachok sandstones (Turkana Grits), Turkana Basin, Kenya. Geological Magazine. 156(8), 1334-1356.

undescribed possible Spinosauridae (Serrano-Martínez, Ortega and Knoll, 2013)
Bathonian-Oxfordian, Middle-Late Jurassic
Tiourarén Formation of the Irhazer Group, Niger
Material- (MUPE HB-87) tooth (23.8x12.7x8.8 mm) (Serrano-Martínez, Ortega and Knoll, 2013; described by Serrano-Martínez, Vidal, Scisio, Ortega and Knoll, in press)
(TP4-2) tooth (17.8x10.7x7.4 mm) (Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015)
(TP4-3) tooth (20.7x8.2x6.4 mm) (Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015)
Comments- Serrano-Martínez et al. (2013) announced what is probably MUPE HB-87 in an abstract, which was later described by Serrano-Martínez et al. (in press). Serrano-Martínez et al. (2015) described two additional teeth, TP4-2 and TP4-3. All of these were viewed as intermediate between basal tetanurines and spinosaurids. They have BW/FABL ratios of 69%, 69% and 78% respectively. HB-87 has mesial and distal serrations which average 20/5 mm and 18/5 mm respectively, the mesial ones which don't continue basally. TP4-2 lacks mesial serrations but has 12 serrations per 5 mm, while TP4-3 lacks serrations. All have sculptured enamel as in spinosaurids, but only the TP specimens have fluting.
References- Serrano-Martínez, Ortega and Knoll, 2013. Isolated theropod teeth from the "Argiles de l'Irhazer" (Middle Jurassic) of Niger. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 210.
Serrano-Martínez, Ortega, Sciscio, Tent-Manclus, Bandera and Knoll, 2015. New theropod remains from the Tiourarén Formation (?Middle Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans. Proceedings of the Geologists' Association. 126(1), 107-118.
Serrano-Martínez, Vidal, Scisio, Ortega and Knoll, in press. Isolated theropod teeth from the Middle Jurassic of Niger and the early dental evolution of Spinosauridae. Acta Palaeontologica Polonica. http://dx.doi.org/10.4202/app.00101.2014

Baryonychinae Charig and Milner, 1986 vide Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998
Definition- (Baryonyx walkeri <- Spinosaurus aegyptiacus) (Holtz et al., 2004; modified from Sereno et al., 1998)
References- Charig and Milner, 1986. Baryonyx, a remarkable new theropod dinosaur. Nature. 324, 359-361.
Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998. A long-snouted predatory dinosaur from Africa and the evolution of the spinosaurids. Science. 282(5392), 1298-1302.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria Second Edition. University of California Press. 71-110.

undescribed possible baryonychine (Fowler, 2007)
Berriasian, Early Cretaceous
Purbeck Limestone Group, England
Material- (DCM-G95a) tooth
Comments- This is slightly labiolingually compressed, recurved and lingually fluted. No serrations are preserved but may have been worn away, and Fowler states it may also be pliosaurid.
References- Fowler, 2007. Recently rediscovered baryonychine teeth (Dinosauria Theropoda): New morphologic data, range extension and similarity to ceratosaurs. Journal of Vertebrate Paleontology. 27(3), 76A.
Fowler, 2007 online. http://www.denverfowler.com/publications/Fowler_2007_SVP.htm

undescribed Baryonychinae (Seeley, 1869)
Late Berriasian-Barremian, Early Cretaceous
Wealden Supergroup, England
Material- (NHMUK 3240) tooth (Fowler, 2007 online)
?(NHMUK 3330) tooth (Fowler, 2007 online)
(NHMUK 10822) tooth (Fowler, 2007)
(NHMUK 26030) tooth (Fowler, 2007 online)
(NHMUK 26031) tooth (Fowler, 2007 online)
(NHMUK 26032) tooth (Fowler, 2007 online)
(NHMUK 36533 in part) two teeth (Fowler, 2007 online)
(NHMUK R212) tooth (Fowler, 2007 online)
(NHMUK R213) tooth (Fowler, 2007 online)
(NHMUK R215) tooth (Lydekker, 1888)
(NHMUK R641) tooth (Fowler, 2007 online)
?(NHMUK R642) tooth (Fowler, 2007 online)
(NHMUK R5226) tooth (Fowler, 2007)
(Woodwardian Museum Ji910) tooth (Seeley, 1869)
Comments- These were referred to Suchosaurus (Seeley, 1869; Lydekker, 1880) or catalogued as crocodylians (recognized by Fowler, 2007), but have inexact published provenence.
References- Seeley, 1869. Index to the fossil remains of Aves, Ornithosauria, and Reptilia from the Secondary system of strata, arranged in the Woodwardian Museum of the University of Cambridge. With a prefatory notice by the Rev. Adam Sedgewick. London, Bell & Daldy. 143 pp.
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.
Fowler, 2007. Recently rediscovered baryonychine teeth (Dinosauria Theropoda): New morphologic data, range extension and similarity to ceratosaurs. Journal of Vertebrate Paleontology. 27(3), 76A.
Fowler, 2007 online. http://www.denverfowler.com/publications/Fowler_2007_SVP.htm

undescribed Baryonychinae (Lydekker, 1888)
Late Berriasian-Valanginian, Early Cretaceous
Hastings Group, England
Material- (NHMUK R635) tooth (Lydekker, 1888)
(NHMUK R1901) two teeth (Fowler, 2007)
References- Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia, and Proterosauria. British Museum of Natural History, London. 309 pp.
Fowler, 2007. Recently rediscovered baryonychine teeth (Dinosauria Theropoda): New morphologic data, range extension and similarity to ceratosaurs. Journal of Vertebrate Paleontology. 27(3), 76A.
Fowler, 2007 online. http://www.denverfowler.com/publications/Fowler_2007_SVP.htm

Baryonychinae indet. (Charig and Milner, 1997)
Berriasian-Early Valanginian, Early Cretaceous
Ashdown Beds Formation of the Hastings Group, England
Material- (BEXHM:1993.485) tooth
Diagnosis- compared to Baryonyx, carinae do not extend to base of crown.
Reference- Charig and Milner, 1997. Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum of London (Geology). 53, 11-70.

Baryonychinae indet. (Martill and Hutt, 1996)
Barremian, Early Cretaceous
Wessex Formation of the Wealden Group, England
Material- (IWCMS 1995.207) tooth (Martill and Hutt, 1996)
(IWCMS 1995.208) tooth (Martill and Hutt, 1996)
(IWCMS 1995.209) tooth (Martill and Hutt, 1996)
(IWCMS 2012.563) dorsal vertebra (Hutt and Newbery, 2004)
(MIWG 3642) two teeth (Martill and Hutt, 1996)
(MIWG 5122) tooth (Martill and Hutt, 1996)
?(MIWG 6527) (~10-11 m) manual phalanx I-1 (Naish, 1999; Naish, Hutt and Martill, 2001)
(UOP.97) tooth (Charig and Milner, 1997)
Comments- The teeth described by Martill and Hutt (1996) as Baryonychidae differ from those in the holotype in having strong labial fluting, but variation of this kind is also known in Cristatusaurus, Spinosaurus and Oxalaia. Thus it should not be used as a diagnostic character for distinguishing spinosaurid species. While they also assign MIWG 5120 to Baryonychidae, Hutt (2001) lists it as Theropoda indet..
Hutt and Newbery (2004) describe a tall-spined dorsal vertebra (cast UOP C001.2004) as Baryonyx walkeri. This vertebra was originally in a private collection but cast as UOP C001.2004, and has since been officially deposited as IWCMS 2012.563 (Mattsson, pers. comm. 2015). It is illustrated in Naish (2011), who stated it differs from the B. walkeri holotype "in possessing
a much taller neural spine and a prominent spinodiapophyseal lamina."
Although often referred to Baryonyx or B. walkeri itself based on the correlation of the Wessex Formation and Upper Weald Clay, the 2021 description of Wesssex baryonychines Ceratosuchops and Riparovenator complicates matters. Neither taxon preserves manual or dorsal material, although if Cristatusaurus is a ceratosuchopsin the tall neural spine would make sense referred to one of the Wessex baryonychines instead of the short-spined Baryonyx.
References- Martill and Hutt, 1996. Possible baryonychid dinosaur teeth from the Wessex Formation (Lower Cretaceous, Barremian) of the Isle of Wight, England. Proceedings of the Geologists' Association. 107, 81-84.
Hutt, 2001. Catalogue of Wealden Group Dinosauria in the Museum of Isle of Wight Geology. In Martill and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association. 411-422.
Naish, Hutt and Martill, 2001. Saurischian dinosaurs 2: Theropods. In Martill and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association (London). 242-309.
Hutt and Newbery, 2004. A new look at Baryonyx walkeri (Charig and Milner, 1986) based upon a recent fossil find from the Wealden. SVPCA 2004. 18.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The Palaeontological Association. 526-559.

undescribed baryonychine (Fowler, 2007 online)
Early Aptian, Early Cretaceous
Atherfield Clay Formation, England
Material- (NHMUK R5165) tooth
Reference- Fowler, 2007 online. http://www.denverfowler.com/publications/Fowler_2007_SVP.htm

Baryonychinae indet. (Torcida, Fuentes, Izquierdo, Montero and Urien, 1997)
Late Hauterivian, Early Cretaceous
Urbión Group, Tenada de la Rosada, Cabezón de la Sierra, Castile and Leon, Spain
Material- (PS.C-15,30) partial tooth (?x16.48x12.48 mm)
(PS.C-15,32) tooth (43.5x21,21x13 mm)
Comments- The stratigraphy is from the Paleobiology Database, while the measurements are from Isasmendi et al. (2020), except the Crown Height of PS.C-15,30 is changed from 43 mm to unknown due to its partial preservation.
References- Torcida, Fuentes, Izquierdo, Montero and Urien, 1997. Dientes de dinosaurios teropodos (cf. Baryonyx) en el Weald de Burgos (Espana). Studia Geologica Salamanticensia. 33, 59-65.
https://paleobiodb.org/classic/basicCollectionSearch?collection_no=89274
Isasmendi, Sáez-Benito, Torices, Navarro-Lorbés and Pereda-Suberbiola, 2020. New insights about theropod palaeobiodiversity in the Iberian Peninsula and Europe: Spinosaurid teeth (Theropoda, Megalosauroidea) from the Lower Cretaceous of La Rioja (Spain). Cretaceous Research. 104600.

Baryonychinae indet. (Torcida, Fuentes, Izquierdo, Montero and Urien, 1997)
Late Hauterivian, Early Cretaceous
El Juguete, Pinilla de los Moros Formation, Urbión Group, Salas de los Infantes, Castile and Leon, Spain
Material- (J.G. 1) (subadult) chevron (160 mm) (Fuentes Vidarte, Meijide Calvo, Izquierdo, Molinero, Montero, Pérez, Urien, Meijide Fuentes and Meijide Fuentes, 1999)
....(J.G. 2) proximal cervical rib
....(J.G. 3) proximal cervical rib
....(J.G. 4) proximal cervical rib
....(J.G. 5) squamosal
....(J.G. 6) postorbital
....(J.G. 7) caudal centrum (90.5 mm)
....(J.G. 8) posterior sacral fragment (fifth centrum 71.5 mm)
....(J.G. 9) cervical neural arch
....(J.G. 10) metacarpal I (65 mm)
....(J.G. 12) proximal metacarpal II
....(J.G. 18) proximal metacarpal III
....(J.G. 21) manual phalanx I-1 (130 mm)
....(J.G. 284) tooth fragment
....(J.G. coll.) 36 fragments
Late Hauterivian, Early Cretaceous
La Tejera-Valdesancho, Pinilla de los Moros Formation, Urbión Group, Barbadillo del Mercado, Castile and Leon, Spain
(PS.TBMV,13) incomplete tooth (?x14x10.32 mm) (Torcida, Fuentes, Izquierdo, Montero and Urien, 1997)
Late Hauterivian, Early Cretaceous
Tenada de Costalomo, Pinilla de los Moros Formation, Urbión Group, Salas de los Infantes, Castile and Leon, Spain
(PS.CLST,2) tooth (26.08x12.96x9.92 mm) (Torcida, Fuentes, Izquierdo, Montero and Urien, 1997)
Comments- The stratigraphy of PS.CLST,2 and PS.TBMV,13 is from the Paleobiology Database, while the measurements are from Isasmendi et al. (2020) except the Crown Height of the latter is changed from 30.04 mm to unknown due to its partial preservation.
Isasmendi et al. (2024 online) state "this specimen does not belong to the genus Baryonyx and, possibly, not even to Spinosauridae, owing to the combination of the presence of a ridge on the anterior surface of the shaft of the chevron, instead of a groove, and the absence of paired anterior processes on the proximal end of the chevron. The described chevron (MDS-Salas de los Infantes JBS,1) lacks an anterior longitudinal groove, which is present in Baryonyx, Camarillasaurus, SM-KK14, and FSAC-KK 11888. In MDS-Salas de los Infantes JBS,1, instead, the anterior surface is convex, with a ridge appearing distally, similar to the chevrons assigned to Vallibonavenatrix. Furthermore, MDS-Salas de los Infantes JBS,1 bears anterior processes, a feature apparently not present in Spinosauridae. It has been reported that these processes are not present in the preserved haemal arches of Baryonyx, Riparovenator, Suchomimus, SM-KK14, and apparently, in Vallibonavenatrix. In Camarillasaurus (MPG-KPC45, 60, and 63), most of the preserved chevrons lack the anterior process, but the left anterior process is present in MPG-KPC47. Likewise, some of the Baryonyx and FSAC KK 11888 (O.W.M. Rauhut, pers. comm.) chevrons seem to show anterior processes, but they are not as developed as in the El Juguete theropod."
References- Torcida, Fuentes, Izquierdo, Montero and Urien, 1997. Dientes de dinosaurios teropodos (cf. Baryonyx) en el Weald de Burgos (Espana). Studia Geologica Salamanticensia. 33, 59-65.
Fuentes Vidarte, Meijide Calvo, Izquierdo, Molinero, Montero, Pérez, Urien, Meijide Fuentes and Meijide Fuentes, 1999. Restos de Baryonyx (Dinosauria, Theropoda) en el weald de Salas de los Infantes (Burgos, España). I Jornadas Internacionales sobre Paleontología de dinosaurios y su entorno. 25-26.
Fuentes Vidarte, Meijide Calvo, Izquierdo, Montero, Pérex, Torcida, Urién, Meijide Fuentes and Meijide Fuentes, 2001. Restos fósiles de Baryonyx (Dinosauria, Theropoda) en el Cretácico inferior de Salas de los Infantes (Burgos, España). Actas de las I Jornadas internacionales sobre Paleontología de Dinosaurios y su entorno. 349-359.
https://paleobiodb.org/classic/basicCollectionSearch?collection_no=68464
https://paleobiodb.org/classic/basicCollectionSearch?collection_no=89271
Isasmendi, Sáez-Benito, Torices, Navarro-Lorbés and Pereda-Suberbiola, 2020. New insights about theropod palaeobiodiversity in the Iberian Peninsula and Europe: Spinosaurid teeth (Theropoda, Megalosauroidea) from the Lower Cretaceous of La Rioja (Spain). Cretaceous Research. 104600.
Isasmendi, Cuesta, Díaz-Martínez, Company, Sáez-Benito, Viera, Torices and Pereda-Suberbiola, 2024 online. Increasing the theropod record of Europe: A new basal spinosaurid from the Enciso Group of the Cameros Basin (La Rioja, Spain). Evolutionary implications and palaeobiodiversity. Zoological Journal of the Linnean Society. Advance article. DOI: 10.1093/zoolinnean/zlad193

Baryonychinae indet. (Falset, Santos-Cubedo, Figols and Fuster, 2009)
Late Hauterivian-Early Barremian, Early Cretaceous
Cantaperdius Formation, Spain
Material- (MG98-I-I) tooth (~35x14x12 mm)
Reference- Falset, Santos-Cubedo, Figols and Fuster, 2009. Primera cita de Baryonychine (Theropoda: Spinosauridae) en el Hauterviense terminal-Barremiense basal (Formación Cantaperdius) de Castellón (España). Journal of Paleontology and Paleoecology, Paleolusitana. 1, 391-396.

Baryonychinae indet. (Sánchez-Hernández, Benton and Naish, 2007)
Early Barremian, Early Cretaceous
El Castellar Formation, Spain
Material- (BNA-2/7) tooth (Gasca et al., 2018)
(CSC1-4) incomplete manual ungual I (~220 mm) (Gasca et al., 2018)
(MOAL-1/1) tooth (22.7x14x10.5 mm) (Gasca et al., 2008)
(MPG PX-23) tooth (1.3x.7x.4 mm) (Sanchez-Hernandez et al., 2007)
(SM-2/D1) tooth (11.6x9x6.8 mm) (Gasca et al., 2008)
(SUE1-2) mid caudal centrum (Gasca et al., 2018)
(VES4-1) mid caudal centrum (Gasca et al., 2018)
Comments- Manual ungual I CSC1-4 was discovered in 2010 and described as Aff. Baryonyx sp. by Gasca et al. (2018). Gasca et al. state it "differs from that of Baryonyx walkeri in having a little less curvature, a straight dorsal edge in the proximal part, slightly more width above the grooves than below - comparing the section at mid-length -, and a certain asymmetry, with the lateral face more flattened."
References- Sánchez-Hernández, Benton and Naish, 2007. Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain. Palaeogeography, Palaeoclimate, Palaeoecology. 249, 180-215.
Gasca, Moreno-Azanza and Canudo, 2008. Dientes de dinosaurios terópodos espinosáuridos de la Formación El Castellar (Cretácico Inferior, Teruel). Palaeontologica Nova. 8, 233-234.
Gasca, Díaz-Martínez, Moreno-Azanza, Canudo and Alonso, 2018. A hypertrophied ungual phalanx from the lower Barremian of Spain: Implications for the diversity and paleoecology of Spinosauridae (Theropoda) in Iberia. Cretaceous Research. 84, 141-152.

Baryonychinae indet. (Sánchez-Hernández, Benton and Naish, 2007)
Early Barremian, Early Cretaceous
Camarillas Formation, Spain
Material- (MPG CR(m)-1) tooth (17x7x4 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG CR(m)-2) tooth (12x6x4 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG CR(m)-3) tooth (18x6x6 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG CR(m)-4) tooth (17x9x7 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG POCA-6) tooth (3.9x1.9x.9 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG POCA-7) tooth (3.1x1.5x? mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG POCA-14) tooth (3x1.8x1.5 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG POCA-15) tooth (2.8x1.5x.8 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG POCA-18) tooth (2x1x.5 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG SC-1) tooth (16x9x8 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPG SC-2) tooth (21x12x7 mm) (Sánchez-Hernández, Benton and Naish, 2007)
(MPZ 2014/444) tooth (18.76x7.57x5.36 mm) (Alonso and Canudo, 2016)
References- Sánchez-Hernández, Benton and Naish, 2007. Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain. Palaeogeography, Palaeoclimate, Palaeoecology. 249, 180-215.
Alonso and Canudo, 2016. On the spinosaurid theropod teeth from the Early Barremian (Early Cretaceous) Blesa Formation (Spain). Historical Biology. 28(6), 823-834.

Baryonychinae indet. (Infante, Canudo and Ruiz-Omeñaca, 2005)
Early Barremian, Early Cretaceous
Mirambel Formation, Spain
Material- (LAD0-2) tooth fragment (?x6.7x5.3 mm)
Comments- Note Isasmendi et al. (2020) incorrectly list this tooth's measurements as 22x9.6x4.2 mm. This tooth preserves both labial and lingual fluting and 4.9 serrations per mm.
References- Infante, Canudo and Ruiz-Omeñaca, 2005. Primera evidencia de dinosaurios terópodos en la Formación Mirambel (Barremiense inferior, Cretácico Inferior) en Castellote, Teruel. Geogaceta. 38, 31-34.
Isasmendi, Sáez-Benito, Torices, Navarro-Lorbés and Pereda-Suberbiola, 2020. New insights about theropod palaeobiodiversity in the Iberian Peninsula and Europe: Spinosaurid teeth (Theropoda, Megalosauroidea) from the Lower Cretaceous of La Rioja (Spain). Cretaceous Research. 104600.

Baryonychinae indet. (Ruiz-Omeñaca, Canudo, Cruzado-Caballero, Infante and Moreno-Azanza, 2005)
Barremian, Early Cretaceous
Blesa Formation, Spain
Material- (CAN1 953) tooth (22.39x8.70x5.96 mm) (Alonso and Canudo, 2016)
(CAN1 957) tooth (17.16x7.92x5.15 mm) (Alonso and Canudo, 2016)
(CAN1 958) tooth (20.75x7.69x5.42 mm) (Alonso and Canudo, 2016)
(CAN1 959) tooth (9.14x4.32x3.66 mm) (Alonso and Canudo, 2016)
(CAN1 977) tooth (Alonso and Canudo, 2016)
(CAN1 983) tooth (6.96x3.03x2.34 mm) (Alonso and Canudo, 2016)
(CAN1 984) tooth (8.91x4.03x3.10 mm) (Alonso and Canudo, 2016)
(CAN1 999) tooth (11.40x4.25x3.85 mm) (Alonso and Canudo, 2016)
(CAN1 1001) tooth (9.58x3.72x2.53 mm) (Alonso and Canudo, 2016)
(CAN1 1616) tooth (16.68x7.09x6.05 mm) (Alonso and Canudo, 2016)
(MPZ 97/468) tooth fragment (14.77x5.45x3.28 mm) (Ruiz-Omeñaca et al., 1997)
(MPZ 2001/207) tooth (28.36x13.29x9.38 mm) (Canudo and Ruiz-Omeñaca, 2003)
(MPZ 2001/208) tooth (18.14x7.84x5.90 mm) (Canudo and Ruiz-Omeñaca, 2003)
(MPZ 2005/303) tooth fragment (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/304) tooth (FABL 8.7 mm) (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/305) tooth fragment (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/306) tooth (FABL 6.6 mm) (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/307) tooth (FABL 6.4 mm) (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/308) tooth fragment (FABL ~9 mm) (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/309) tooth (FABL 4.7 mm) (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/310) tooth (FABL 2.5 mm) (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/311) tooth (FABL 3 mm) (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/312) tooth fragment (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/313) tooth fragment (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/314) tooth fragment (Canudo and Ruiz-Omeñaca, 2005)
(MPZ 2005/315) tooth fragment (Canudo and Ruiz-Omeñaca, 2005)
(TPFM coll.) several teeth (Ruiz-Omeñaca et al., 2005)
Comments- MPZ 97/468 was originally described as Theropoda indet. B by Ruiz-Omeñaca et al. (1997), and later assigned to Baryonychidae indet. (Ruiz-Omeñaca et al., 1998). Canudo and Ruiz-Omeñaca (2003) illustrated MPZ 2001/207 and 2001/208 as Baryonychinae indet.. These three specimens and MPZ 2005/303-315 were described in detail by Ruiz-Omeñaca et al. (2005).
The teeth are generally similar to Baryonyx, but differ in being more labiolingually compressed, having labial fluting and a greater variation in serration density (6-13 per mm vs. 7 per mm).
References- Ruiz-Omeñaca, Canudo and Cuenca-Bescós, 1997. Primera evidencia de un área de alimentación de dinosaurios herbívoros en el Cretácico Inferior de España (Teruel). Monografías de la Academia de Ciencias de Zaragoza. 10, 1-48.
Ruiz-Omeñaca, Canudo and Cuenca-Bescós, 1997. Primera cita de dinosaurios barionícidos (Saurischia: Theropoda) en el Barremiense superior (Cretácico inferior) de Vallipón (Castellote, Teruel). Beca del Museo de Mas de las Matas. 17, 201-223.
Canudo and Ruiz-Omeñaca, 2003. Los restos directos de dinosaurios teropódos (excluyendo Aves) en España [Direct remains of theropod dinosaurs (excluding Aves) in Spain]. Ciencias de la Tierra. 26, 347-373.
Ruiz-Omeñaca, Canudo, Cruzado-Caballero, Infante and Moreno-Azanza, 2005. Baryonychine teeth (Theropoda: Spinosauridae) from the Lower Cretaceous of La Cantalera (Jose, NE Spain). Kaupia. 14, 59-63.
Alonso and Canudo, 2016. On the spinosaurid theropod teeth from the Early Barremian (Early Cretaceous) Blesa Formation (Spain). Historical Biology. 28(6), 823-834.
Alonso, Gasca, Navarro-Lorbés, Rubio and Canudo, 2018. A new contribution to our knowledge of the large-bodied theropods from the Barremian of the Iberian Peninsula: The "Barranco del Hocino" site (Spain). Journal of Iberian Geology. 44, 7-23.

unnamed possible Baryonychinae (Ruiz-Omeñaca, Canudo and Cuenca-Bescós, 1997)
Late Barremian, Early Cretaceous
Artoles Formation, Spain
Material- (MPZ 98/59) tooth (19.0x10.0x8.0 mm) (Ruiz-Omeñaca, Canudo and Cuenca-Bescós, 1997)
(MPZ 98/60) tooth (16.0x8.0x6.4 mm) (Ruiz-Omeñaca, Canudo and Cuenca-Bescós, 1997)
(MPZ 98/61) tooth (~17.0x~7.4x~6.3 mm) (Ruiz-Omeñaca, Canudo and Cuenca-Bescós, 1997)
(MPZ 2014/438) tooth (38.34x15.30x12.56 mm) (Alonso and Canudo, 2016)
(MPZ 2014/439) tooth (21.61x8.94x8.06 mm) (Alonso and Canudo, 2016)
(MPZ 2014/440) tooth (30.79x13.03x11.11 mm) (Alonso and Canudo, 2016)
(MPZ 2014/441) tooth (25.48x9.86x8.55 mm) (Alonso and Canudo, 2016)
(MPZ 2014/442) tooth (7.91x4.42x3.94 mm) (Alonso and Canudo, 2016)
Comments- These differ from Baryonyx in having both lingual and labial fluting, and in lacking mesial serrations. This opens the possibility of them being from a basal spinosaurine.
Reference- Ruiz-Omeñaca, Canudo and Cuenca-Bescós, 1997. Primera cita de dinosaurios barionícidos (Saurischia: Theropoda) en el Barremiense superior (Cretácico inferior) de Vallipón (Castellote, Teruel). Beca del Museo de Mas de las Matas. 17, 201-223.
Alonso and Canudo, 2016. On the spinosaurid theropod teeth from the Early Barremian (Early Cretaceous) Blesa Formation (Spain). Historical Biology. 28(6), 823-834.

Baryonychinae indet. (Royo and Gomez, 1927)
Barremian-Aptian, Early Cretaceous
Wealden Supergroup equivalent, Spain
Material- ?teeth (Royo and Gomez, 1927)
Comments- Buffetaut (2010) notes Royo and Gomez (1927) mentioned Suchosaurus teeth from the Wealden of Spain.
References- Royo and Gomez, 1927. Sur le facies wealdien d’Espagne. Comptes Rendus de la Societe geologique de France. 10, 125-128.
Buffetaut, 2010. Spinosaurs before Stromer: Early finds of spinosaurid dinosaurs and their interpretations. Geological Society, London, Special Publications. 343, 175-188.

Baryonychinae indet. (Torcida Fernández, Izquierdo Montero, Huerta Hurtado, Montero Huerta and Pérez Martínez, 2003)
Late Barremian-Early Aptian, Early Cretaceous
Castrillo de la Reina Formation, Spain
Material- (MDS-OTII,81) tooth (?x21.03x14.74 mm) (Alonso, Canudo, Fernández-Baldor and Huerta, 2017)
(MDS-OTII,84) tooth (?x5.21x3.55 mm) (Alonso, Canudo, Fernández-Baldor and Huerta, 2017)
(MDS-OTII,85) tooth (?x7.06x5.58 mm) (Alonso, Canudo, Fernández-Baldor and Huerta, 2017)
(MDS-OTII,86) tooth (?x10.5x8.72 mm) (Alonso, Canudo, Fernández-Baldor and Huerta, 2017)
(MDS-OTII,99) tooth (29.23x14.94x10.55 mm) (Alonso, Canudo, Fernández-Baldor and Huerta, 2017)
(PS-JTS 20) tooth (?x9x7 mm) (Torcida Fernández, Izquierdo Montero, Huerta Hurtado, Montero Huerta and Pérez Martínez, 2003)
(PS C-1 11) tooth (?x14x10 mm) (Torcida Fernández, Izquierdo Montero, Huerta Hurtado, Montero Huerta and Pérez Martínez, 2003)
Comments- These teeth are similar to Baryonyx except for having both labial and lingual fluting. PS-JTS 20 has 6 serrations per mm mesially (though lacking them basally), and 6-8 per mm distally. PS C-1 11 has 6 per mm mesially and 5-6 per mm distally.
References- Torcida Fernández, Izquierdo Montero, Huerta Hurtado, Montero Huerta and Pérez Martínez, 2003. Dientes de dinosaurios (Theropoda, Sauropoda), en el Cretácico Inferior de Burgos (España). In Pérez-Lorente (ed.). Dinosaurios y otros Reptiles Mesozoicos en España (IER, Ciencias de la Tierra, 26). 335-346.
Alonso, Canudo, Fernández-Baldor and Huerta, 2017. Isolated theropod teeth associated with sauropod remains from El Oterillo II (Early Cretaceous) site of Salas de los Infantes (Burgos, Spain). Journal of Iberian Geology. 43(2), 193-215.

undescribed baryonychine (Suteethorn, Buffetaut, Wongo, Suteethorn and Tong, 2018)
Late Barremian?, Early Cretaceous
Pra Prong, Sao Khua Formation?, Thailand
Material- teeth
Comments- In addition to fluted Siamosaurus teeth, Suteethorn et al. (2018) mention "teeth with no ribbing of the crown. The enamel shows the fine wrinkling seen on most spinosaurid teeth. The carinae are well-marked. The mesial carina seems to be unserrated. The distal carina bears very small irregular serrations which disappear easily when the tooth is worn."
Reference- Suteethorn, Buffetaut, Wongo, Suteethorn and Tong, 2018. Morphological diversity of spinosaurid teeth from the Pra Prong locality (Lower Cretaceous of eastern Thailand). 5th International Palaeontological Congress. Abstract Book, 666.

undescribed Baryonychinae (Sone, Hirayama, He, Yoshida and Komatsu, 2015)
Early Cretaceous
Tembeling Group, Malaysia
Material- (UM10575) tooth
(UM10576) tooth
Comments- Samathi et al. (2019) state "the teeth show sharp vertical ridges and serrated carinae with minute denticles and display a veined micro-ornament over the surface ... subcircular tooth crowns in cross-section and the presence of striations."
References- Sone, Hirayama, He, Yoshida and Komatsu, 2015. First dinosaur fossils from Malaysia: Spinosaurid and ornithischian teeth. 2nd International Symposium on Asian Dinosaurs. 18.
Samathi, Chanthasit and Sander, 2019. A review of theropod dinosaurs from the Late Jurassic to mid-Cretaceous of southeast Asia. Annales de Paléontologie. 105(3), 201-215.

undescribed Baryonychinae (Congleton, 1990)
Aptian, Early Cretaceous
Gres de Gaba Member of the Koum Formation, Cameroon
Material- (CAM 320) tooth
(CAM 322) tooth
(CAM 349) tooth
(CAM 350) tooth
(CAM 351) tooth
(CAM 352) tooth
(CAM 353) tooth
(CAM 354) tooth
(CAM 355) tooth
(CAM 356) tooth
(CAM 357) tooth
(CAM 358) tooth
(CAM 360) tooth
(SMU 72033) tooth
Comments- These were assigned to cf. Spinosauridae by Congleton (1990) and to Baryonychinae by Bertin (2010) due to their serrations.
References- Congleton, 1990. Vertebrate paleontology of the Koum Basin, northern Cameroon, and archosaurian paleobiogeography in the Early Cretaceous. MS thesis. Southern Methodist University. 245 pp.
Bertin, 2010. A catalogue of the material and review of the Spinosauridae. PalArch’s Journal of Vertebrate Palaeontology. 7(4), 1-39.

unnamed Baryonychinae (Le Loeuff, Métais, Dutheil, Rubinos, Buffetaut, Ois Lafont, Cavin, Moreau, Tong, Blanpied and Sbeta, 2010)
Hauterivian-Barremian, Early Cretaceous
Jawsh quarry, Cabao Formation, Libya
Material- (UT-JAW2) tooth (27x~12x~9 mm)
(UT-JAW coll.) tooth
Comments- Le Loeuff et al. (2010) state UT-JAW2 is "slightly compressed labio-lingually with a granular enamel. The tooth is fluted and its mesial and distal carinae show extremely small serrations (7-8/mm)." They refer it to Baryonychinae as cf. Baryonyx, though they note "they seem to be broader in relation to the crown height than is typical for Baryonyx."
Reference- Le Loeuff, Métais, Dutheil, Rubinos, Buffetaut, Ois Lafont, Cavin, Moreau, Tong, Blanpied and Sbeta, 2010. An Early Cretaceous vertebrate assemblage from the Cabao Formation of NW Libya. Geological Magazine. 147(5), 750-759.

Ceratosuchopsini Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021
Definition- (Ceratosuchops inferodios <- Baryonyx walkeri, Spinosaurus aegyptiacus) (Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021)
Comments- This tribe was established by Barker et al. (2021) based on Cau's megamatrix, and included Ceratosuchops, Riparovenator and Suchomimus, but not Baryonyx or Vallibonavenatrix among baryonychines.
Reference- Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports. 11:19340.

Cristatusaurus Taquet and Russell, 1998
= Suchomimus Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998
C. lapparenti Taquet and Russell, 1998
= Suchomimus tenerensis Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998
= Baryonyx tenerensis (Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998) Sues, Frey, Martill and Scott, 2002
Aptian, Early Cretaceous
Tegama Bed of the Elrhaz Formation, Niger
Holotype- (MNN GDF 366) (subadult) premaxillae, partial maxilla, partial dentary
Paratypes- (MNN GDF 357, 358, 361) dorsal vertebrae (135 mm)
(MNN GDF 359) posterior (11th?) dorsal neural arch (centrum ~100 mm)
(MNN GDF 365) (adult) premaxilla
Referred- (MNN GAD 70) specimen including cervical vertebrae, sacral vertebrae, caudal vertebrae (Samathi et al., 2021 online)
(MNN GAD 72) (juvenile) tibia (700 mm) (Samathi, Chanthasit and Sander, 2019)
(MNN GAD 87) specimen including caudal vertebrae (Samathi et al., 2021 online)
(MNN GAD 94-52) specimen including chevron (Samathi et al., 2021 online)
(MNN GAD 97) (subadult) tibia (760 mm), astragalus (Samathi, Chanthasit and Sander, 2019)
(MNN GAD 98) astragalus (Samathi, Chanthasit and Sander, 2019)
?(MNN GAD 151) tooth (53x22x15 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 154) tooth (50x19x14 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 161) tooth (68x20x13 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 166) tooth (54x18x14 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 210) tooth (25x14x8 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 374) tooth (65x21x13 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 504) tooth (63x18x13 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 507) tooth (54x21x16 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNN GAD 513) cervical vertebrae 2-10, dorsal vertebrae 1-5, ribs, gastralia, girdle and limb elements, manual and pedal elements, furcula (Lipkin et al., 2007)
?(MNN GAD 518) tooth (45x14x10 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 520) tooth (58x22x15 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 544) tooth (34x12x10 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 570) tooth (23x14x7 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 582) tooth (8x5x3 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 625) tooth (43x20x10 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 1966) tooth (55x17x12 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 1966 a) tooth (47x18x15 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 1966 b) tooth (52x17x17 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
?(MNN GAD 1966 c) tooth (63x19x15 mm) (Fanti, Cau, Martinelli and Contessi, 2014)
(MNN GDF 214) nasal, prefrontal, frontal, braincase (Ibrahim and Sereno, 2011)
(MNN GDF 500; holotype of Suchomimus tenerensis) (11.0 m, 2.9-4.8 tons) third cervical rib, fifth cervical rib, eighth cervical rib, first dorsal vertebra, second dorsal vertebra, third dorsal vertebra, fifth dorsal vertebra, sixth dorsal vertebra, seventh dorsal vertebra, eighth dorsal vertebra, ninth dorsal vertebra, tenth dorsal neural spine, eleventh dorsal centrum, thirteenth dorsal vertebra, fourteenth dorsal vertebra, fifteenth dorsal vertebra, sixteenth dorsal vertebra, ten dorsal ribs, gastralia, sacral centra, sacral neural spines 3-5, caudal transverse processes 2-5, caudal neural spines 1-5, six mid caudal centra, distal caudal vertebra, three chevrons, scapula, coracoid, humerus (560 mm), radius (255 mm), ulna, manual ungual I (264 mm), manual ungual II (165 perp. to art,), metacarpal III (130 mm), manual ungual III (120 perp. to art.), ilium (1 m), pubis, ischium, femur (1.075 m), tibia (960 mm), metatarsal III (380 mm), pedal phalanx (Sereno et al., 1998)
(MNN GDF 501; paratype of Suchomimus tenerensis) premaxillae, maxillae (Sereno et al., 1998)
(MNN GDF 502; paratype of Suchomimus tenerensis) quadrate (Sereno et al., 1998)
(MNN GDF 503; paratype of Suchomimus tenerensis) partial dentary (Sereno et al., 1998)
(MNN GDF 504; paratype of Suchomimus tenerensis) partial dentary (Sereno et al., 1998)
(MNN GDF 505; paratype of Suchomimus tenerensis) partial dentary (Sereno et al., 1998)
(MNN GDF 506; paratype of Suchomimus tenerensis) axis (Sereno et al., 1998)
(MNN GDF 507; paratype of Suchomimus tenerensis) posterior cervical vertebra (Sereno et al., 1998)
(MNN GDF 508; paratype of Suchomimus tenerensis) posterior dorsal vertebra (Sereno et al., 1998)
(MNN GDF 510; paratype of Suchomimus tenerensis) caudal vertebra (Sereno et al., 1998)
(MNN GDF 511; paratype of Suchomimus tenerensis) caudal vertebra (Sereno et al., 1998)
?(MNN GDF coll.) dorsal neural arch (Taquet, 1976)
(MNN GDF coll.) teeth, bones (Sereno et al., 1998)
(MNN GDF coll.) snout (Ibrahim and Sereno, 2011)
Diagnosis- (after Taquet and Russell, 1988) sagittal crest on premaxilla.
(modified after Sereno et al., 1998) elongate subnarial process that nearly excludes maxilla from external naris (unknown in other baryonychines); broadened posterior dorsal, sacral and proximal caudal neural spines; robust humeral tuberosities; hook-shaped radial entepicondyle.
Other diagnoses- Sereno et al. (1998) listed 'hypertrophied ulnar olecranon process that is offset from the humeral articulation' as a diagnostic character, but Carrano et al. (2012) note it is present in Baryonyx too. They also listed heightened posterior dorsal, sacral and proximal caudal neural spines, but Riparovenator has tall caudal neural spines as well, and a Wessex dorsal possibly belonging to it or Ceratosuchops has a heightened dorsal neural spine.
Comments- Contra Sereno et al. (1998), Charig and Milner (1997) and Carrano et al. (2012), Cristatusaurus is not a nomen dubium. Taquet and Russell (1998) are correct in noting the "brevirostrine" condition distinguishes Cristatusaurus from Baryonyx, except that they define this as having a sagittal crest (pg. 351) as opposed to the more obvious interpretation of having a short snout. A Suchomimus paratype (MNN GDF 501) has a sagittal crest too. This may be due to ontogeny, but the same can be said of the characters Sereno et al. proposed to distinguish Suchomimus from Baryonyx, and Ceratosuchops and Riparovenator are now known to lack crests as well. In addition, Cristatusaurus and Suchomimus resemble each other more than Baryonyx in having a less rounded anterodorsal margin to their premaxillae (also in Ceratosuchops but not Riparovenator), and having a comparatively larger second alveolus (though it is smaller in Suchomimus than Cristatusaurus; also in Riparovenator but not Ceratosuchops). Since both are from the same formation, share an apparently apomorphy, and have no obvious differences besides alveolar proportions, they are synonymized here.
Several authors have suggested synonymizing Cristatusaurus/Suchomimus with Baryonyx (Charig and Milner, 1997; Milner pers. comm. 1999 to Naish et al., 2001; Sues et al., 2002). While Charig and Milner's (1997) decision to sink the Cristatusaurus material into Baryonyx sp. indet. made sense prior to the naming of Cristatusaurus or Suchomimus, one cannot now sink a named species like lapparenti into "sp. indet.", nor can one sink Cristatusaurus into Baryonyx without placing Suchomimus there too (as nothing suggests lapparenti is closer to walkeri than to tenerensis; quite the opposite, as seen above). Thus Buffetaut and Ouaja (2002) were wrong to formally sink Cristatusaurus into Baryonyx sp. but leave Suchomimus as provisionally valid. Similarly, even though Sues et al. state "There exists at present no evidence to indicate the presence of more than one taxon of spinosaurid in the faunal assemblage from GAD 5", they incorrectly refer to this taxon as Baryonyx tenerensis instead of B. lapparenti (which would have priority if lapparenti and tenerensis are one taxon). The combination Baryonyx lapparenti has yet to appear in the literature, however. Ultimately, this is a subjective decision since there is no objective definition of 'genus', and the more recent naming of Vallibonavenatrix, Iberospinus, Protathlitis and especially Ceratosuchops and Riparovenator shows the community accepts baryonychine species as non-congeneric. The latter two taxa were recovered closer to Cristatusaurus (as Suchomimus) by their describers, so would also be species of Baryonyx if lapparenti was accepted as such. Finally, sinking all baryonychines into Baryonyx would also imply Suchosaurus was congeneric as well, creating a taxonomic issue since it has 145 years of priority over Baryonyx.
Taquet (1976) describes a dorsal neural arch as identical to Becklespinax and very similar to Spinosaurus, so this may belong to Cristatusaurus. Ibrahim and Sereno (2011) remark on "complete snout and braincase material" of Suchomimus, the latter no doubt MNN GDF 214 as mentioned by Paulina-Carabajal et al. (2019) and Barker et al. (2021). Fanti et al. (2014) list many "Baryonychinae indet." teeth from Gadoufoua, here provisionally referred to Cristatusaurus.
Samathi et al. (2019) measured the holotype tibia as 960 mm instead of Sereno et al.'s 945 mm, and provides some additional details.
References- Taquet, 1984. Une curieuse spécialisation du crane de certains Dinosaures carnivores du Crétacé: le museau long et etroit des Spinosauridés. Comptes Rendus des Seances de l'Academie des Sciences, Série 2. 299(5), 217-222.
Charig and Milner, 1997. Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum of London (Geology). 53, 11-70.
Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998. A long-snouted predatory dinosaur from Africa and the evolution of the spinosaurids. Science. 282(5392), 1298-1302.
Taquet and Russell, 1998. New data on spinosaurid dinosaurs from the Early Cretaceous of the Sahara. Comptes Rendus de l'Académie des Sciences à Paris, Sciences de la terre et des planetes. 327, 347-353.
Naish, Hutt and Martill, 2001. Saurischian dinosaurs 2: Theropods. In Martill and Naish (eds.). Dinosaurs of the Isle of Wight. The Palaeontological Association, London. 242-309.
Buffetaut and Ouaja, 2002. A new specimen of Spinosaurus (Dinosauria, Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the evolutionary history of the Spinosauridae. Bulletin de la Societe Geologique de France. 173(5), 415-421.
Sues, Frey, Martill and Scott, 2002. Irritator challengeri, a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil. Journal of Vertebrate Paleontology. 22(3), 535-547.
Hutt and Newbery, 2004. A new look at Baryonyx walkeri (Charig and Milner, 1986) based upon a recent fossil find from the Wealden. SVPCA 2004. 18.
Lipkin, Sereno and Horner, 2007. The furcula in Suchomimus tenerensis and Tyrannosaurus rex (Dinosauria: Theropoda: Tetanurae). Journal of Paleontology. 81(6), 1523-1527.
Ibrahim and Sereno, 2011. New data on spinosaurids (Dinosauria: Theropoda) from Africa. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 130.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.
Fanti, Cau, Martinelli and Contessi, 2014. Integrating palaeoecology and morphology in theropod diversity estimation: A case from the Aptian-Albian of Tunisia. Palaeogeography, Palaeoclimatology, Palaeoecology. 410, 39-57.
Paulina-Carabajal, Ezcurra and Novas, 2019. New information on the braincase and endocranial morphology of the Late Triassic neotheropod Zupaysaurus rougieri using Computed Tomography data. Journal of Vertebrate Paleontology. e1630421.
Samathi, Chanthasit and Sander, 2019. Two new basal coelurosaurian theropod dinosaurs from the Lower Cretaceous Sao Khua Formation of Thailand. Acta Palaeontologica Polonica. 64(2), 239-260.
Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports. 11:19340.
Samathi, Sander and Chanthasit, 2021 online. A spinosaurid from Thailand (Sao Khua Formation, Early Cretaceous) and a reassessment of Camarillasaurus cirugedae from the Early Cretaceous of Spain. Historical Biology. Latest Articles. DOI: 10.1080/08912963.2021.1874372

Ceratosuchops Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021
C. inferodios Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021
Barremian, Early Cretaceous
Wessex Formation of the Wealden Group, England
Holotype- (IWCMS 2014.95.1) prefrontal, frontals, parietal, orbitosphenoids, laterosphenoids
....(IWCMS 2014.95.2) otoccipital, supraoccipital
....(IWCMS 2014.95.3) basisphenoid, basioccipital, prootics, otoccipital
....(IWCMS 2014.95.5) anterior premaxillae
....(IWCMS 2021.30) posterior premaxillae
Paratype- ....(IWCMS 2014.95.4) postorbital
Diagnosis- (after Barker et al., 2021) premaxilla with tuberosity at anterior edge of external naris; narrow subcondylar recess of basioccipital; subcondylar recess ventrally separated from occipital condyle; deep, elongate sulci medial to oval scars on basisphenoid; anteroposteriorly thick interbasipterygoidal web; supraoccipital dorsal process possessing a gently curving posteroventral surface in coronal section.
Comments- Discovered between 2014 and 2021, the premaxillae and braincase are smaller than Riparovenator, while the isolated postorbital was referred "as the dimensions of the articular facets of the postorbital, while damaged, match those of the respective frontal and laterosphenoid articulations." Barker et al. (2021) used Cau's megamatrix to recover it sister to Riparovenator in their new Ceratosuchopsini within Baryonychinae.
Reference- Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports. 11:19340.

Riparovenator Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021
R. milnerae Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021
Barremian, Early Cretaceous
Wessex Formation of the Wealden Group, England
Holotype- (IWCMS 2014.95.6) anterior premaxillae
....(IWCMS 2014.96.1) partial prefrontal, frontals, partial parietal, orbitosphenoid, laterosphenoid
....(IWCMS 2014.96.2) partial orbitosphenoid, laterosphenoid
....(IWCMS 2014.96.3) lacrimal fragment, prefrontal
....(IWCMS 2020.448.1) posterior braincase
....(IWCMS 2020.448.2) cultriform process of basisphenoid
Paratypes- ....(IWCMS 2014.95.7) posterior nasal
....(IWCMS 2020.447.1-39) at least twenty-two fragmentary to complete mid to distal caudal vertebrae, eighteen partial to complete mid to distal chevrons
Diagnosis- (after Barker et al., 2021) deeply notched dorsal orbital margin between prefrontal and postorbital process of frontal; deeply inset facial nerve (VII) foramen that is largely obscured from lateral view; deep subcondylar recess (depth over 1/3 of its mediolateral width; depth less than 1/5 in other baryonychines); reduced exposure of articular surface of the basipterygoid process in lateral view.
Comments- Discovered between 2014 and 2020, the premaxillae and braincase are larger than Ceratosuchops while the isolated nasal fragment was referred "due to its size (the ventral frontal facet appears too large for the nasal process of the Ceratosuchops frontal) and similar state of preservation" The caudal elements were referred "due to their close proximity (c. 10m) to the holotype basicranial complex." Barker et al. (2021) used Cau's megamatrix to recover it sister to Ceratosuchops in their new Ceratosuchopsini within Baryonychinae.
Reference- Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports. 11:19340.

Spinosaurinae Stromer, 1915 vide Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998
Definition- (Spinosaurus aegyptiacus <- Baryonyx walkeri) (Holtz et al., 2004; modified from Sereno et al., 1998)
Diagnosis- (after Sues et al., 2002) tooth crowns with distinct but non-serrated carinae; fluted enamel on both the labial and lingual surfaces.
References- Stromer, 1915. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîje-Stufe (unterstes Cenoman). 3. Das Original des Theropoden Spinosaurus aegyptiacus nov. gen., nov. spec. Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften Mathematisch-physikalische Klasse Abhandlung. 28(3), 1-31.
Sereno, Beck, Dutheil, Gado, Larsson, Lyon, Marcot, Rauhut, Sadleir, Sidor, Varricchio, Wilson and Wilson, 1998. A long-snouted predatory dinosaur from Africa and the evolution of the spinosaurids. Science. 282(5392), 1298-1302.
Sues, Frey, Martill and Scott, 2002. Irritator challengeri, a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil. Journal of Vertebrate Paleontology. 22(3), 535-547.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria Second Edition. University of California Press. 71-110.

Camarillasaurus Sanchez-Hernandez and Benton, 2014
= "Camarillasaurus" Sanchez-Hernandez and Benton, online 2012
C. cirugedae Sanchez-Hernandez and Benton, 2014
= "Camarillasaurus cirugedae" Sanchez-Hernandez and Benton, online 2012
Early Barremian, Early Cretaceous
Camarillas Formation, Aragon, Spain
Holotype- (MPG-KPC1) scapular fragment
(MPG-KPC2) scapular fragment
(MPG-KPC3, 4) partial third to fifth synsacral centra
(MPG-KPC5) incomplete proximal chevron
(MPG-KPC6) distal anterior dorsal ?rib
(MPG-KPC7) incomplete first or second dorsal rib
(MPG-KPC8) proximal tibia
(MPG-KPC9) partial posterior dorsal vertebra
(MPG-KPC10) distal caudal vertebra (~80 mm)
(MPG-KPC11) distal caudal vertebra (~80 mm)
(MPG-KPC12) incomplete mid caudal vertebra
(MPG-KPC13) mid caudal vertebra (~80 mm)
(MPG-KPC14) partial proximal caudal centrum
(MPG-KPC15) partial proximal caudal centrum
(MPG-KPC16) partial ?first sacral centrum
(MPG-KPC17) partial proximal caudal centrum
(MPG-KPC18) partial ?sacral central fragment
(MPG-KPC19) partial proximal caudal centrum
(MPG-KPC20) partial proximal caudal centrum
(MPG-KPC21) partial proximal caudal vertebra (115 mm)
(MPG-KPC22) proximal caudal central fragment
(MPG-KPC23) 'coracoid'
(MPG-KPC25-29) fragments
(MPG-KPC30) scapular fragment
(MPG-KPC31) ?caudal neural spine fragment
(MPG-KPC32) ?caudal neural spine fragment
(MPG-KPC33) ?caudal neural spine fragment
(MPG-KPC34) ?scapular fragment
(MPG-KPC35) ?scapular fragment
(MPG-KPC36) ?scapular fragment
(MPG-KPC37) ?scapular fragment
(MPG-KPC38) ?scapular fragment
(MPG-KPC39) incomplete caudal neural arch
(MPG-KPC40-42) fragments
?(MPG-KPC43) partial lateral tooth
(MPG-KPC44) partial proximal chevron
(MPG-KPC45) dorsal rib fragment
(MPG-KPC46) partial proximal caudal centrum
(MPG-KPC47) dorsal rib fragment
(MPG-KPC48) dorsal rib fragment
(MPG-KPC49) dorsal rib fragment
(MPG-KPC50) dorsal rib fragment
(MPG-KPC51) anterior-mid ?cervical neural arch fragment
Referred- .... proximal caudal vertebra, femur, pedal ungual (Rauhut, Canudo and Castanera, 2019)
Diagnosis- (after Sanchez-Hernandez and Benton, 2014) caudal vertebrae with articular surfaces that have well developed edges and are unusually broad; extremely long tibia proximal end, with ratio of anteroposterior/mediolateral axis of 2.8.
Other diagnoses- While Sanchez-Hernandez and Benton (2014) included 'chevron with a deep broad longitudinal groove along the length of the shaft arising from a fossa placed below the haemal canal on the anterior and posterior side' in the diagnosis, Samathi et al. (2021) noted it "is present in other spinosaurids such as Suchomimus (MNBH GAD G94-52, A.S. pers. obs.) and the Khok Kruat spinosaurid (SM-KK14, A.S. pers. obs.)." Similarly, the original authors propose 'tibia with a narrow and deep longitudinal groove placed anterior to the crista fibularis on the medial [actually lateral] surface', but Samathi et al. state this "is also found in the Spinosaurus neotype (FSAC-KK 11,888, A.S. pers. obs.), although this feature" ... "might be present due to pathology or crushing because it is present on only one of the two preserved tibiae" and is also "found in one of the tibiae of Suchomimus (MNBH GAD 500, A.S. pers. obs.)." Finally they proposed 'articular surface on the distal end of the chevron blade' based on supposed ventral chevron tip MPG-KPC6, but as theropod chevrons don't articulate ventrally with anything, this fragment is more likely to be a distal anterior dorsal rib (articulating with sternal ribs) if it belongs to the specimen.
Comments- Sanchez-Hernandez and Benton (2014) described this as a basal ceratosaur more derived than Limusaurus, but less than Elaphrosaurus, MNN TIG6 and neoceratosaurs using Carrano and Sampson's ceratosaur matrix. Note the holotype is only listed as MPG-KPC1-46, but numbers through 51 are then described in the detailed list, except vertebra MPG-KPC24 and 25-29 and 40-42 that are presumably the "several isolated broken fragments of bones." Rauhut et al. (2019) reported "Further excavation at the type locality yielded some additional elements, presumably of the type individual, including an anterior caudal vertebra, a complete femur, and a pedal ungual."
Wang et al. (2016) noted "A fragmentary vertebra tentatively identified as a cervical [MPG-KPC24] most likely represents an anterior dorsal based on the position of the parapophyses and presence of a hypapophysis" and that "the morphology of the hypapophysis in Camarillasaurus more strongly resembles those found in some coelurosaurs (e.g. Nomingia, Avimimus, Mononykus) than the structure on dorsal 1 of Elaphrosaurus." Samathi (2019) found "It is too small to belong to the same individual or even the same taxon as Camarillasaurus", and Samathi et al. (2021) excluded the element from the type material. It is listed here under Maniraptora incertae sedis.
Rauhut et al. suggested "A supposed cervical vertebra is a posterior dorsal vertebra", probably referring to MPG-KPC9 considered "most likely a cervical" by Sanchez-Hernandez and Benson. The offset central faces are thus due to deformation, and the absent pleurocoel and parapophysis not located on the centrum would support Rauhut et al.'s identification.
Rauhut et al. also suggested "several supposed dorsal vertebrae are caudal vertebrae", which would seem likely for MPG-KPC17, MPG-KPC20 and MPG-KPC21 at least, which have the transversely compressed centra with ventral grooves typical of proximal caudal vertebrae. MPG-KPC51 was identified as "a posterior cervical vertebra or an anterior dorsal" by Sanchez-Herndandez and Benton, but may be a left fragment of an anterior-mid cervical upside down, with the labeled 'postzygapophysis' being an epipophysis and the horizontal ridge being the postzygapophysodiapophyseal lamina. MPG-KPC39 is described as "a broken dorsal neural arch" but the supposed prezygapophyses are clearly postzygapophyses because their articular surfaces angle ventrally, and the element is a caudal based on the nearly vertical zygapophyseal surfaces and unexpanded hypantrum. The triangular and rectangular fossae indicated in figure 5B2 are probably simply erosional. MPG-KPC31-33 "may be broken tops of neural spines", but are too short to be from dorsal vertebrae and may be caudal neural spine tips (or transverse processes) instead.
Wang et al. also correctly noted "Though six sacral vertebrae were inferred in Camarillasaurus, it should be noted that the authors only identified the remains of five vertebrae as sacrals." Although Sanchez-Hernandez and Benton identified the fused sacrals as 2-4, Malafaia et al. (2020) figured them as 3-5 based on comparison to Vallibonavenatrix.
Rauhut et al. note "the 'sternal plates' are parts of the scapulae", referring to MPG-KPC1-2, and the same may be true of MPG-KPC34-38 that are listed as "isolated remains of sternal plates." The supposed coracoid strongly differs from ceratosaurian or megalosauroid theropods in being anterodistally expanded as in many paravians, having a thick lip on the lateral side of the posterior edge, and having a ventrally located foramen that is most likely damage. The element is near certainly not a coracoid, although a proper identification would benefit from figures in multiple perspectives.
Contrary to Sanchez-Hernandez and Benton's phylogenetic conclusion, Rauhut et al. stated "is a megalosauroid tetanuran, and probably a spinosaurid", and the latter has since been confirmed by Samathi, Samathi et al. and Malafaia et al... Samathi et al. added it to Carrano et al.'s tetanurine analysis and found "The spinosaurid character in Camarillasaurus is the lack of an anterior process on the chevron", while a "likely spinosaurine character in Camarillasaurus is pachyostosis in the tibia", matching Barker et al.'s (2021) recovery of it using Cau's megamatrix as the basalmost spinosaurine.
References- Sanchez-Hernandez and Benton, 2014 (online 2012). Filling the ceratosaur gap: A new ceratosaurian theropod from the Early Cretaceous of Spain. Acta Palaeontologica Polonica. 59(3), 581-600.
Wang, Stiegler, Amiot, Wang, Du, Clark and Xu, 2016. Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology. 27(1), 144-148.
Rauhut, Canudo and Castanera, 2019. A reappraisal of the Early Cretaceous theropod dinosaur Camarillasaurus from Spain. XVII Conference of the EAVP, Program and Abstracts. 96.
Samathi, 2019. Theropod dinosaurs from Thailand and southeast Asia Phylogeny, evolution, and paleobiogeography. PhD thesis, Rheinischen Friedrich-Wilhelms-Universität Bonn. 249 pp.
Malafaia, Gasulla, Escaso, Narvaez and Ortega, 2020. An update of the spinosaurid (Dinosauria: Theropoda) fossil record from the Lower Cretaceous of the Iberian Peninsula: Distribution, diversity, and evolutionary history. Journal of Iberian Geology. 46, 431-444.
Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports. 11:19340.
Samathi, Sander and Chanthasit, 2021 online. A spinosaurid from Thailand (Sao Khua Formation, Early Cretaceous) and a reassessment of Camarillasaurus cirugedae from the Early Cretaceous of Spain. Historical Biology. Latest Articles. DOI: 10.1080/08912963.2021.1874372

Oxalaia Kellner, Azevedo, Machado, Carvalho and Henriques, 2011
O. quilombensis Kellner, Azevedo, Machado, Carvalho and Henriques, 2011
Early Cenomanian, Late Cretaceous
Alcântara Formation of the Itapecuru Group, Brazil
Holotype- (MN 6117-V) (skull ~1.35 m) incomplete premaxillae
Paratype- ?(MN 6119-V) maxillary fragment
Referred- ?(UFMA 1.10.229) distal caudal centrum (Medeiros and Schultz, 2002)
?(UFMA 1.10.240) distal caudal centrum (94 mm) (Medeiros and Schultz, 2002)
?(UFMA 1.20.070; morphotype 1) tooth (80 mm) (Medeiros and Schultz, 2002)
?(UFMA 1.20.443; morphotype 2) tooth (~101 mm) (Medeiros, 2006)
?(UFMA 1.20.444 SL; 25 morphotype 1, 95 morphotype 2 and 153 intermediates) 273 complete to fragmentary teeth (13-98 mm) (Medeiros, 2006)
Diagnosis- (after Kellner et al., 2011) maximum expansion of the distal end of the premaxillae between the 3rd and 4th alveoli; anterior projection of the maxillae between the premaxillae in the palatal region very thin; diastema between the 5th and 6th premaxillary teeth present but shorter than in Spinosaurus; ventral portion of the premaxillae very sculptured.
Other diagnoses- Kellner et al. (2011) also listed the lack of serrations in their diagnosis, but this is true of all spinosaurines. They listed "presence of two replacement teeth associated with the 3rd functional tooth", but Isasmendi et al. (2020) noted "two replacement teeth at different stages of development within the same alveolus ... has been documented in the maxilla of Irritator", the dentary of "Iberospinus, and now in the CPI 477 maxilla." The condition in Iberospinus where only the third and fifth dentary tooth positions have two replacement teeth suggests its presence in Oxalaia's third premaxillary tooth is developmental happenstance, so that positional specificity should not be used as an autapomorphy.
Comments- Oxalaia's holotype was first mentioned in abstracts (Machado and Kellner, 2008; Machado et al., 2009) before being described as a new taxon by Kellner et al. (2011).
Isasmendi et al. (2023) noted that "In MN 6119-V, the preserved alveoli are visible in lateral view with laterocumbent teeth as occurs in Spinosaurinae and unlike baryonychines. Furthermore, the paradental groove in MN 6119-V is wavy or sinusoidal in a similar way to that of African spinosaurines (e.g., MNHN SAM 124, MSNM V4047 and NNHMUK PV R 16420), and differing from the straighter one of baryonychines." While these suggest spinosaurine affinities, they furthermore proposed "The medial shelf of MN 6119-V is subcylindrical, but in ventral view it does not seem to be strongly directed medially, unlike the specimens MNHN SAM 124, MSNM V4047 and NHMUK PV R 16420 that have a proportionally wider medial shelf", and "The height between the external naris and the alveolar margin is more similar to that of baryonychines and Irritator, which could indicate it was more anteriorly located than in MSNM V4047 and NHMUK PV R 16420." Thus, "it seems that O. quilombensis cannot be attributed to Spinosaurus aegyptiacus and, instead, would be more closely related to Irritator."
The isolated teeth are all unserrated and thus referred to Spinosaurinae. However, they vary morphologically with two extremes and numerous intermediates. Morphotype 1 is nearly straight with labial and lingual fluting, rounded in section and lacks enamel wrinkles at its base. Morphotype 2 is straight to gently curved, lacks fluting, is round to labiolingually compressed in section and sometimes has basal enamel wrinkles. While Medeiros considered morphotype 2 to be a different, otherwise unknown theropod taxon, I find it more likely that the numerous intermediates and similar size ranges indicate positional variation within the jaw. Kellner et al. noted morphotype 1 resembled Oxalaia from the same formation, so all of the teeth are tentatively referred to that taxon here.
Medeiros and Schultz (2002) referred two distal caudals to Sigilmassasaurus, which Ibrahim et al. (2014) recently synonymized with Spinosaurus. As the spinosaurine Oxalaia is from the same formation, it is likely the caudals belong to it.
References- Medeiros and Vilas Boas, 1999. Ocorrência de uma paleocomunidade continental do Cenomaniano (Cretáceo Superior) do Nordeste do Brasil. In: Jornadas Argentinas de Paleontologia de Vertebrados. Resúmenes, La Plata, UNLP. 15, 18.
Medeiros and Schultz, 2001. Uma paleocomunidade de vertebrados do Cretáceo médio, bacia de São Luís. In Rossetti, Góes Truckenbrodt (eds.). O Cretáceo na bacia de São Luís - Grajaú, Museu Emílio Goeldi. 209-221.
Medeiros and Schultz, 2002. A fauna dinossauriana da Laje do Coringa, Cretáceo médio do Nordeste do Brasil. Arquivos do Museu Nacional. 60(3), 155-162.
Medeiros, 2005. Spinosaurid teeth variation in mid-Cretaceous Alcântara Formation, Maranhão state, Brazil. Resumos do XIX Congresso Brasileiro de Paleontologia e VI Congresso Latino-Americano de Paleontologia. [pp unknown]
Medeiros, 2006. Large theropod teeth from the Eocenomanian of northeastern Brazil and the occurrence of Spinosauridae. Revista Brasileira de Paleontologia. 9(3), 333-338.
Machado and Kellner, 2008. An overview of the Spinosauridae (Dinosauria, Theropoda) with comments on the Brazilian material. Journal of Vertebrate Paleontology. 28(3), 109A.
Machado, Azevedo, Carvalho, Henriques and Kellner, 2009. A new spinosaurid from the Cretaceous Alcantara Formation (Maranhao), Northeastern Brazil. Journal of Vertebrate Paleontology. 29(3), 138A-139A.
Kellner, Azevedo, Machado, Carvalho and Henriques, 2011. A new dinosaur (Theropoda, Spinosauridae) from the Cretaceous (Cenomanian) Alcântara Formation, Cajual Island, Brazil. Anais da Academia Brasileira de Ciências. 83(1), 99-108.
Ibrahim, Sereno, Dal Sasso, Maganuco, Fabbri, Martill, Zouhri, Myhrvold and Iurino, 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science. 345(6204), 1613-1616.
Sales and Schultz, 2017. Spinosaur taxonomy and evolution of craniodental features: Evidence from Brazil. PloS One. 12(11), e0187070.
Smyth, Ibrahim and Martill, 2020. Sigilmassasaurus is Spinosaurus: A reappraisal of African spinosaurines. Cretaceous Research. 114:104520.
Lacerda, Grillo and Romano, 2022 (2021 online). Rostral morphology of Spinosauridae (Theropoda, Megalosauroidea): Premaxilla shape variation and a new phylogenetic inference. Historical Biology. 34(11), 2089-2109.
Isasmendi, Navarro-Lorbés, Sáez-Benito, Viera, Torices and Pereda-Suberbiola, 2023 (2022 online). New contributions to the skull anatomy of spinosaurid theropods: Baryonychinae maxilla from the Early Cretaceous of Igea (La Rioja, Spain). Historical Biology. 35(6), 909-923.

Irritator Martill, Cruikshank, Frey, Small and Clarke, 1996
= “Angaturama” Anonymous, 1995
= Angaturama Kellner and Campos, 1996
I. challengeri Martill, Cruikshank, Frey, Small and Clarke, 1996
= Angaturama limai Kellner and Campos, 1996
Albian, Early Cretaceous
Romualdo Member of Santana Formation, Brazil
Holotype- (SMNS 58022) incomplete skull (~840 mm), teeth, partial mandibles
Referred- (GP/2T-5; holotype of Angaturama limai) premaxillae, anterior maxilla, anterior nasal
Diagnosis- (after Sues et al., 2002) nasals with prominent median bony crest that terminates posteriorly in knob-like, somewhat dorsoventrally flattened projection; dorsal surface of parietals facing posterodorsally and vertical axis of braincase inclined anteroventrally; posterior surface of basisphenoid with deep, dorsoventrally oval median recess; surangular with broad lateral shelf (also in Baryonyx).
(after Carrano et al., 2012) midline ridge on dorsal surface of conjoined premaxillae extending further anteriorly than in Baryonyx and Cristatusaurus.
Comments- Angaturama was originally printed on a Brazilian postage stamp in 1995.
References- Martill, Cruickshank, Frey, Small and Clarke, 1996. A new crested maniraptoran dinosaur from the Santana Formation (Lower Cretaceous) of Brazil. Journal of the Geological Society of London. 153, 5-8.
Kellner and Campos, 1996. First Early Cretaceous theropod dinosaur from Brazil with comments on Spinosauridae. Neues Jahrbuch fuer Geologie und Palaeontologie Abhandlungen. 199(2), 151-166.
Sues, Frey, Martill and Scott, 2002. Irritator challengeri, a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil. Journal of Vertebrate Paleontology. 22(3), 535-547.
Carrano, Benson and Sampson, 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 10(2), 211-300.

unnamed Spinosaurinae (Alonso and Canudo, 2016)
Barremian, Early Cretaceous
Blesa Formation, Spain
Material- (CAN1 960) tooth (16.19x6.22x5.96 mm) (Alonso and Canudo, 2016)
(CAN1 968) tooth (17.15x5.69x4.44 mm) (Alonso and Canudo, 2016)
(CAN1 973) tooth (22.19x9.68x8.34 mm) (Alonso and Canudo, 2016)
(CAN1 990) tooth (17.93x7.23x6.42 mm) (Alonso and Canudo, 2016)
(HOC 17) tooth (23.25x10.27x8.61 mm) (Alonso et al., 2018)
(HOC 28) tooth fragment (Alonso et al., 2018)
References- Alonso and Canudo, 2016. On the spinosaurid theropod teeth from the Early Barremian (Early Cretaceous) Blesa Formation (Spain). Historical Biology. 28(6), 823-834.
Alonso, Gasca, Navarro-Lorbés, Rubio and Canudo, 2018. A new contribution to our knowledge of the large-bodied theropods from the Barremian of the Iberian Peninsula: The "Barranco del Hocino" site (Spain). Journal of Iberian Geology. 44, 7-23.

unnamed Spinosaurinae (Navarro-Lorbés and Torices, 2018)
Late Barremian-Early Aptian, Early Cretaceous
Enciso Group, El Horcajo, Trevijano, La Rioja, Spain
Material- (CP-UR-3; = Tre-3) incomplete tooth (37.12x12.72x10.75 mm) (Navarro-Lorbés and Torices, 2018)
(CP-UR-5) tooth (?x18.71x15.4 mm) (Isasmendi, Sáez-Benito, Torices, Navarro-Lorbés and Pereda-Suberbiola, 2020)
(CP-UR-8) tooth (42.50x12.71x12.04 mm) (Isasmendi, Sáez-Benito, Torices, Navarro-Lorbés and Pereda-Suberbiola, 2020)
(CP-UR-18) (juvenile) tooth (14.10x5.11x4.97 mm) (Isasmendi, Sáez-Benito, Torices, Navarro-Lorbés and Pereda-Suberbiola, 2020)
Comments-Navarro-Lorbés and Torices (2018) described these as Morphotype 3- "Medium sized teeth (CH = 40-50 mm). Ellipsoidal to subcircular basal cross section and very elongated crown apically. Curved posteriorly and with a slight curvature to the lingual region too. Fluted enamel covering all the surface of the crown. Some of the specimens present irregularities in the carinae, discarding them as microdenticles because of their irregularities and their little development." While Navarro-Lorbés and Torices only analyzed and figured one (as Tre-3), and never stated how many of their 19 new theropod and crocodyliform teeth were Morphotype 3, Isasmendi et al. (2020) included three of them in their discriminant analysis, which provided specimen numbers and measurements for what turns out to be four teeth in total. They found "CPUR-18 falls within the smaller teeth of the sample and is similar in size to those of La Cantalera-1 and Galve. CP-UR-3 and CP-UR-8 are more closely grouped with the teeth attributed to Spinosaurus", and thus "The similarity in size between CP-UR-18 and the teeth of La Cantalera-1 could indicate that its producer was a juvenile spinosaurine theropod."
References- Navarro-Lorbés and Torices, 2018. Preliminary analysis of theropod teeth from "El Horcajo", Lower Cretaceous, La Rioja, Spain. XVI Encuentro de Jovenes Investigadoras en Paleontologia, Libro de Resumenes. 83-86.
Isasmendi, Sáez-Benito, Torices, Navarro-Lorbés and Pereda-Suberbiola, 2020. New insights about theropod palaeobiodiversity in the Iberian Peninsula and Europe: Spinosaurid teeth (Theropoda, Megalosauroidea) from the Lower Cretaceous of La Rioja (Spain). Cretaceous Research. 104600.

Ichthyovenator Allain, Xaisanavong, Richir and Khentavong, 2012
I. laosensis Allain, Xaisanavong, Richir and Khentavong, 2012
Late Barremian-Early Cenomanian (Aptian?), Early-Late Cretaceous
Grès supérieurs Formation, Laos
Holotype- (MDS BK10-01 to 15) twelfth dorsal vertebra (135 mm), partial thirteenth dorsal neural spine, posterior dorsal rib, dorsal rib (uncollected), first sacral neural spine, second sacral neural spine, fused partial second and third sacral vertebrae (third ~90 mm), third sacral neural spine, fourth sacral neural spine, fifth sacral neural spine, third sacral ribs, first caudal vertebra (101 mm), second caudal vertebra (95 mm), ilia (one incomplete; 920, 910 mm), pubes (one fragmentary; 650 mm), ischia (one incomplete; 496, 494 mm)
Referred- (presacrals are MDS BK10-16 to 25) three teeth, cervical series, first dorsal vertebra, seven caudal vertebrae, pubis (Allain, 2014)
(caudals are MDS BK97-02) teeth, vertebrae (Allain, 2014)
Diagnosis- (after Allain et al., 2012) sinusoidal dorsosacral sail; twelfth dorsal neural spine is 410% of centrum length with anterodistal finger-like process; posterior dorsal ribs articulated with sternal complex; fan-shaped sacral neural spines 3 and 4; transverse processes of first caudal vertebra with sigmoid profile in dorsal view; deep prezygapophyseal centrodiapophyseal fossae on first caudal vertebra; long iliac blade with ilium/pubis length ratio 1.46; proximal pubic plate with obturator and pubic notches; large ischial plate with ischial foramen (also in Monolophosaurus); mediolaterally flattened ischial shaft.
Comments- The holotype was discovered in 2010.
Allain (2014) discuss a new specimen preserving a first dorsal vertebra which is "nearly identical to" Sigilmassasaurus (now a junior synonym of Spinosaurus), showing an articular surface much wider than high (ratio of ~1.5) and absent interzygapophyseal laminae.
Relationships- Allain et al. (2012) found Ichthyovenator to be a basal baryonychine in their phylogenetic analysis. Yet adding it to the much larger matrix of Carrano et al. 2012 results in it being sister to Concavenator, with both sister to Allosauria in Carnosauria. Given the Sigilmassasaurus-like cervicodorsals and newly recovered teeth in that specimen which are unrecurved and unserrated, Allain (2014) proposed Ichthyovenator was "more closely related to Spinosaurinae." Indeed, Barker et al. (2021) recovered it as a spinosaurine using Cau's matrix.
Several characters led to its placement in Allain et al.'s analysis. Ichthyovenator is a megalosauroid due to having a vertical ridge anterior to the hyposphene in the dorsal vertebra. But this is miscoded and is actually absent. It is a spinosaurid due to the tall dorsal neural spine, but these are also present in Concavenator, which was not included in their analysis. It is placed in Baryonychinae due to four characters- the basally webbed dorsal neural spine; an accessory centrodiapophyseal lamina on the dorsal, which is also present in the unincluded Concavenator; a mediolaterally expanded pubic boot; and a posterior pubic boot "reduced to a small flange". But this is correlated with the last character, as a reduced posterior pubic boot (with insignificant anterior boot) leads to a transversely expanded distal pubis, and indeed the only taxa coded as having the first state are also the only ones coded as having the second. It is excluded from Avetheropoda due to the small pubic boot as well. It is excluded from Allosauroidea because the posterior dorsal neural spine is not anteriorly inclined, but this is also true in Concavenator. It's also excluded from Allosauroidea because the ischial obturator foramen is closed. Yet the two coelurosaurs are miscoded as lacking an obturator notch; when that's corrected, Ichthyovenator's closed foramen becomes an autapomorphy. So of the nine characters, three are due to Allain et al. not including Concavenator in their matrix. Two are due to miscodings. The two pubic boot characters are correlated, so are really only one. Yet this character was not used by Carrano et al.. Finally, the neural spine webbing is valid and used by Carrano et al..
Several characters also led to its placement when added to Carrano et al.'s matrix. Ichthyovenator is sister to Concavenator due to three characters- tall dorsal neural spine (also in spinosaurids); accessory centrodiapophyseal lamina (also in baryonychines); peg-and-socket ilioischial articulation (not included by Allain et al.). It is an avetheropod because of four characters- narrow brevis fossa (coded more strictly in Allain et al.); m. cuppedicus shelf (not included by Allain et al.); open pubic obturator notch (however, Suchomimus and Baryonyx are miscoded by Carrano et al.); large and oval pubic obturator notch/foramen (not included by Allain et al.). It is excluded from Megalosauroidea due to lacking a vertical ridge anterior to the hyposphene (miscoded by Allain et al.) and having a vertical ilial ridge (not included by Allain et al.). So of these nine characters, two are correctly coded as also present in baryonychines, and another is miscoded as being absent in baryonychines. Five others aren't used by Allain et al., and another was miscoded by Allain et al.
In Allain et al.'s matrix, after recoding Ichthyovenator's hyposphene ridge and the coelurosaurs' obturator notches, and excluding the correlated pubic boot character, Ichthyovenator is still a baryonychine. Constraining it to be a non-allosauroid carnosaur takes five extra steps, which is down from the nine it took before (though it was slightly more nested in Carnosauria before, but only by two steps). If we add the five characters Allain et al. didn't use that support this position (the brevis fossa expansion counts too since it is an additional state of a character), then either alternative is equally parsimonious.
In Carrano et al.'s matrix, after recoding Baryonyx and Suchomimus for their obturator notches, Ichthyovenator is now a non-allosauroid carnosaur and Concavenator is back to Carcharodontosauridae (making them sister taxa like before is now one step longer). It now takes three more steps to make Ichthyovenator a spinosaurid, so that's dropped from five. If we add on the transversely wide pubic boot character that they didn't include, we could say it only takes two more steps to make Ichthyovenator a spinosaurid.
In conclusion, both matrices give almost the same answer- Ichthyovenator is either equally likely or two steps more likely to be a basal carnosaur instead of a baryonychine. The main flaw was that Allain et al. didn't include several useful characters, which is expected considering their total character number is 51% of Carrano et al.'s. Allain's (2014) new report of serrationless and unrecurved teeth could at least even the odds in favor of a spinosaurid identity, though these do contradict it being a baryonychine. This is perhaps why when those two characters are newly coded for Ichthyovenator in Carrano et al.'s matrix, the same general results are found as reported above (three more steps needed to move it from a basal carnosaur to a spinosaurid). However, given the morphology of the undescribed teeth and cervicodorsals, the taxon is provisionally placed in Spinosaurinae here.
References- Allain, Xaisanavong, Richir and Khentavong, 2012. The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the Early Cretaceous of Laos. Naturwissenschaften. 99(5), 369-377.
Allain, 2014. New material of the theropod Ichthyovenator from Ban Kalum type locality (Laos): Implications for the synonymy of Spinosaurus and Sigilmassasaurus and the phylogeny of Spinosauridae. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 78.
Evers, Rauhut, Milner, McFeeters and Allain, 2015. A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the "middle" Cretaceous of Morocco. PeerJ. 3:e1323.
Barker, Hone, Naish, Cau, Lockwood, Foster, Clarkin, Schneider and Gostling, 2021. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports. 11:19340.

Siamosaurus Buffetaut and Ingavat, 1986
S. suteethorni Buffetaut and Ingavat, 1986
Late Barremian, Early Cretaceous
Phu Wiang 1, Sao Khua Formation, Thailand
Holotype- (DMR TF 2043a) tooth (47.7x16.6x12.5 mm)
Paratypes- (DMR TF 2043b) tooth (24.3x?x? mm)
(DMR TF 2043c) tooth
(DMR TF 2043d) tooth
(DMR TF 2043e) tooth
(DMR TF 2043f) tooth
(DMR TF 2043g) tooth
(DMR TF 2043h) tooth
(DMR TF 2043i) tooth
Late Barremian, Early Cretaceous
Kalasin 2, Sao Khua Formation, Thailand
Referred- teeth (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Kalasin 11, Sao Khua Formation, Thailand
tooth (Buffetaut and Suteethorn, 1999)
Late Barremian, Early Cretaceous
Phu Din Daeng, Sao Khua Formation, Thailand
(PRC147) incomplete tooth (?x~16x? mm) (Tong, Buffetaut, Suteethorn, Suteethorn, Cuny, Cavin, Deesri, Martin, Wongko, Naksri and Claude, 2019)
(PRC coll.) teeth (Tong, Buffetaut, Suteethorn, Suteethorn, Cuny, Cavin, Deesri, Martin, Wongko, Naksri and Claude, 2019)
Late Barremian, Early Cretaceous
Phu Sung, Sao Khua Formation, Thailand
tooth fragment (Chanthasit, Suteethorn, Naksri, Tong, Wongko and Sonoda, 2019)
Late Barremian, Early Cretaceous
Phu Wiang 5A, Sao Khua Formation, Thailand
tooth (Martin, Suteethorn and Buffeaut, 1999)
Late Barremian, Early Cretaceous
Sao Khua Formation, Thailand
tooth (Kobayashi, Takai and Hayami, 1964)
Late Barremian?, Early Cretaceous
Pra Prong, Sao Khua Formation?, Thailand
teeth (Suteethorn, Buffetaut, Wongo, Suteethorn and Tong, 2018)
Diagnosis- (after Buffetaut and Ingavat, 1986) serrationless carinae; oval cross section; ~15 flutes on both labial and lingual sides.
Comments- Buffetaut and Ingavat (1983) briefly described and figured the type teeth, referring them "either to an unusual theropod dinosaur, or to some unknown crocodilian." Buffetaut and Ingavat (1986) later fully described and named the teeth Siamosaurus suteethorni, referred to ?Spinosauridae. Buffetaut and Suteethorn (1999) noted that in Kobayashi et al. (1964) "the so-called 'ichthyosaur' tooth apparently belongs to the enigmatic dinosaur Siamosaurus." Buffetaut and Suteethorn (1999) state "isolated teeth of Siamosaurus are frequently found at many vertebrate localities in the Sao Khua Formation, sometimes, as at Wat Sakawan, in association with sauropod remains." Martin et al. (1999) list a few localities with Siamosaurus teeth. Samathi et al. (2019) note "in fact, there are very small serrations on unworn teeth (E. Buffetaut pers. comm.; A.S.pers. obs.)", which would remove the reason to assign this to Spinosaurinae. Suteethorn et al. (2018) note a spinosaurid morph at the Pra Prong locality "with a strongly ornamented crown, bearing up to 25 well-marked ridges on each face, and with a wrinkled enamel between the ridges and at the apex." Buffetaut is preparing a redescription of the taxon (pers. comm. to Bertin, 2010).
References- Kobayashi, Takai and Hayami, 1964. On some Mesozoic fossils from the Khorat Series of east Thailand and a note on the Khorat Series. Japanese Journal of Geology and Geography. 34, 181-192.
Buffetaut and Ingavat, 1983. Vertebrates from the continental Jurassic of Thailand. CCOP Technical Bulletin. 16, 68-75.
Buffetaut and Ingavat, 1986. Unusual theropod dinosaur teeth from the Upper Jurassic of Phu Wiang, northeastern Thailand. Revue de Paléobiologie. 5, 217-220.
Buffetaut and Suteethorn, 1999. The dinosaur fauna of the Sao Khua Formation of Thailand and the beginning of the Cretaceous radiation of dinosaurs in Asia. Palaeogeography, Palaeoclimatology, Palaeoecology. 150, 13-23.
Martin, Suteethorn and Buffeaut, 1999. Description of the type and referred material of Phuwiangosaurus sirindhornae Martin, Buffetaut and Suteethorn, 1994, a sauropod from the Lower Cretaceous of Thailand. Oryctos. 2, 39-91.
Bertin, 2010. A catalogue of the material and review of the Spinosauridae. PalArch’s Journal of Vertebrate Palaeontology. 7(4), 1-39.
Suteethorn, Buffetaut, Wongo, Suteethorn and Tong, 2018. Morphological diversity of spinosaurid teeth from the Pra Prong locality (Lower Cretaceous of eastern Thailand). 5th International Palaeontological Congress. Abstract Book, 666.
Chanthasit, Suteethorn, Naksri, Tong, Wongko and Sonoda, 2019. New vertebrate fossil site from the Early Cretaceous Sao Khua Formation, Sakon Nakhon Province, northeastern Thailand. Open Journal of Geology. 9, 619-622.
Samathi, Chanthasit and Sander, 2019. A review of theropod dinosaurs from the Late Jurassic to mid-Cretaceous of southeast Asia. Annales de Paléontologie. 105(3), 201-215.
Tong, Buffetaut, Suteethorn, Suteethorn, Cuny, Cavin, Deesri, Martin, Wongko, Naksri and Claude, 2019. Phu Din Daeng, a new Early Cretaceous vertebrate locality on the Khorat Plateau, NE Thailand. Annales de Paléontologie. 105(3), 223-237.
S. sp. (Buffetaut and Suteethorn 1998)
Aptian, Early Cretaceous
Ban Pia Fan, Khok Kruat Formation, Thailand
Material- (SM-KK 14; Khok Kruat spinosaurid) fourth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, tenth cervical vertebra, dorsal vertebrae, neural spine, ribs, chevron, metacarpal?, pelvic elements including pubis (Buffetaut, Suteethorn and Tong, 2004)
?...(SM-KK coll.) teeth (Buffetaut, Suteethorn, Le Loeuff, Khansubha, Tong and Wongko, 2005)
Aptian, Early Cretaceous
Khok Pa Suam, Khok Kruat Formation, Thailand
(PM2016-1-003) tooth (44.3x16.7x13.1 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
(PM2016-1-004) tooth (43.3x16.9x13.1 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
(PM2016-1-005) tooth (39.8x19.6x14.7 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
Aptian, Early Cretaceous
Sam Ran, Khok Kruat Formation, Thailand
(PM2016-1-001) tooth (20x12.8x9.5 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
(PM2016-1-002) tooth (33.6x15.4x14.3 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
Aptian, Early Cretaceous
Lam Pao Dam, Khok Kruat Formation, Thailand
(PM2016-1-006) tooth (56.8x15x14.6 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
(PM2016-1-007) tooth (75.1x32.6x22.7 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
(PM2016-1-008) tooth (46.3x16x11 mm) (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
Aptian, Early Cretaceous
Khok Kruat Formation, Thailand
(PM2016-1 coll.) almsot fifty teeth (Wongko, Buffetaut, Khamha and Lauprasert, 2019)
Description- Buffetaut and Suteethorn (1998) say "a form similar to the earlier Siamosaurus seems to be present" in the Khok Kruat Formation.
Buffetaut et al. (2005) state "at Khok Pa Suam, more unusual teeth showing an uncompressed crown, extremely reduced serrations and conspicuous ribbing have been found. Although they resemble crocodilian teeth, their type of curvature suggests that they belong to theropods. They are strongly reminiscent of ... Siamosaurus suteethorni ... and tentatively referred to a spinosaurid."
Wongko et al. (2019) describe a few teeth each from the Khok Pa Suam, Sam Ran and Lam Pao Dam localities out of "almost sixty spinosaurid teeth ... collected by Staffs of the Department of
Mineral Resources during the last decade."
Buffetaut et al. (2005) also mention a partial skeleton of a spinosaurid from another locality and figure a neural spine intermediate in length between Baryonyx and Spinosaurus. Siamosaurus teeth "have been found close to the above-mentioned skeletal elements, but they may be evidence of scavenging rather than anatomical association."
Milner et al. (2017) state "have been found close to the above-mentioned skeletal elements, but they may be evidence of scavenging rather than anatomical association."
Samathi et al. (2019) identify the cervical numbers based on Evers et al.'s revision of spinosaurid presacral vertebrae, and note it differs from Ichthyovenator in that "SM-KK 14 has an anteriorly concave pubis and a posteriorly curved chevron." They figure the pubis as well.
References- Buffetaut and Suteethorn, 1998. Early Cretaceous dinosaurs from Thailand and their bearing on the early evolution and biogeographical history of some groups of Cretaceous dinosaurs. In Lucas, Kirkland and Estep (eds.). New Mexico Museum of Natural History Bulletin. 14, 205-210.
Buffetaut, Suteethorn and Tong, 2004. Asian spinosaur confirmed. SVPCA 2004 Abstracts. [pp unknown]
Buffetaut, Suteethorn, Le Loeuff, Khansubha, Tong and Wongko, 2005. The dinosaur fauna from the Khok Kruat Formation (Early Cretaceous) of Thailand. In Wannakao, Youngme, Srisuk and Lertsirivorakul (eds.). Proceedings of the International Conference on Geology, Geotechnology and Mineral Resources of Indochina. 575-581.
Milner, Buffetaut and Suteethorn, 2007. A tall-spined spinosaurid theropod from Thailand and the biogeography of spinosaurs. Journal of Vertebrate Paleontology. 27(3), 118A.
Samathi, Chanthasit and Sander, 2019. A review of theropod dinosaurs from the Late Jurassic to mid-Cretaceous of southeast Asia. Annales de Paléontologie. 105(3), 201-215.
Wongko, Buffetaut, Khamha and Lauprasert, 2019. Spinosaurid theropod teeth from the red beds of the Khok Kruat Formation (Early Cretaceous) in northeastern Thailand. Tropical Natural History. 19(1), 8-20.
S? fusuiensis (Hou, Yeh and Zhao, 1975) new comb.
= Sinopliosaurus fusuiensis Hou, Yeh and Zhao, 1975
Aptian, Early Cretaceous
Xinlong Formation, Guangxi, China
Holotype- (IVPP V4793) five teeth (one lost) (69x16.5x13 mm)
Referred- (NP03) tooth (Amoit et al., 2010)
(NP07) tooth (Amoit et al., 2010)
teeth (Dong, 1992)
several teeth (Mo et al., 2014)
Diagnosis- provisionally indeterminate relative to Siamosaurus suteethorni.
Comments- Note the Xinlong Formation used to be called the Napai Formation (Mo et al., 2015). These teeth were discovered in 1973 and originally described by Hou et al. (1975) as a new species of the pliosaur genus Sinopliosaurus. The type of that genus, S. weiyuanensis, preserves only teleosaurid teeth in addition to crocodilian vertebrae and a pliosaurid ischium and femur (the type specimen) (Young, 1948). Buffetaut et al. (2008) later redescribed the S. fusuiensis teeth and discovered they were spinosaurid. In particular, they resemble spinosaurines in a lack of serrations, but baryonychines in the presence of ribbing. They are identical with those of Siamosaurus suteethorni and both differ from Baryonyx in having stronger ribbing. They suggested the teeth belong to a taxon "closely related to, if not identical with Siamosaurus suteethorni", which I have formalized here by placing fusuiensis in Siamosaurus.
Dong (1992) mentioned spinosaurine teeth from the Napai Formation, but whether these refer to the S? fusuiensis material is uncertain. Mo et al. (2014) note spinosaurid teeth in three additional localities.
References- Young, 1948. Fossil crocodiles in China, with notes on dinosaurian remains associated with the Kansu crocodiles. Bulletin of the Geological Society of China. 28, 235-288.
Hou, Yeh and Zhao, 1975. Fossil reptiles from Fusui, Kwangshi. Vertebrata Palasiatica. 13, 23-33.
Dong, 1992. Dinosaurian Fauna's of China. Ocean Press/Springer-Verlag, Beijing/Berlin. 188 pp.
Buffetaut, Suteethorn, Tong and Amiot, 2008. An Early Cretaceous spinosaurid theropod from southern China. Geological Magazine. 145(5), 745-748.
Amiot, Buffetaut, Lecuyer, Wang, Boudad, Ding, Fourel, Hutt, Martineau, Medeiros, Mo, Simon, Suteethorn, Sweetman, Tong, Zhang and Zhou, 2010. Oxygen isotope evidence for semiaquatic habits among spinosaurid theropods. Geology. 38, 139-142.
Mo, Huang, Xie and Buffetaut, 2014. A megatheropod tooth from the Early Cretaceous of Fusui, Guangxi, Southern China. Acta Geologica Sinica (English Edition). 88(1), 6-12.
Mo, Buffetaut, Tong, Amiot, Cavin, Cuny, Suteethorn, Suteethorn and Jiang, 2015. Early Cretaceous vertebrates from the Xinlong Formation of Guangxi (southern China): A review. Geological Magazine. doi:10.1017/S0016756815000394
S? sp. (Hasegawa, Buffetaut, Manabe and Takakuwa, 2003)
Barremian, Early Cretaceous
Lower Member of the Sebayashi Formation, Japan
Material- (GMNH-PV-999) tooth (51x20x14 mm) (Hasegawa, Buffetaut, Manabe and Takakuwa, 2003)
(KDC-PV-0003) partial tooth (?x7.4x6.8 mm) (Kubota, Takakuwa and Hasegawa, 2017)
Comments- This is almost identical to Siamosaurus according to Hasegawa et al. (2003) and Buffetaut et al. (2008).
References- Hasegawa, Buffetaut, Manabe and Takakuwa, 2003. A possible spinosaurid tooth from the Sebayashi Formation (Lower Cretaceous), Gunma, Japan. Bulletin of Gunma Museum of Natural History. 7, 1-5.
Buffetaut, Suteethorn, Tong and Amiot, 2008. An Early Cretaceous spinosaurid theropod from southern China. Geological Magazine. 145(5), 745-748.
Kubota, Takakuwa and Hasegawa, 2017. Second discovery of a spinosaurid tooth from the Sebayashi Formation (Lower Cretaceous), Kanna Town, Gunma Prefecture, Japan. Bulletin of Gunma Museum of Natural History. 21, 1-6.

unnamed spinosaurine (Lü, Xu, Jiang, Jia, Li., Yuan, Zhang and Ji, 2009)
Aptian-Albian, EarlyCretaceous
Haoling Formation, Henan, China
Material- (41HIII-00012) tooth (10x5x? mm)
Comments- While originally assigned to the Mangchuan Formation, Xu et al. 2012 split this into three new formations and revised their ages.
Reference- Lü, Xu, Jiang, Jia, Li., Yuan, Zhang and Ji, 2009. A preliminary report on the new dinosaurian fauna from the Cretaceous of the Ruyang Basin, Henan Province of Central China. Journal of the Paleontological Society of Korea. 25, 43-56.

undescribed Spinosaurinae (Pause, 1979)
Early-Middle Campanian, Late Cretaceous
El Atrun, Quseir Formation, Kharga Oasis, Egypt
Material- tooth (Pause, 1979)
teeth (Churcher, 1995)
Early-Middle Campanian, Late Cretaceous
Bee's Friday Site, Quseir Formation, Dakhla Oasis, Egypt
(ROM? coll.) teeth and/or bones (Churcher, 1999)
Early-Middle Campanian, Late Cretaceous
Wadi el-Battikh (Lungfish Bluff), Quseir Formation, Dakhla Oasis, Egypt
(ROM? coll.) material (Churcher and De Iluliis, 2001)
Comments- According to Churcher (1995), Pause's 1979 thesis reported "a damaged tooth assigned to Spinosaurus aegyptiacus" from "from east of the Darb el-Arbain, at El Atrun, near Maks Bahri, at km 11 south of Baris", but this to too late to be congeneric with the Cenomanian Spinosaurus.
Churcher further noted "On a visit to the El Atrun exposure in 1993, Dr. Dale A. Russell of the Canadian Museum of Nature, Ottawa, and I found isolated teeth of two carnivorous dinosaurs, tentatively assigned to Spinosaurus and Carcharodontosaurus..."
Churcher and Russell (1992) stated in the Dakhla (spelled Dakhleh by them) Oasis, "theropod and sauropod teeth and bones were found a few metres below the Duwi Formation." Churcher's (1999) faunal list matches this with Bee's Friday Site, and says "Subsequently specimens assignable to Spinosaurus and Carcharodontosaurus were obtained", suggesting this was the theropod material mentioned in 1992.
Churcher and de Iluliis (2001) state that at Lungfish Bluff "Identification of Mut Formation shales here is based on the presence of terrestrial dinosaurs (e.g. Spinosaurus)."
References- Pause, 1979 . Uber eine fossile Schildkrote (Podocnemis) aus der mittleren Kreide der sudlichen Kharga-Oasen (SW-Agypten). Thesis, Institut fur Geologie und Palaontologie der Technischen Universitat Berlin. 42 pp.
Churcher and Russell, 1992. Terrestrial vertebrates from Campanian strata in Wadi el-Gedid (Kharga and Dakhleh Oases), Western Desert of Egypt. Journal of Vertebrate Paleontology. 12(3), 23A.
Churcher, 1995. Giant Cretaceous lungfish Neoceratodus tuberculatus from a deltaic environment in the Quseir (=Baris) Formation of Kharga oasis, Western Desert of Egypt. Journal of Vertebrate Paleontology. 15(4), 845-849.
Churcher, 1999. A note on the Late Cretaceous vertebrate fauna of the Dakhleh Oasis. In Churcher and Mills (eds.). Reports from the Survey of Dakhleh Oasis, Western Desert of Egypt, 1977-1987. Dakhleh Oasis Project: Monograph 2. Oxbow Monograph 99. 55-67.
Churcher and De Iluliis, 2001. A new species of Protopterus and a revision of Ceratodus humei (Dipnoi: Ceratodontiformes) from the Late Cretaceous Mut Formation of eastern Dakhleh Oasis, Western Desert of Egypt. Palaeontology. 44(2), 305-323.

Spinosaurus Stromer, 1915
= Sigilmassasaurus Russell, 1996
= Spinosaurini Stromer, 1915 vide Arden, Klein, Zouhri and Longrich, 2019
Definition- (Spinosaurus aegyptiacus <- Irritator challengeri, Oxalaia quilimboensis) (Arden, Klein, Zouhri and Longrich, 2019)
S. aegyptiacus Stromer, 1915
= Sigilmassasaurus brevicollis Russell, 1996
= Spinosaurus marocannus Russell, 1996
Early Cenomanian, Late Cretaceous
Baharija Formation, Egypt
Holotype- (IPHG 1912 VIII 19; destroyed) (~13.2 m; aegyptiacus morphotype) (skull ~1.45 m) maxillary fragment, incomplete dentary (mandible ~1.34 m), splenial, angular, fifteen teeth (~30x15x14, 22x14x10.5, 19x10x9.5, 40x20x16, ~52x22x20, 68x30x~23, 85x34x~24, 47x20x17, 49x21x18, 52x22x18, 41x~19x17, 40x18x15 mm), third cervical neural arch, ~fifth cervical vertebra (185 mm), four dorsal vertebrae (195, 170, ~190, ~210 mm), three dorsal neural arches, anterior dorsal rib, two middle dorsal ribs, posterior dorsal rib, gastralium, six gastralia fragments, incomplete first sacral vertebra (155 mm), second sacral centrum, partial third sacral centrum, proximal caudal vertebra (90 mm)
Referred- (IPHG 1911 XII 21; destroyed) dorsal centrum (145 mm) (Stromer, 1934)
(IPHG 1912 VIII 20; destroyed) tooth (Stromer, 1934)
(IPHG 1912 VIII 21; destroyed) tooth (Stromer, 1934)
(IPHG 1912 VIII 22; destroyed) three cervical centra (~110-120, 120, 140 mm), two partial cervical neural arches, dorsal neural spine, two dorsal ribs, distal fibula (Stromer, 1934)
(IPHG 1922 X45; Spinosaurus B; destroyed) (~9.9 m; brevicollis morphotype) two partial teeth, first dorsal centrum (117 mm), anterior dorsal vertebra (~140 mm), anterior dorsal centrum (~135 mm), incomplete posterior dorsal vertebra (140 mm), posterior dorsal vertebra (160 mm), partial dorsal rib, two proximal ?dorsal ribs, partial gastralia, two ilial fragments, distal femur, tibiae (565, 600 mm), proximal phalanx III-1, phalanx IV-? (45 mm), phalanx IV-? (49 mm), pedal ungual (100 mm) (Stromer, 1934)
(IPHG 1922 X 81; destroyed) chevron fragment (Stromer, 1934)
Albain, Early Cretaceous
Continental Intercalaire, Algeria
(MNHN SAM 124; from Gara Samani) (~14.3 m) (skull ~1.42 m) partial premaxillae, partial maxillae, dentary fragment (Taquet and Russell, 1998)
(MNHN SAM 125; from Gara Samani) premaxillary fragment (Taquet and Russell, 1998)
(MNHN SAM 126; from Gara Samani) cervical centrum (Taquet and Russell, 1998)
(MNHN SAM 127; from Gara Samani) cervical centrum (Taquet and Russell, 1998)
(MNHN SAM 128; from Gara Samani) dorsal neural arch (Taquet and Russell, 1998)
(MNNHN coll.; from Alrar) manual phalanx (300+ mm) (Lapparent, 1960)
(MNNHN coll.; from Alrar) pedal ungual III (100 mm) (Lapparent, 1960)
(MNNHN coll.; from Djoua) manual ungual (85 mm) (Lapparent, 1960)
(Univ. Paris 6; lost; from Djoua) two partial teeth (Haug, 1905)
Aptian-Albian, Early Cretaceous
Chicla Formation, Libya
tooth (Le Loeuff and Metais, 2010)
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Neotype- (FSAC-KK 11888) (~11.3 m; ~17 year old subadult; aegyptiacus morphotype) (skull ~1.12 m) nasal, lacrimal(?), prefrontal, squamosal, quadratojugal, incomplete quadrates (240 mm), dentary fragments, teeth, axis (80 mm), seventh cervical vertebra (~155 mm), partial tenth cervical rib, incomplete fourth dorsal neural spine, partial fifth dorsal vertebra, partial sixth dorsal vertebra (170 mm), fragmentary seventh dorsal vertebra (170 mm), incomplete eighth dorsal vertebra (180 mm), partial ninth dorsal neural spine, several partial dorsal ribs, gastralium, third sacral centrum (145 mm), fourth sacral centrum (140 mm), fifth sacral centrum (135 mm), ~fourth caudal centrum, fragmentary ~eighth caudal neural arch, ~eleventh partial caudal neural spine, ~fourteenth caudal vertebra, five partial chevrons, metacarpal II fragment, phalanx II-1 (175 mm), proximal phalanx II-2, ilia (~710 mm), pubes, ischia (520 mm), femora (610 mm), tibiae (668 mm), fibulae (605 mm), metatarsal I (105 mm), phalanx I-1 (115 mm), pedal ungual I, incomplete metatarsal II, phalanx II-1 (100 mm), phalanx II-2, pedal ungual II, phalanx III-1 (90 mm), phalanx III-2 (65 mm), phalanx III-3, pedal ungual III, phalanx IV-1 (60 mm), phalanx IV-2 (50 mm), phalanx IV-3 (40 mm), phalanx IV-4 (35 mm), pedal ungual IV (80 mm) (Ibrahim et al., 2014)
Referred- (BSPG 2006 I 53) (brevicollis morphotype) tenth cervical vertebra (123 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2006 I 54) (brevicollis morphotype) first dorsal vertebra (139 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2006 I 55) (brevicollis morphotype) first dorsal vertebra (81 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2006 I 56) (brevicollis morphotype) eighth cervical vertebra (119 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2006 I 57) (aegyptiacus morphotype) incomplete sixth cervical vertebra (198 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2008 I 68) (aegyptiacus morphotype) tibia (480 mm) (Rauhut, 2009)
(BSPG 2011 I 115) (brevicollis morphotype) ninth cervical vertebra (123 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2011 I 116) (brevicollis morphotype) tenth cervical vertebra (123 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2011 I 117) (brevicollis morphotype) sixth cervical centrum (184 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2011 I 118) (brevicollis morphotype) ?fifth cervical centrum (212 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
?(BSPG 2013 I 95) (brevicollis morphotype?) anterior dorsal vertebra (168 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(BSPG 2013 I 97) (aegyptiacus morphotype) incomplete ~sixth cervical vertebra (140 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(CMN 41768; paratype of Spinosaurus maroccanus) (aegyptiacus morphotype) mid cervical vertebra (183 mm) (Russell, 1996)
(CMN 41772; paratype of Sigilmassasaurus brevicollis) ?posterior dorsal vertebra (162 mm) (Russell, 1996)
(CMN 41774; paratype of Sigilmassasaurus brevicollis) (brevicollis morphotype) first dorsal vertebra (67 mm) (Russell, 1996)
(CMN 41775; paratype of Sigilmassasaurus brevicollis) mid caudal vertebra (94 mm) (Russell, 1996)
(CMN 41776; paratype of Sigilmassasaurus brevicollis) ?mid dorsal vertebra (~150 mm) (Russell, 1996)
(CMN 41790; paratype of Sigilmassasaurus brevicollis) (brevicollis morphotype) first dorsal vertebra (~127 mm) (Russell, 1996)
(CMN 41800; bone taxon I) distal manual phalanx (Russell, 1996)
(CMN 41820; bone taxon J) manual ungual (81 mm) (Russell, 1996)
(CMN 41850; paratype of Sigilmassasaurus brevicollis) (brevicollis morphotype) third dorsal vertebra (152 mm) (Russell, 1996)
(CMN 41851; paratype of Sigilmassasaurus brevicollis) ?posterior dorsal vertebra (157 mm) (Russell, 1996)
(CMN 41852; bone taxon G) incomplete humerus (~650 mm) (Russell, 1996)
(CMN 41853; paratype of Sigilmassasaurus brevicollis) mid caudal vertebra (110 mm) (Russell, 1996)
(CMN 41854; paratype of Sigilmassasaurus brevicollis) mid caudal vertebra (110 mm) (Russell, 1996)
(CMN 41855; paratype of Sigilmassasaurus brevicollis) distal caudal vertebra (59 mm) (Russell, 1996)
(CMN 41856; paratype of Sigilmassasaurus brevicollis) (brevicollis morphotype) first dorsal vertebra (146 mm) (Russell, 1996)
(CMN 41857; holotype of Sigilmassasaurus brevicollis) (brevicollis morphotype) first dorsal vertebra (121 mm) (Russell, 1996)
(CMN 41858; paratype of Sigilmassasaurus brevicollis) (brevicollis morphotype) second dorsal vertebra (Russell, 1996)
(CMN 41869; bone taxon M) (~1 ton) proximal femur (Russell, 1996)
(CMN 50402; paratype of Sigilmassasaurus brevicollis) ?mid dorsal vertebra (Russell, 1996)
(CMN 50407; paratype of Sigilmassasaurus brevicollis) ?mid or posterior dorsal vertebra (98 mm) (Russell, 1996)
(CMN 50428; paratype of Sigilmassasaurus brevicollis) ?mid dorsal vertebra (Russell, 1996)
(CMN 50790; paratype of Spinosaurus maroccanus) (aegyptiacus morphotype) mid cervical vertebra, two cervical ribs (Russell, 1996)
(CMN 50791; holotype of Spinosaurus maroccanus) (brevicollis morphotype) ~sixth cervical vertebra (195 mm) (Russell, 1996)
(CMN 50794; bone taxon I) manual phalanx I-1 (250 mm) (Russell, 1996)
(CMN 50800; paratype of Sigilmassasaurus brevicollis) ?mid or posterior dorsal vertebra (88 mm) (Russell, 1996)
(CMN 50805; bone taxon I) manual phalanx II-? (111 mm) (Russell, 1996)
(CMN 50813; paratype of Spinosaurus maroccanus) anterior dorsal neural arch (Russell, 1996)
(CMN 50832; paratype of Spinosaurus maroccanus) anterior dentary, tooth (Russell, 1996)
(CMN 50833; paratype of Spinosaurus maroccanus) mid dentary (Russell, 1996)
(Eberharde coll.; cast CMN 41977; bone taxon J) manual ungual (189 mm) (Russell, 1996)
(FSAC-KK 3209) (aegyptiacus morphotype) fused frontals (one partial) (135 mm) (Arden et al., 2019)
(FSAC-KK 3210) (aegyptiacus morphotype) fused frontoparietals (Arden et al., 2019)
(FSAC-KK 7715) (brevicollis morphotype) fused frontals (Arden et al., 2019)
(FSAC-KK 11889) ulna (Ibrahim et al., 2014)
(GMNH-FV 2400) tooth (50x33x30 mm) (Hasegawa et al., 2010)
(GZG.V.19990) tooth (29.8x13x11.5 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.19991) tooth (41.8x15x10.6 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.19992) tooth (28.1x12.9x12 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.19993) tooth (33.9x3.6x11 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.19994) tooth (30.2x11x9.6 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20000) tooth (40x14.5x11.9 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20001) tooth (32.2x14.1x11 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20002) tooth (?x18x13.5 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20003) tooth (19x7.8x7 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20007) tooth (32.1x13.5x11.1mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20010) tooth (30x12.5x9.5 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20011) tooth (37.9x13.4x10.1 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20015) tooth (40.5x15.4x12 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20017) tooth (40x12x15.8 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20018) tooth (30x11x10 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20019) tooth (39.5x17x13.5 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20020) tooth (39x13.9x10.5 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20021) tooth (?x13.6x11.5 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20022) tooth (32.2x11.5x9 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20024) tooth (35.5x13.8x10.8 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20026) tooth (41x12.2x10 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20028) tooth (31x10.5x9.1 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20029) tooth (24.5x11.1x8 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20030) tooth (20.5x11.9x9.2 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20032) tooth (45.9x15.9x12.8 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20033) tooth (54x22.4x17 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20034) tooth (48x17.5x12.1 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20035) tooth (61x21x17.3 mm) (Richter, Mudroch and Buckley, 2013)
(GZG.V.20036) tooth (57.7x21.6x17.3 mm) (Richter, Mudroch and Buckley, 2013)
(IMGP 969-1) maxillary fragment (Buffetaut, 1989)
(IMGP 969-2) ?dentary fragment (Buffetaut, 1989)
(IMGP 969-3) tooth (22 mm) (Buffetaut, 1989)
(JP Cr698) pedal ungual ?II (Singer, 2015 online)
(JP Cr coll.) several teeth (Singer, 2015 online)
(MNHM.KK04) partial ilium (Zitouni, Laurent, Dyke and Jalil, 2019)
(MHNM.KK374) (juvenile; aegyptiacus morphotype) quadrate (78 mm) (Hendrickx, 2006; described in Hendrickx, Mateus and Buffetaut, 2016)
(MHNM.KK375) (subadult; aegyptiacus morphotype) quadrate (145 mm) (Hendrickx, 2006; described in Hendrickx, Mateus and Buffetaut, 2016)
(MHNM.KK376) (brevicollis morphotype) partial quadrate (Hendrickx, 2006; described in Hendrickx, Mateus and Buffetaut, 2016)
(MHNM.KK377) (subadult; aegyptiacus morphotype) quadrate (130 mm) (Hendrickx, 2006; described in Hendrickx, Mateus and Buffetaut, 2016)
(MHNM.KK378) (adult; aegyptiacus morphotype) quadrate (220 mm) (Hendrickx, 2006; described in Hendrickx, Mateus and Buffetaut, 2016)
(MSNM V4047) (brevicollis morphotype) (~17.7 m) (skull ~1.75 m) premaxillae, partial maxillae, partial nasals (Dal Sasso et al., 2005)
(MSNM V6353) pedal phalanx III-1 (Ibrahim et al., 2014)
(MSNM V6849) distal caudal vertebra (Ibrahim et al., 2014)
(MSNM V6872) ~third dorsal vertebra (Ibrahim et al., 2014)
(MSNM V6874) ~tenth cervical vertebra (Ibrahim et al., 2014)
(MSNM V6877) ~tenth cervical vertebra (Ibrahim et al., 2014)
(MSNM V6881) second dorsal vertebra (Ibrahim et al., 2014)
(MSNM V6886) proximal manual phalanx (Ibrahim et al., 2014)
(MSNM V6888) pedal phalanx (Ibrahim et al., 2014)
(MSNM V6893) metacarpal III (Ibrahim et al., 2014)
(MSNM V6896) (subadult; aegyptiacus morphotype) quadrate (145 mm) (Ibrahim et al., 2014; described in Hendrickx, Mateus and Buffetaut, 2016)
(MSNM V6897) pedal phalanx (Ibrahim et al., 2014)
(MSNM V6900) (subadult) ilium (Ibrahim et al., 2014)
(MSNM V7142) distal metatarsal (Ibrahim et al., 2014)
(MSNM V7143) ~first dorsal vertebra (Ibrahim et al., 2014)
(NHMUK R16420) (aegyptiacus morphotype) premaxillae, partial maxillae (Milner, 2003)
(NHMUK R16421) (brevicollis morphotype) incomplete dentary (Milner, 2003)
(NHMUK R16427) (brevicollis morphotype) mid cervical neural arch (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(NHMUK R16429) (aegyptiacus morphotype) incomplete anterior cervical vertebra (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(NHMUK R16434) (brevicollis morphotype) first dorsal vertebra (113 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(NHMUK R16435) (brevicollis morphotype) third dorsal vertebra (150 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(NHMUK R16436) (brevicollis morphotype) second dorsal vertebra (132 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(NHMUK R16665) rostrum (Cuff and Rayfield, 2013)
(NHMUK R36637) (aegyptiacus morphotype) ~eighth cervical centrum (120 mm) (Evers, Rauhut, Milner, McFeeters and Allain, 2015)
(P.P. No. 481) cervical vertebra (D'Anastasio and Capasso, 2004)
(Piccini coll.; cast MPCM 13574) pedal ungual III (120 mm) (Novas et al., 2005)
(SGM-Din 3) (brevicollis morphotype) anterior cervical vertebra (Sereno et al., 1996)
(SGM-Din 4) anterior cervical centrum (Brusatte and Sereno, 2007)
(SGM-Din 5) (brevicollis morphotype) mid cervical vertebra (Brusatte and Sereno, 2007)
(UCPC-2) nasal fragment (Dal Sasso et al., 2005)
(UCRC PV4) (subadult) partial dentary (Ibrahim et al., 2014)
(UCRC PV5) mid caudal vertebra (Ibrahim et al., 2014)
(UCRC PV6) distal caudal vertebra (Ibrahim et al., 2014)
(UCRC PV7) distal caudal vertebra (Ibrahim et al., 2014)
(UCRC PV8) manual phalanx I-1 (Ibrahim et al., 2014)
teeth (Sadleir, 1998)
(private coll.) (multiple individuals) partial skull (1.05 m), partial mandibles (Hendrickx, online)
(private coll.) (~8 m; multiple individuals) partial skull, partial skeleton (Hendrickx, online)
(private coll.) (multiple individuals) premaxillae, quadrate, posterior mandible, cervical vertebrae, caudal vertebrae (Hendrickx, pers. comm.)
(private coll.) premaxillae, dentary (Hendrickx, pers. comm.)
(private coll.) anterior dentary (Hendrickx, pers. comm.)
Barremian-Albian(?), Early Cretaceous
Lower Kem Kem beds(?), Morocco
Material- (CMN 41629) first dorsal vertebra (150 mm) (Russell, 1996)
(CMN 41862) distal caudal vertebra (58 mm) (Russell, 1996)
Albian-Cenomanian, Early-Late Cretaceous
Tegama Group?, Morocco
(MUZ PIG 1647.II.1) tooth (126x34x29 mm) (Niedzwiedzki and Gierlinski, 2002)
(MUZ PIG 1647.II.2) tooth (114x44x30 mm) (Niedzwiedzki and Gierlinski, 2002)
Albian-Cenomanian, Early-Late Cretaceous
Morocco
(LINMH 001) tooth (69x22x17 mm) (Kellner and Mader, 1997)
(LINHM 002) tooth (60x17x13 mm) (Kellner and Mader, 1997)
(UCPC OT6) cervical vertebra (Carrano et al. 2012)
five teeth (Heckeberg, 2009)
Cenomanian, Late Cretaceous
Echkar Formation, Niger
(MNNHN IGU11) mid cervical centrum (90 mm) (Brusatte and Sereno, 2007)
(MNNHN coll.; from In Abangarit) pedal ungual II (90 mm) (Lapparent, 1960)
teeth (Brusatte and Sereno, 2007)
Early Albian, Early Cretaceous
Chenini Formation, Tunisia
(MGGCTUN coll.; Morphotypes 3 and 5) sixteen teeth (17-70 mm) [check Fanti et al. for details]
(Office National des Mines BM231) anterior dentary (Buffetaut and Ouaja, 2002)
teeth (Schluter and Schwarzhans, 1978)
several teeth (Bouaziz et al., 1988)
Diagnosis- (after Carrano et al., 2012) no midline crest on conjoined premaxillae; premaxilla entirely excluded from borders of external naris; extremely elongate dorsal neural spines.
(after Ibrahim et al., 2014) external naris and narial fossa small and retracted near orbit on side of posterior snout; strongly constricted hourglass-shaped and elongated dorsal centra; greatest anteroposterior dorsal neural spine width below spine apex; dorsal neural spines composed of dense bone with narrow central zone of cancellous bone; proximal one-third of dorsal neural spines textured externally by vertical striae; long bones lack open medullary cavity; length of ilium greater than femur; femur strongly bowed anteriorly with fourth trochanter hypertrophied, extending along ~25% of the femoral shaft; pedal digit I long, digit I-1 phalanx longest nonungual phalanx in pes; pedal unguals broader than deep with length almost four times proximal depth; pedal unguals with flat ventral surface.
Comments- The first Spinosaurus material was discovered in 1898, consisting of teeth described by Haug (1905) as the ichthyodectid fish ?Saurocephalus. Schluter and Schwarzhans (1978) mistakenly referred Spinosaurus teeth to Plesiosaurus sp..
The holotype was discovered in 1912 and described by Stromer (1915). IPHG 1922 X45 was originally described as a second, unnamed, species of Spinosaurus by Stromer (1934). It was distinguished from the S. aegyptiacus holotype in part by its low neural spines and called it Spinosaurus B. Both it and the Spinosaurus B specimen were destroyed in 1944 in a British bombing of Munich, and only fragments were described over most of the century after Stromer's description. Several undescribed specimens held in private collections are of note. One is a chimaerical skull auctioned in 2005 and refused by the NHMUK and MNHN. Another is a partial skeleton reconstructed from isolated bones and auctioned in 2009 without a sale.
Le Loeuff and Metais (2010) note in an abstract "A single dinosaur tooth was also discovered in the course of sedimentological exploration for Total Libya. It is a spinosaurid tooth belonging to a spinosaurine close to Spinosaurus..." This is from the Chicla Formation of Libya, called the Kiklah Formation in the abstract.
Rauhut (2009) briefly described and figured a tibia (BSPG 2008 I 68) he assigned to Coelurosauria, but the shape is very similar to Spinosaurus. Zitouni et al. (2019) described ilium MHNM.KK04 as an abelisaurid, but was noted to be tetanurine by Malafaia et al. (2020) and specifically reidentified as Spinosaurus by Smyth et al. (2020).
Spinosaurus marocannus- Russell (1996) described dentary fragments, cervical vertebrae and a dorsal neural arch from the Cenomanian of Morocco as Spinosaurus marocannus. He distinguished this from S. aegyptiacus by the longer mid cervical vertebrae (centrum, not including anterior ball, ~1.5 times height of posterior articular surface). Sereno et al. (1998) rejected its validity because the only character given in the diagnosis to distinguish it from S. aegyptiacus was the longer cervical centra, and they thought this varied within the vertebral column. S. aegyptiacus preserves two cervicals, one of which is probably the third, and the other from the middle of the neck. The latter's centrum has a length to posterior height ratio of 1.05. The two relatively complete cervicals of S. marocannus have ratios of 1.39 and 1.60. Ratios in Baryonyx's holotype range from 1.25-1.81, which is the same amount of variation supposedly separating the Spinosaurus species. Neural spines are comparable in height between Spinosaurus species, and anteroposterior length of the spine varies with centrum elongation as seen in Baryonyx. The latter shows similar differences between cervicals 5 and 6 as seen between S. aegyptiacus and S. marocannus. Russell states besides the elongation noted above and of the neural peduncle (also seen in Baryonyx), the cervicals of the two Spinosaurus species are very similar. The dentaries are also "essentially indistinguishable". Thus, I must agree with Sereno et al. that S. marocannus is a junior synonym of S. aegyptiacus.
Ibrahim et al. 2014- This paper revolutionized our view of Spinosaurus based on the description of a neotype far more complete than any previously published specimen. The neotype was discovered in 2008, and shows Spinosaurus has unusually short hindlimbs and a small pelvis. This and other evidence suggests the taxon was semiaquatic. The presence of a nearly complete specimen allowed the authors to refer other previously known remains to Spinosaurus. This includes the humerus CMN 41852 described by Russell (1996) as theropod bone taxon G and the unguals described by him as theropod bone taxon J. As Russell states the phalanges he calls bone taxon I match the unguals, those are referred to Spinosaurus here too. Russell states this manual material matches an ungual and phalanx (misidentified as a metatarsal) from Algeria referred to Carcharodontosaurus saharicus by Lapparent (1960), so these are also referred to Spinosaurus here. It should be noted that some authors such as Evers et al. (2015) are skeptical the neotype material belongs to one individual or taxon, though a full evaluation must await its detailed description.
Sigilmassasaurus- Russell (1996) suggested Stromer's (1934) Spinosaurus B was referrable to his new genus Sigilmassasaurus. Sereno et al. (1996) later referred the specimen of Spinosaurus B and the material of Sigilmassasaurus to Carcharodontosaurus, based on the supposedly broad cervical vertebra found with the holotype. This was rejected by Novas et al. (2005), who described differences between Sigilmassasaurus cervicals and pedal unguals and those of carcharodontosaurids, so viewed the genus as valid. They also describe the similarity of caudal vertebrae referred to Spinosaurus B and Sigilmassasaurus to Ouranosaurus in their quadrangular shape and elongate posterodorsally projecting neural spines. Canale et al. (2008) proposed these presacral vertebrae belong to iguanodonts, but contra what they write, ornithischians lack pleurocoels and camerate internal structure. Evers et al. (2012) noted Sigilmassasaurus differs from carcharodontosaurids but shares characters with megalosaurs and specifically spinosaurids. McFeeters et al. (2013) were more reserved, placing it in Orionides, though noting shared characters with megalosaurians and some spinosaurids. They referred dorsals except CMN 41858 to Tetanurae indet., and caudals to Dinosauria indet.. Novas et al. (2013) believed similarities between the pedal unguals of Spinosaurus B and Australovenator meant the former might be megaraptoran. The discovery of the nearly complete Spinosaurus neotype by Ibrahim et al. (2014) allowed those authors to solve the mystery of Sigilmassasaurus and Spinosaurus B and synonymize them with Spinosaurus aegyptiacus. They show all of the elements, including pedal unguals and caudals, closely match those of the Spinosaurus neotype. Furthermore, Allain (2014) found the first dorsal vertebra of the possible spinosaurine Ichthyovenator is nearly identical to Sigilmassasaurus, cementing this identification.
D'Anastasio and Capasso (2004) describe pathology in a vertebra they assign to Spinosaurus maroccanus, but which was reassigned to Sigilmassasaurus by McFeeters et al. (2013), and so is now Spinosaurus aegyptiacus. A pedal ungual from the Echkar Formation of Niger referred to Carcharodontosaurus saharicus by Lapparent (1960) matches the Spinosaurus B morphotype, so can now be referred to Spinosaurus. Brusatte and Sereno (2007) describe a cervical centrum from that formation as a paratype of Carcharodontosaurus iguidensis. This centrum resembles Sigilmassasaurus, so can also now be referred to Spinosaurus. A pedal ungual from Alrar referred to Carcharodontosaurus saharicus by Lapparent (1960) matches the Spinosaurus B morphotype, so can now be assigned to Spinosaurus.
Russell suggested CMN 41629 could be from a separate species than the upper Kem Kem taxon, based on differences from the specimens he described as Sigilmassasaurus brevicollis (broader and more flexed cervicodorsal centra; smaller cervicodorsal pleurocoel with inflated centrum wall dorsal to it; more ventrally projected cervicodorsal parapophyses; broader cervicodorsal hypapophysis; less constricted distal caudal centra; smaller distal caudal neural canal). Positional variation is unlikely, as the parapophyses are at the same level as the S. brevicollis holotype. Ontogenetic variation is similarly unlikely, as it is comparable in size with the holotype. Its dark color may indicate it derives from the base of the Kem Kem beds, which may make it Early Cretaceous. He also questionably referred a distal caudal vertebra (CMN 41862) to this species. Russell noted that the Spinosaurus B material was intermediate in cervicodorsal morphology between CMN 41629 and Sigilmassasaurus brevicollis and from a different formation, so believed it to be a potentially intermediate species. Evers et al. (2015) described differences between Sigilmassasaurus and holotypic Spinosaurus vertebrae and between Spinosaurus B and neotypic Spinosaurus limb elements, proposing Sigilmassasaurus (including Spinosaurus B) is a valid taxon. Similarly, Hendrickx (2006) described differences between five Kem Kem spinosaurine quadrates, which was later published as Hendrickx et al. (2016). Notably, the latter were able to compare their quadrates with that of the aegyptiacus neotype, allowing referral of this element to each morphotype. Arden et al. (2019) suggested differences between Kem Kem frontals indicated two taxa, that "FSAC-KK-7115, differs from the smaller specimens [FSAC-KK3209 and 3210] in being proportionately shorter and broader, with more weakly concave orbital margins, more deeply notched postorbital processes, broad overlapping contact of the frontals with the prefrontals, and a taller sagittal crest. The dorsal projection of the orbital margins is more weakly developed as well." Furthermore, "the bone surface of the small FSAC-KK-3209 is dense and avascular whereas that of the larger FSAC-KK-7115 is highly vascularized", which "indicates that the smaller specimens are more mature than the larger one, which was not yet somatically mature." They furthermore suggested that "Two different morphotypes of rostrum are present as well. The larger rostrum morph has been referred to Spinosaurus cf. aegyptiacus (dal Sasso et al., 2005 [MSNM V4047]) but given its size, it may instead pertain to Sigilmassasaurus. A second, smaller spinosaurine rostrum morph (Cuff and Rayfield, 2013 [NHMUK R16665]) is characterized by concave dorsal margin of the rostrum and a more bulbous premaxilla and overall more gracile proportions." Chiarenza and Cau (2016) noted the dentary (NHMUK R16421) described by Milner (2003) differs from the aegyptiacus holotype in "the overall stouter proportion of the bone, in the shape of the alveolar margin, and in the number and placement of the alveoli", which may indicate it belongs to Sigilmassasaurus. While these differences do suggest more than one taxon is represented by Spinosaurus material, the vast majority of specimens could not be assigned to e.g. aegyptiacus vs. brevicollis due to the lack of associated elements, destruction of the aegyptiacus holotype and Spinosaurus B specimen, and lack of description for the aegyptiacus neotype. Given this and the taxonomic uncertainty that would result from the holotype and neotype being potentially different species, all material is here retained as Spinosaurus aegyptiacus pending description of the neotype. Specimens listed above are identified to each morphotype when possible however.
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