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Spinosaurus aegyptiacus

Reptilia - Spinosauridae

Taxonomy

Spinosaurus aegyptiacus was named by Stromer (1915). Its type specimen is BSP 1915, a partial skeleton, and it is a 3D body fossil. Its type locality is Gebel el Dist (7p), which is in a Cenomanian marginal marine sandstone/sandstone in the Baharîje Formation of Egypt.

Synonyms

Oxalaia quilombensis was named by Kellner et al. (2011). Its type specimen is MN 6117-V, a partial skull (Fused premaxillae), and it is a 3D body fossil. Its type locality is Laje do Coringa, Ilha do Cajual, which is in a Cenomanian estuary/bay sandstone/conglomerate in the Alcântara Formation of Brazil.

It was synonymized subjectively with Spinosaurus aegyptiacus by Smyth et al. (2020).
Spinosaurus marrocanus was named by Russell (1996). Its type specimen is NMC 50791, a set of vertebrae, and it is a 3D body fossil. Its type locality is Tafilalt region (NMC/CMN), which is in an Albian/Cenomanian fluvial-lacustrine sandstone in Morocco.

It was corrected as Spinosaurus maroccanus by Fanti et al. (2014); it was synonymized subjectively with Sigilmassasaurus brevicollis by Evers et al. (2015), Arden et al. (2019) and Lakin and Longrich (2019); it was considered a nomen dubium by Sereno et al. (1998); it was considered a nomen dubium by Buffetaut and Ouaja (2002) and Rauhut (2003); it was considered a nomen dubium by Kellner et al. (2011) and Ibrahim et al. (2014); it was misspelled as Spinosaurus maroccanus by Hendrickx et al. (2016); it was synonymized subjectively with Spinosaurus aegyptiacus by Taquet (2010), Cavin et al. (2010), Carrano et al. (2012), Smyth et al. (2020) and Ibrahim et al. (2020).
Sigilmassasaurus brevicollis was named by Russell (1996). Its type specimen is NMC 41857 and is a 3D body fossil. Its type locality is Tafilalt region (NMC/CMN), which is in an Albian/Cenomanian fluvial-lacustrine sandstone in Morocco.

It was synonymized subjectively with Carcharodontosaurus saharicus by Sereno et al. (1998) and Taquet (2010); it was synonymized subjectively with Spinosaurus aegyptiacus by Ibrahim et al. (2014), Smyth et al. (2020) and Ibrahim et al. (2020); it was considered a nomen dubium by Candeiro et al. (2011).

Synonymy list

Year	Name and author
1915	Spinosaurus aegyptiacus Stromer p. 28 figs. Pl. 1-2
1923	Spinosaurus aegyptiacus Huene p. 457 fig. 3
1926	Spinosaurus aegyptiacus Huene p. 89
1927	Spinosaurus aegyptiacus Huene p. 266
1928	Spinosaurus aegyptiacus Depéret and Savornin p. 264
1934	Spinosaurus aegyptiacus Stromer p. 8
1936	Spinosaurus aegyptiacus Stromer p. 64
1939	Spinosaurus aegyptiacus Kuhn p. 77
1957	Spinosaurus aegyptiacus Lapparent p. 109
1963	Spinosaurus aegyptiacus Furon p. 154
1964	Spinosaurus aegyptiacus Tatarinov p. 538
1967	Spinosaurus aegyptiacus Charig p. 714
1968	Spinosaurus aegypticus Maleev p. 95
1970	Spinosaurus aegyptiacus Steel p. 38
1970	Spinosaurus aegyptiacus Swinton p. 143
1974	Spinosaurus aegyptiacus Langston p. 84
1974	Spinosaurus aegyptiacus Molnar p. 1014
1980	Spinosaurus aegyptiacus Molnar and Pledge p. 285 fig. 3
1984	Spinosaurus aegypticus Taquet p. 81
1984	Spinosaurus aegyptiacus Taquet p. 217
1986	Spinosaurus aegyptiacus Bonaparte p. 74
1986	Spinosaurus aegyptiacus Chabli p. 71
1988	Spinosaurus aegyptiacus Bouaziz et al. pp. 336-338
1988	Spinosaurus aegyptiacus Paul p. 273
1989	Spinosaurus aegyptiacus Buffetaut p. 81
1990	Spinosaurus aegyptiacus Buffetaut et al. p. 192
1990	Spinosaurus aegyptiacus Molnar et al. p. 192
1991	Spinosaurus aegyptiacus Molnar p. 166
1992	Spinosaurus aegyptiacus Buffetaut
1996	Spinosaurus aegyptiacus Kellner p. 614
1996	Spinosaurus aegyptiacus Kellner and Campos p. 155
1996	Spinosaurus marrocanus Russell pp. 355-356 figs. 4-8
1996	Sigilmassasaurus brevicollis Russell pp. 361-362
1997	Spinosaurus aegyptiacus Le Loeuff p. 454
1997	Spinosaurus aegyptiacus Milner p. 699
1997	Spinosaurus marrocanus Milner p. 699
1997	Spinosaurus aegyptiacus Novas p. 685
1997	Spinosaurus aegyptiacus Vickaryous and Ryan p. 488
1998	Spinosaurus aegyptiacus Sereno et al. p. 1302
1998	Spinosaurus marrocanus Taquet and Russell p. 348
2002	Spinosaurus aegyptiacus Buffetaut and Ouaja p. 418
2002	Spinosaurus aegyptiacus Sues et al. p. 543
2003	Spinosaurus aegyptiacus Rauhut p. 34
2004	Spinosaurus aegyptiacus Holtz, Jr. et al. p. 73
2004	Spinosaurus marrocanus Holtz, Jr. et al. p. 73
2004	Sigilmassasaurus brevicollis Holtz, Jr. et al. p. 74
2005	Spinosaurus aegyptiacus Dal Sasso et al. p. 888
2005	Sigilmassasaurus brevicolis Novas et al. p. 169
2005	Sigilmassasaurus brevicollis Novas et al. p. 229
2006	Spinosaurus aegyptiacus Candeiro et al. p. 936
2006	Spinosaurus aegyptiacus Medeiros p. 335
2007	Spinosaurus aegyptiacus Buffetaut
2007	Sigilmassasaurus brevicollis Medeiros et al. p. 418
2007	Spinosaurus marrocanus Regagba et al. p. 149
2007	Spinosaurus aegyptiacus Sánchez-Hernández et al. p. 190
2009	Spinosaurus aegyptiacus Lü et al. p. 50
2010	Spinosaurus aegyptiacus Buffetaut p. 175
2010	Spinosaurus aegyptiacus Cavin et al. p. 399
2010	Spinosaurus aegyptiacus Hasegawa et al. p. 12
2010	Spinosaurus aegyptiacus Lü et al. p. 386
2010	Spinosaurus aegyptiacus Taquet p. 96
2011	Spinosaurus aegyptiacus Gianechini et al. p. 14
2011	Sigilmassasaurus brevicollis Hone et al. p. 502
2011	Spinosaurus aegyptiacus Hone et al. p. 502
2011	Oxalaia quilombensis Kellner et al. p. 100 figs. 1-4
2011	Spinosaurus aegyptiacus Kellner et al. p. 101
2011	Spinosaurus aegyptiacus Mateus et al. p. 54
2011	Spinosaurus aegyptiacus Vila et al. p. 157
2011	Spinosaurus aegyptiacus Wongko et al. p. 62
2012	Oxalaia quilombensis Candeiro et al. p. 223
2012	Spinosaurus aegyptiacus Carrano et al. p. 243
2012	Spinosaurus aegyptiacus Le Loeuff et al. p. 489
2012	Oxalaia quilombensis Lindoso et al. p. 119
2012	Spinosaurus aegyptiacus Richter et al.
2013	Spinosaurus aegyptiacus Bertozzo et al. p. 120
2013	Oxalaia quilombensis Fernandes de Azevedo et al. p. 131
2013	Oxalaia quilombensis Furtado et al. p. 106
2013	Sigilmassasaurus brevicollis McFeeters et al. p. 636
2013	Sigilmassasaurus brevicollis Novas et al. p. 186
2013	Spinosaurus aegyptiacus Richter et al. p. 302
2014	Spinosaurus aegyptiacus Fanti et al. p. 40
2014	Spinosaurus maroccanus Fanti et al. p. 40
2014	Spinosaurus aegyptiacus Ibrahim et al. p. S9
2015	Spinosaurus aegyptiacus Benyoucef et al. p. 52
2015	Spinosaurus aegyptiacus Evers et al. p. 2
2015	Sigilmassasaurus brevicollis Evers et al. p. 8–9
2016	Sigilmassasaurus brevicollis Chiarenza and Cau p. 10
2016	Spinosaurus aegyptiacus Chiarenza and Cau p. 10
2016	Spinosaurus maroccanus Hendrickx et al. p. 2
2016	Spinosaurus aegyptiacus Hendrickx et al. p. 12
2016	Sigilmassasaurus brevicollis Hendrickx et al. p. 15
2016	Spinosaurus aegyptiacus Serrano-Martínez et al. p. 403
2017	Spinosaurus aegyptiacus Ezcurra and Agnolin p. 485
2018	Oxalaia quilombensis Aranciaga Rolando et al. p. 107
2018	Spinosaurus aegyptiacus Aureliano et al. p. 283
2018	Spinosaurus aegyptiacus Gasca et al. p. 141
2018	Sigilmassasaurus brevicollis Hassler et al. p. 2
2018	Spinosaurus aegyptiacus Hassler et al. p. 2
2019	Spinosaurus aegyptiacus Arden et al. p. 275
2019	Sigilmassasaurus brevicollis Arden et al. p. 276
2019	Oxalaia quilimboensis Arden et al. p. 279
2019	Sigilmassasaurus brevicollis Lakin and Longrich p. 129
2019	Spinosaurus aegyptiacus Lakin and Longrich p. 130
2020	Spinosaurus aegyptiacus Ibrahim et al. p. 9
2020	Oxalaia quilombensis Malafaia et al. p. 432
2020	Sigilmassasaurus brevicollis Malafaia et al. p. 432
2020	Spinosaurus aegyptiacus Malafaia et al. p. 432
2020	Oxalaia quilombensis Pereira et al. p. 1
2020	Spinosaurus aegyptiacus Smyth et al. p. 3
2021	Spinosaurus aegyptiacus McFeeters p. 182
2021	Oxalaia quilombensis de França et al. p. 1817
2021	Spinosaurus aegyptiacus de França et al. p. 1817
2022	Spinosaurus aegyptiacus Mateus and Estraviz-López p. 2
2022	Spinosaurus aegyptiacus Salem et al. p. 2
2022	Sigilmassasaurus brevicollis Salem et al. p. 9

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Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Deuterostomia
phylum	Chordata	Haeckel 1874
subphylum	Vertebrata
superclass	Gnathostomata
—	Osteichthyes	()
subclass	Sarcopterygii	()
subclass	Dipnotetrapodomorpha	(Nelson 2006)
subclass	Tetrapodomorpha	()
—	Tetrapoda
—	Reptiliomorpha
—	Anthracosauria
subclass	Amphibiosauria	Kuhn 1967
—	Cotylosauria	()
—	Amniota
—	Sauropsida
class	Reptilia
subclass	Eureptilia	()
—	Romeriida

Rank	Name	Author
—	Diapsida	()
—	Archosauromorpha	(Huene 1946)
—	Crocopoda
—	Archosauriformes	Gauthier 1986
—	Eucrocopoda
—	Archosauria	()
informal	Avemetatarsalia
—	Ornithodira
—	Dinosauromorpha
—	Dinosauriformes
—	Dinosauria	()
—	Theropoda	()
—	Neotheropoda
—	Averostra	Paul 2002
—	Tetanurae
superfamily	Megalosauroidea	()
family	Spinosauridae
subfamily	Spinosaurinae
genus	Spinosaurus
species	aegyptiacus

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

†Spinosaurus aegyptiacus Stromer 1915

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Invalid names: Oxalaia quilombensis Kellner et al. 2011 [synonym], Sigilmassasaurus brevicollis Russell 1996 [synonym], Spinosaurus marrocanus Russell 1996 [synonym]

Diagnosis

Reference	Diagnosis
D. A. Russell 1996 (Sigilmassasaurus brevicollis)	As for genus; for species-level characters see under "Description" and "Discussion", Sigilmassasaurus sp., below.
A. W. A. Kellner et al. 2011 (Oxalaia quilombensis)	Spinosaurid with the following combination of characters that distinguish it from other members of this clade (autapomorphies are marked with an asterisk): teeth unserrated (differing from Cristatusaurus and Suchomimus); 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; presence of two replacement teeth associated with the 3rd functional tooth; diastema between the 5th and 6th premaxillary teeth present but shorter than in Spinosaurus; ventral portion of the premaxillae very sculptured
M. T. Carrano et al. 2012	Spinosaurid with: (1) no midline crest on conjoined premaxillae (Dal Sasso et al. 2005); (2) premaxilla entirely excluded from borders of external naris (Dal Sasso et al. 2005); and (3) extremely elongate dorsal neural spines (Stromer 1915).
B. McFeeters et al. 2013 (Sigilmassasaurus brevicollis)	Articular surfaces of cervical centra much wider than high, with width/height ratio approximately 1.5 (versus 1.0–1.25 in most theropods; Russell 1996); interzygapophyseal laminae absent, so that the short neural spine contacts the dorsal margin of the neural canal anteriorly and posteriorly.
N. Ibrahim et al. 2014	Spinosaurid with adult body length ~15 m characterized by the following cranial features: external naris and narial fossa small and retracted near the orbit on the side of the posterior snout; premaxilla excluded from the border of the external naris. Distinguishing postcranial features include strongly constricted hourglass-shaped and elongated dorsal centra; dorsal neural spine height up to ten times greater than centrum height; greatest anteroposterior dorsal neural spine width below spine apex; dorsal neural spines composed of dense bone with a 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 larger than dorsoventral length of 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 the pes; pedal unguals broader than deep with length almost four times proximal depth; pedal unguals with flat ventral surface.
S. W. Evers et al. 2015 (Sigilmassasaurus brevicollis)	Very large spinosaurid theropod dinosaur. The taxon can be diagnosed on the basis of the following autapomorphies: mid-cervical vertebrae with offset, transversely convex, strongly rugose triangular platform at the posterior end of the ventral side that is confluent with a ventral keel anteriorly; anteriorly broad centroprezygapophyseal lamina with no or strongly reduced centroprezygapophyseal fossa already in anterior mid-cervical vertebrae; reduced neural arch lamination with no or incomplete distinction between anterior and posterior centrodiapophyseal laminae in posterior cervicals and first dorsal; small elongate fossa on either side of the base of the neural spine in last cervical and first dorsal vertebrae. Furthermore, Sigilmassasaurus brevicollis differs from most other theropods in the combination of the following characters: anterior articular surface of posterior cervical and anterior dorsal vertebrae more than 1.5 times wider than high and wider than length of centrum (also in Ichthyovenator); well-developed anterior tubercle present on the anterior articular surface in posterior cervical and anterior dorsal vertebrae; interzygapophyseal laminae absent in posterior cervicals and anteriormost dorsals, resulting in ventrally open spinopre- and spinopostzygapophyseal fossae (also in Ichthyovenator); posterior cervical and anterior dorsal vertebrae with massive transverse processes with deeply penetrating pneumatic openings at the base anteriorly and posteriorly (also in D1 in Ichthyovenator); epipophyses weakly developed in mid-cervicals and absent in posterior cervicals; posterior cervical and anteriormost dorsal vertebrae with anteroposteriorly short, posteriorly inclined, low and spike-like neural spines (modified from Russell, 1996; McFeeters, 2011; Evers, 2012; McFeeters et al., 2013).
R. S. H. Smyth et al. 2020	From Ibrahim et al. (2014), except where otherwise noted. Spinosaurid with adult body length ~15 m characterized by the following cranial features: external naris and narial fossa small and retracted near the orbit on the side of the posterior snout; premaxilla excluded from the border of the external naris. Distinguishing postcranial axial features include mid-cervical vertebrae with offset, transversely convex, strongly rugose triangular platform at the posterior end of the ventral side that is confluent with a ventral keel anteriorly (autapomorphic); reduced neural arch lamination with no or incomplete distinction between anterior and posterior centrodiapophyseal laminae in posterior cervicals and first dorsal; small elongate fossa on either side of the base of the neural spine in last cervical and first dorsal vertebrae (after Evers et al., 2015). Strongly constricted hourglass-shaped and elongated dorsal centra; dorsal neural spine height up to ten times greater than centrum height (autapomorphic); greatest anteroposterior dorsal neural spine width below spine apex; dorsal neural spines composed of dense bone with a thin central zone of cancellous bone (autapomorphic); proximal one-third of dorsal neural spines textured externally by vertical striae. Amphicoelous caudal vertebrae; centra possess a longitudinal groove on the ventral surface; neural spine height over seven times greater than centrum height in distal vertebrae; base of mid-distal neural spines flattened proximodistally (autapomorphic); spinodiapophyseal laminae of caudal vertebrae are the most developed laminae (autapomorphic); mid-distal caudal vertebrae with reduced zygapophyses; mid-distal chevrons similar to proximal ones in size and shape (chevrons do not shorten, maintain a slight curve to the rear, and are not increasingly hooklike), (after Ibrahim et al., 2020b). Appendicular autapomorphic features include: long bones lack conspicuous medullary cavity; length of ilium larger than dorsoventral length of femur; femur strongly bowed anteriorly with fourth trochanter hypertrophied, extending along ~25% of the femoral shaft; pedal digit I long, digit I-1st phalanx longest non-ungual phalanx in the pes; pedal unguals broader than deep with length almost four times proximal depth; pedal unguals with flat ventral surface.