Sigilmassasaurus

''Sigilmassasaurus'' ( ; "Sijilmassa lizard") is an extremely controversial genus of spinosaurid dinosaur that lived approximately 100 to 94 million years ago during the Late Cretaceous Period in what is now northern Africa. Named in 1996 by Canadian paleontologist Dale Russell, it contains a single species, ''Sigilmassasaurus brevicollis''. The identity of the genus has been debated by scientists, with some considering its fossils to represent material from the closely related species ''Spinosaurus aegyptiacus'', while others have classified it as a separate taxon, forming the clade Spinosaurini with ''Spinosaurus'' as its sister taxon.

''Sigilmassasaurus'' was a moderately-built, ground-dwelling, bipedal carnivore, like most other theropods. It may have had strong neck musculature as evidenced by the morphology of its vertebrae (backbones). ''Sigilmassasaurus'' may have had semiaquatic habits and a partially piscivorous diet. It coexisted with other large theropods in the Kem Kem Group.

History of research

Fossils of ''Sigilmassasaurus'' were recovered at the Kem Kem Formation in the Tafilalt Oasis region of Morocco, near the site of the ancient city of Sijilmassa, for which it was named. Canadian paleontologist Dale Russell named ''Sigilmassasaurus'' in 1996, from the ancient city and the Greek word ''sauros'' ("lizard"). A single species was named, ''S. brevicollis'', which is derived from the Latin ''brevis'' ("short") and ''collum'' ("neck"), because the neck vertebrae are very short from front to back. ''Sigilmassasaurus'' comes from red sandstone sediments in southern Morocco, which are known by various names, including the ''Grès rouges infracénomaniens'', Continental Red Beds, and lower Kem Kem Beds. The rocks date back to the Cenomanian, the earliest stage of the Late Cretaceous Period, approximately 100 to 94 million years ago.

The holotype, or original specimen, of ''S. brevicollis'', CMN 41857, is a single posterior neck vertebra, although Russell referred about fifteen other vertebrae found in the same formation to the species. Other material had been found in Egypt, and was referred to by German paleontologist Ernst Stromer as "''Spinosaurus'' B". Russell in 1996 considered the Egyptian specimen, IPHG 1922 X45, to belong to ''Sigilmassasaurus'' or a closely related animal, naming it as a ''Sigilmassasaurus'' sp. A second ''Sigilmassasaurus ''sp. was named by Russel based on specimen CMN 41629, an anterior dorsal vertebra. "''Spinosaurus'' B" would be intermediate in build between this latter ''Sigilmassasaurus'' sp. and ''S. brevicollis''. Russell created the family Sigilmassasauridae for these animals. The neck vertebrae of these dinosaurs are wider from side to side, about 50%, than they are long from front to back. Whether the neck as a whole was particularly short, is unknown: the holotype vertebra is a cervicodorsal, from the transition between the neck and the back, which would not be long anyway. The exact position of ''Sigilmassasaurus'' within the theropod family tree is unknown, but it belongs somewhere inside the theropod subgroup known as Tetanurae and most likely was a member of the family Spinosauridae.

Disputed validity

The validity of ''Sigilmassaurus'', however, did not go unchallenged shortly after it was named. In 1996, Paul Sereno and colleagues described a ''Carcharodontosaurus'' skull (SGM-Din-1) from Morocco, as well as a neck vertebra (SGM-Din-3) which resembled that of "''Spinosaurus'' B," which they therefore synonymized with ''Carcharodontosaurus.'' A 1998 study went further, calling ''Sigilmassasaurus'' itself a junior synonym of ''Carcharodontosaurus.''

In 2005, however, Argentine paleontologist Fernando Novas and colleagues found that SGM-Din-3, which was used to synonymize ''Carcharodontosaurus'' and "''Spinosaurus'' B", was not actually associated with SGM-Din-1, the ''Carcharodontosaurus'' skull described in 1996, and shows clear differences with the holotype of ''Carcharodontosaurus''. Other features of "''Spinosaurus'' B" also differed from ''Carcharodontosaurus'', lending support to the notion that it (and therefore ''Sigilmassasaurus'') is a separate taxon. The same study claimed that the tail vertebrae by Russell assigned to the species were in fact those of iguanodonts. A study in 2013 by Bradley McFeeters and colleagues considered ''Sigilmassasaurus'' as valid and an indeterminate member of the Tetanurae.

In 2014, German-Moroccan paleontologist Nizar Ibrahim and colleagues referred the specimens of ''Sigilmassasaurus'' to ''Spinosaurus aegyptiacus'', together with "''Spinosaurus'' B" and created a neotype for ''S. aegyptiacus.'' ''Spinosaurus maroccanus'' was considered a ''nomen dubium'' following the conclusions of previous papers.Supplementary Information
/ref> In a 2015 re-description of ''Sigilmassasaurus'' by Serjoscha Evers and his team, it was considered a valid genus within the Spinosauridae. The authors also proposed ''Spinosaurus maroccanus'' as a junior synonym of ''Sigilmassasaurus'', and rejected the proposal of a ''Spinosaurus aegyptiacus'' neotype.

A study by British paleontologist Thomas Arden and colleagues in 2018 concluded that ''Sigilmassasaurus'' was a valid genus and formed a tribe with ''Spinosaurus'' termed Spinosaurini. The largest specimen of ''Spinosaurus cf. aegyptiacus'', MSNM V4047, was tentatively assigned to ''S. brevicollis''. On the basis of vertebrae, the researchers suggested that ''Sigilmassasaurus'' may have grown larger than ''Spinosaurus''. Although in the absence of associated material, it is difficult to be certain what material belongs to which genus. Below is a cladogram based on the analysis by Arden and colleagues:

A 2020 study by British paleontologist Symth and colleagues suggested that ''Sigilmassasaurus'' is synonymous with ''Spinosaurus'' and the Brazilian spinosaurine genus ''Oxalaia'', with both genera falling into the ''Spinosaurus'' hypodigm