Sigilmassasaurus

Sigilmassasaurus (/siːdʒɪlˌmɑːsəˈsɔːrəs/ see-jil-MAH-sə-SOR-əs; "Sijilmassa lizard") is a potentially dubious genus of tetanuran theropod dinosaur that lived approximately 100 to 94 million years ago during the middle of the Cretaceous Period in what is now northern Africa. Sigilmassasaurus was a moderately-built, ground-dwelling, bipedal carnivore, like most other theropods.

Sigilmassasaurus
Temporal range: Late Cretaceous, 100–94 Ma
Sigilmassasaurus vertebra
Middle neck vertebra, specimen CMN 50791
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Spinosauridae
Tribe: Spinosaurini
Genus: Sigilmassasaurus
Russell, 1996
Species:
S. brevicollis
Binomial name
Sigilmassasaurus brevicollis
Russell, 1996
Synonyms

Discovery and naming

Fossils of this dinosaur 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.[1]

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. These rocks date back to the Cenomanian, the earliest stage of the Late Cretaceous Period, approximately 100 to 94 million years ago.[2]

Disputed validity

Neck reconstructions of Sigilmassasaurus and Baryonyx
Neck reconstructions of Sigilmassasaurus (top) and Baryonyx

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 Ernst Stromer as "Spinosaurus B".[3] Russell in 1996 considered this 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 by him 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.[1] 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.[4]

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.[5] A later in 1998 study went further, calling Sigilmassasaurus itself a junior synonym of Carcharodontosaurus.[6]

In 2005, however, 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 clearly differ 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.[7] A study in 2013 confirmed that Sigilmassasaurus was valid and an indeterminate member of the Tetanurae.[8]

Spinosaurus jaw
Spinosaurine snout specimen (MNSM V4047), Milan Natural History Museum

In 2014, Nizar Ibrahim and colleagues referred the specimens of Sigilmassasaurus to Spinosaurus aegyptiacus, together with "Spinosaurus B" as the neotype, and Spinosaurus maroccanus was considered as a nomen dubium following the conclusions of the other papers.[9][10][11] In a 2015 re-description of Sigilmassasaurus by Serjoscha Evers and his team, it was considered a valid genus, belonging to Spinosauridae. This re-description also proposed Spinosaurus maroccanus as a junior synonym of Sigilmassaurus, and rejected the proposal of a Spinosaurus aegyptiacus neotype.[4]

A study by Thomas Arden and colleagues in 2018 concluded 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.[12]

Paleobiology

Diet and Feeding

On the bottoms of its cervical vertebrae, Sigilmassasaurus bore a series of highly rugged bony structures. These were suggested by Evers and colleagues as being possible evidence for substantial neck musculature, since the attachment sites of muscles and ligaments are often indicated by scarring on the bone surface. The neck muscles inferred from Sigilmassasaurus in particular would have enabled it to rapidly snatch fish out of the water, as indicated by the use of similarly-placed musculature in modern birds and crocodilians.[4] This has also been proposed for the related genus Irritator, on account of the prominent sagittal crest running towards the back of its head.[13] However, Evers and colleagues noted that a more thorough biomechanical analysis is required for confirmation of this condition in Sigilmassasaurus.[4]

Paleoecology

Several large theropods (more than one tonne) are known from the Cenomanian of northern Africa, raising questions about how such animals would have coexisted. Species of Spinosaurus, the longest known theropod, has been found in both Morocco and Egypt, as has the huge Carcharodontosaurus. Two smaller theropods, Deltadromeus and Bahariasaurus, have also been found in Morocco and Egypt, respectively, and may be closely related or possibly the same genus. Sigilmassasaurus, from Morocco, and "Spinosaurus B", from Egypt, represent a fourth type of large predator. This situation resembles that in the Late Jurassic Morrison Formation of North America, which boasts up to five theropod genera over one tonne in weight, as well as several smaller genera.[14][15] Differences in head shape and body size among the large North African theropods may have been enough to allow niche partitioning as seen among the many different predator species found today in the African savanna.[16]

References

  1. ^ a b Russell, D.A. (1996). "Isolated Dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco". Bulletin - Museum National d'Histoire Naturelle Section C: Sciences de la Terre. 18: 349–402.
  2. ^ Sereno, PC; Dutheil, DB; Iarochene, M; Larsson, HCE; Lyon, GH; Magwene, PM; Sidor, CA; Varricchio, DJ; Wilson, JA (1996). "Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation". Science. 272 (5264): 986–991. Bibcode:1996Sci...272..986S. doi:10.1126/science.272.5264.986. PMID 8662584.
  3. ^ Stromer, E. (1934). "Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 13. Dinosauria". Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung, Neue Folge (in German). 22: 1–79.
  4. ^ a b c d Evers, S. W.; Rauhut, O. W. M.; Milner, A. C.; McFeeters, B.; Allain, R. (2015). "A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the "middle" Cretaceous of Morocco". PeerJ. 3: e1323. doi:10.7717/peerj.1323. PMC 4614847. PMID 26500829.
  5. ^ Sereno, Paul C.; Dutheil, Didier B.; Iarochene, M.; Larsson, Hans C. E.; Lyon, Gabrielle H.; Magwene, Paul M.; Sidor, Christian A.; Varricchio, David J.; Wilson, Jeffrey A. (1996). "Predatory Dinosaurs from the Sahara and Late Cretaceous Faunal Differentiation". Science. 272 (5264): 986–991. doi:10.1126/science.272.5264.986. ISSN 0036-8075. PMID 8662584.
  6. ^ Sereno, Paul C.; Beck, Allison L.; Dutheil, Didier B.; Gado, Boubacar; Larsson, Hans C. E.; Lyon, Gabrielle H.; Marcot, Jonathan D.; Rauhut, Oliver W. M.; Sadleir, Rudyard W. (1998). "A Long-Snouted Predatory Dinosaur from Africa and the Evolution of Spinosaurids". Science. 282 (5392): 1298–1302. doi:10.1126/science.282.5392.1298. ISSN 0036-8075. PMID 9812890.
  7. ^ Novas, Fernando; de Valais, Silvina; Vickers Rich, Patricia; Rich, Tom (2005). "A large Cretaceous theropod from Patagonia, Argentina, and the evolution of carcharodontosaurids". Die Naturwissenschaften. 92 (5): 226–30. doi:10.1007/s00114-005-0623-3. PMID 15834691.
  8. ^ McFeeters, Bradley; Ryan, Michael J.; Hinic-Frlog, Sanja; Schröder-Adams, Claudia (2013). "A reevaluation of Sigilmassasaurus brevicollis (Dinosauria) from the Cretaceous of Morocco". Canadian Journal of Earth Sciences. 50 (6): 636–649. Bibcode:2013CaJES..50..636M. doi:10.1139/cjes-2012-0129.
  9. ^ dal Sasso, C.; Maganuco, S.; Buffetaut, E.; Mendez, M.A. (2005). "New information on the skull of the enigmatic theropod Spinosaurus, with remarks on its sizes and affinities". Journal of Vertebrate Paleontology. 25 (4): 888–896. doi:10.1671/0272-4634(2005)025[0888:NIOTSO]2.0.CO;2. ISSN 0272-4634.
  10. ^ Ibrahim, Nizar; Sereno, Paul C.; Dal Sasso, Cristiano; Maganuco, Simone; Fabri, Matteo; Martill, David M.; Zouhri, Samir; Myhrvold, Nathan; Lurino, Dawid A. (2014). "Semiaquatic adaptations in a giant predatory dinosaur". Science. 345 (6204): 1613–6. Bibcode:2014Sci...345.1613I. doi:10.1126/science.1258750. PMID 25213375. Supplementary Information
  11. ^ Sereno, P.C.; Beck, A.L.; Dutheil, D.B.; Gado, B.; Larsson, H.C.E.; Lyon, G.H.; Marcot, J.D.; Rauhut, O.W.M.; Sadleir, R.W.; Sidor, C.A.; Varricchio, D.D.; Wilson, G.P; Wilson, J.A. (1998). "A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids". Science. 282 (5392): 1298–1302. Bibcode:1998Sci...282.1298S. doi:10.1126/science.282.5392.1298. PMID 9812890.
  12. ^ Arden, T.M.S.; Klein, C.G.; Zouhri, S.; Longrich, N.R. (2018). "Aquatic adaptation in the skull of carnivorous dinosaurs (Theropoda: Spinosauridae) and the evolution of aquatic habits in Spinosaurus". Cretaceous Research. In Press: 275–284. doi:10.1016/j.cretres.2018.06.013.
  13. ^ Sues, H. D.; Frey, E.; Martill, D. M.; Scott, D. M. (2002). "Irritator challengeri, a spinosaurid (Dinosauria: Theropoda) from the Lower Cretaceous of Brazil". Journal of Vertebrate Paleontology. 22 (3): 535–547. doi:10.1671/0272-4634(2002)022[0535:ICASDT]2.0.CO;2.
  14. ^ M Henderson, Donald (1998). "Skull and tooth morphology as indicators of niche partitioning in sympatric Morrison Formation theropods". Gaia. 15.
  15. ^ Holtz, Thomas; E. Molnar, Ralph; Currie, Philip (2004), "Basal Tetanurae", The Dinosauria: Second Edition, pp. 71–110
  16. ^ Farlow, James O.; Planka, Eric R. (2002). "Body Size Overlap, Habitat Partitioning and Living Space Requirements of Terrestrial Vertebrate Predators: Implications for the Paleoecology of Large Theropod Dinosaurs". Historical Biology. 16 (1): 21–40. doi:10.1080/0891296031000154687. ISSN 0891-2963.
1996 in paleontology

Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 1996.

2018 in archosaur paleontology

The year 2018 in archosaur paleontology was eventful. Archosaurs include the only living dinosaur group — birds — and the reptile crocodilians, plus all extinct dinosaurs, extinct crocodilian relatives, and pterosaurs. Archosaur palaeontology is the scientific study of those animals, especially as they existed before the Holocene Epoch began about 11,700 years ago. The year 2018 in paleontology included various significant developments regarding archosaurs.

This article records new taxa of fossil archosaurs of every kind that have been described during the year 2018, as well as other significant discoveries and events related to paleontology of archosaurs that occurred in the year 2018.

Bahariya Formation

The Bahariya Formation (also transcribed as Baharija Formation) is a fossiliferous geologic formation dating back to the Early Cenomanian, which outcrops within the Bahariya depression in Egypt, and is known from oil exploration drilling across much of the Western Desert where it forms an important oil reservoir.

Baryonyx

Baryonyx () is a genus of theropod dinosaur which lived in the Barremian stage of the Early Cretaceous Period, about 130–125 million years ago. The first skeleton was discovered in 1983 in the Weald Clay Formation of Surrey, England, and became the holotype specimen of B. walkeri, named by palaeontologists Alan J. Charig and Angela C. Milner in 1986. The generic name, Baryonyx, means "heavy claw" and alludes to the animal's very large claw on the first finger; the specific name, walkeri, refers to its discoverer, amateur fossil collector William J. Walker. The holotype specimen is one of the most complete theropod skeletons from the UK (and remains the most complete spinosaurid), and its discovery attracted media attention. Specimens later discovered in other parts of the United Kingdom and Iberia have also been assigned to the genus.

The holotype specimen, which may not have been fully grown, was estimated to have been between 7.5 and 10 m (25 and 33 ft) long and to have weighed between 1.2 and 1.7 t (1.3 and 1.9 short tons). Baryonyx had a long, low, and narrow snout, which has been compared to that of a gharial. The tip of the snout expanded to the sides in the shape of a rosette. Behind this, the upper jaw had a notch which fitted into the lower jaw (which curved upwards in the same area). It had a triangular crest on the top of its nasal bones. Baryonyx had a large number of finely serrated, conical teeth, with the largest teeth in front. The neck formed an S-shape, and the neural spines of its dorsal vertebrae increased in height from front to back. One elongated neural spine indicates it may have had a hump or ridge along the centre of its back. It had robust forelimbs, with the eponymous first-finger claw measuring about 31 cm (12 in) long.

Now recognised as a member of the family Spinosauridae, Baryonyx's affinities were obscure when it was discovered. Some researchers have suggested that Suchosaurus cultridens is a senior synonym (being an older name), and that Suchomimus tenerensis belongs in the same genus; subsequent authors have kept them separate. Baryonyx was the first theropod dinosaur demonstrated to have been piscivorous (fish-eating), as evidenced by fish scales in the stomach region of the holotype specimen. It may also have been an active predator of larger prey and a scavenger, since it also contained bones of a juvenile Iguanodon. The creature would have caught and processed its prey primarily with its forelimbs and large claws. Baryonyx may have had semiaquatic habits, and coexisted with other theropod, ornithopod, and sauropod dinosaurs, as well as pterosaurs, crocodiles, turtles and fishes, in a fluvial environment.

Cajual Island

The island of Cajual is located in the Baía de São Marcos near to Alcântara, Maranhão, Brazil.The island is an important Brazilian paleontological site, where fossils of animals such as Spinosaurus and Sigilmassasaurus were found, and also such plants as conifers and ferns. The finds include the remains of the largest carnivorous dinosaur in Brazil. Traces of the maxillary and nostril of the specimen were found. Known as Oxalaia quilombensis, the species is part of the family of dinosaurs known as Spinosauridae, with elongated skulls and spines along the back.

The presence of fossils also present in Africa proves that the continents were once united, forming Gondwana. The "Lage of the Joker", where the fossils are found, is up to 2 meters below the water at high tide, so it can only be visited for only a few hours a day.

Currently, the island of Cajual is home to a small community of quilombolas.

Cenomanian

The Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding age. Both age and stage bear the same name.

As a unit of geologic time measure, the Cenomanian age spans the time between 100.5 ± 0.9 Ma and 93.9 ± 0.8 Ma (million years ago). In the geologic timescale it is preceded by the Albian and is followed by the Turonian. The Upper Cenomanian starts approximately at 95 M.a.

The Cenomanian is coeval with the Woodbinian of the regional timescale of the Gulf of Mexico and the early part of the Eaglefordian of the regional timescale of the East Coast of the United States.

At the end of the Cenomanian an anoxic event took place, called the Cenomanian-Turonian boundary event or the "Bonarelli Event", that is associated with a minor extinction event for marine species.

Ichthyovenator

Ichthyovenator (meaning "fish hunter") is a genus of spinosaurid theropod dinosaur that lived during the Early Cretaceous of what is now Laos, likely from the Aptian stage (113–125 million years ago). Ichthyovenator is known from fossils collected in the Grès supérieurs Formation. Like other members of its family, it had elongated neural spines forming a sail on its back, although Ichthyovenator's was unusual due to its particular wave-like curvature and being split in two over the hips. Ichthyovenator was initially thought to belong to the Baryonychinae subfamily, but more recent analyses place it in the Spinosaurinae.

Irritator

Irritator is a genus of spinosaurid theropod dinosaur that lived in what is now Brazil during the Albian stage of the Early Cretaceous Period, about 110 million years ago. It is known from a nearly complete skull found in the Romualdo Formation of the Araripe Basin. Fossil dealers had acquired this skull and illegally sold it to the State Museum of Natural History Stuttgart. In 1996, the specimen became the holotype of the type species Irritator challengeri. The genus name comes from the word "irritation", reflecting the feelings of paleontologists who found the skull had been heavily damaged and altered by the collectors. The species name is an homage to the fictional character Professor Challenger from Arthur Conan Doyle's novels.

Many paleontologists regard Angaturama limai—known from a snout tip that was described later in 1996—as a potential junior synonym of Irritator. Both animals hail from the same stratigraphic units of the Araripe Basin. It was also previously proposed that Irritator and Angaturama's skull parts belonged to the same specimen. Although this has been cast into doubt, more overlapping fossil material is needed to confirm whether they are the same animal or not. Other spinosaurid skeletal material, some of which could belong to Irritator or Angaturama, was retrieved from the Romualdo Formation, allowing for a replica skeleton to be made and mounted for display at the National Museum of Rio de Janeiro in 2009.

Estimated at between 6 and 8 meters (20 and 26 ft) in length, Irritator weighed around 1 tonne (1.1 short tons), making it one of the smallest spinosaurids known. Its long, shallow and slender snout was lined with straight and unserrated conical teeth. Lengthwise atop the head ran a thin sagittal crest, to which powerful neck muscles were likely anchored. The nostrils were positioned far back from the tip of the snout, and a rigid secondary palate on the roof of the mouth would have strengthened the jaw when feeding. Belonging to a subadult, Irritator challengeri's holotype remains the most completely preserved spinosaurid skull yet found. The Angaturama snout tip expanded to the sides in a rosette-like shape, bearing long teeth and an unusually tall crest. One possible skeleton indicates it, like other spinosaurids, had enlarged first-finger claws and a sail running down its back.

Irritator had been mistaken initially for a pterosaur, and later a maniraptoran dinosaur. In 1996, the animal was identified as a spinosaurid theropod. The holotype skull was thoroughly prepared before being redescribed in 2002, confirming this classification. Both Irritator and Angaturama belong to the Spinosaurinae subfamily. A generalist diet—like that of today's crocodilians—has been suggested; Irritator might have preyed mainly on fish and any other small prey animals it could catch. Fossil evidence is known of an individual that ate a pterosaur, either from hunting or scavenging it. Irritator may have had semiaquatic habits, and inhabited the tropical environment of a coastal lagoon surrounded by dry regions. It coexisted with other carnivorous theropods as well as turtles, crocodyliforms, and a large number of pterosaur and fish species.

Kem Kem Beds

The Kem Kem Beds (also referred to by various names including the Continental Red Beds and Continental intercalaire) is a geological formation along the border between Morocco and Algeria in southeastern Morocco, whose strata date back to the Late Cretaceous.Dinosaur remains are among the fossils that have been recovered from the formation. Recent fossil evidence in the form of isolated large abelisaurid bones and comparisons with other similarly aged deposits elsewhere in Africa indicates that the fauna of the Kem Kem Beds (specifically in regard to the numerous predatory theropod dinosaurs) may have been mixed together due to the harsh and changing geology of the region when in reality they would likely have preferred separate habitats and likely would be separated by millions of years.

List of African dinosaurs

This is a list of dinosaurs whose remains have been recovered from Africa. Africa has a rich fossil record, but it is patchy and incomplete. It is rich in Triassic and Early Jurassic dinosaurs. African dinosaurs from these time periods include Coelophysis, Dracovenator, Melanorosaurus, Massospondylus, Euskelosaurus, Heterodontosaurus, Abrictosaurus, and Lesothosaurus. In the Middle Jurassic, the sauropods Atlasaurus, Chebsaurus, Jobaria, and Spinophorosaurus, flourished, as well as the theropod Afrovenator. The Late Jurassic is well represented in Africa, mainly thanks to the spectacular Tendaguru Formation. Veterupristisaurus, Ostafrikasaurus, Elaphrosaurus, Giraffatitan, Dicraeosaurus, Janenschia, Tornieria, Tendaguria, Kentrosaurus, and Dysalotosaurus are among the dinosaurs whose remains have been recovered from Tendaguru. This fauna seems to show strong similarities to that of the Morrison Formation in the United States and the Lourinha Formation in Portugal. For example, similar theropods, ornithopods and sauropods have been found in both the Tendaguru and the Morrison. This has important biogeographical implications.

The Early Cretaceous in Africa is known primarily from the northern part of the continent, particularly Niger. Suchomimus, Elrhazosaurus, Rebbachisaurus, Nigersaurus, Kryptops, Nqwebasaurus, and Paranthodon are some of the Early Cretaceous dinosaurs known from Africa. The Early Cretaceous was an important time for the dinosaurs of Africa because it was when Africa finally separated from South America, forming the South Atlantic Ocean. This was an important event because now the dinosaurs of Africa started developing endemism because of isolation.

The Late Cretaceous of Africa is known mainly from North Africa. During the early part of the Late Cretaceous, North Africa was home to a rich dinosaur fauna. It includes Spinosaurus, Carcharodontosaurus, Rugops, Bahariasaurus, Deltadromeus, Paralititan, Aegyptosaurus, and Ouranosaurus.

List of dinosaur genera

This list of dinosaurs is a comprehensive listing of all genera that have ever been included in the superorder Dinosauria, excluding class Aves (birds, both living and those known only from fossils) and purely vernacular terms.

The list includes all commonly accepted genera, but also genera that are now considered invalid, doubtful (nomen dubium), or were not formally published (nomen nudum), as well as junior synonyms of more established names, and genera that are no longer considered dinosaurs. Many listed names have been reclassified as everything from birds to crocodilians to petrified wood. The list contains 1559 names, of which approximately 1192 are considered either valid dinosaur genera or nomina dubia.

Sijilmasa

Sijilmasa (Arabic: سجلماسة‎; also transliterated Sijilmassa, Sidjilmasa, Sidjilmassa and Sigilmassa) was a medieval Moroccan city and trade entrepôt at the northern edge of the Sahara in Morocco. The ruins of the town extend for five miles along the River Ziz in the Tafilalt oasis near the town of Rissani. The town's history was marked by several successive invasions by Berber dynasties. Up until the 14th century, as the northern terminus for the western trans-Sahara trade route, it was one of the most important trade centres in the Maghreb during the Middle Ages.

Spinosauridae

Spinosauridae (meaning "spined lizards") is a family of megalosauroidean theropod dinosaurs. The genus Spinosaurus, from which the family, subfamily, and tribe borrow their names, is the longest terrestrial predator known from the fossil record, and likely reached lengths of 15 m (49 ft). Most spinosaurids lived during the Cretaceous Period, with possible origins in the Late Jurassic, and fossils of them have been recovered worldwide, including in Africa, Europe, South America, Asia, and Australia, although none have been formally named from the latter continent. Spinosaur remains have generally been attributed to the Early to Mid Cretaceous, with the exception of the Ostafrikasaurus from the Late Jurassic.

Spinosaurids were large bipedal carnivores with elongated, crocodile-like skulls lined with conical teeth bearing little to no serrations, and small crests on top of their heads. The teeth in the front end of their lower jaws fanned out into a spoon-shaped structure similar to a rosette, which gave the animal a characteristic look. Their shoulders were robust, prominent and bore stocky forelimbs with giant "hooked" claws on the first finger of their hands. Many genera had unusually tall neural spines on their vertebrae, which supported sails or humps of skin or fat tissue.

Direct fossil evidence and anatomical adaptations indicate that spinosaurids were at least partly piscivorous, with additional fossil finds indicating they also hunted pterosaurs and small to medium-sized dinosaurs. Osteological analyses have suggested a semiaquatic lifestyle for some members of this clade.

Spinosaurus

Spinosaurus (meaning "spine lizard") is a genus of theropod dinosaur that lived in what now is North Africa, during the upper Albian to upper Turonian stages of the Cretaceous period, about 112 to 93.5 million years ago. This genus was known first from Egyptian remains discovered in 1912 and described by German paleontologist Ernst Stromer in 1915. The original remains were destroyed in World War II, but additional material has come to light in the early 21st century. It is unclear whether one or two species are represented in the fossils reported in the scientific literature. The best known species is S. aegyptiacus from Egypt, although a potential second species, S. maroccanus, has been recovered from Morocco.

Spinosaurus was among the largest of all known carnivorous dinosaurs, nearly as large as or even larger than Tyrannosaurus, Giganotosaurus and Carcharodontosaurus. Estimates published in 2005, 2007, and 2008 suggested that it was between 12.6–18 metres (41–59 ft) in length and 7 to 20.9 tonnes (7.7 to 23.0 short tons) in weight. New estimates published in 2014 and 2018 based on a more complete specimen, supported the earlier research, finding that Spinosaurus could reach lengths of 15–16 m (49–52 ft). The latest estimates suggest a weight of 6.4–7.5 tonnes (7.1–8.3 short tons). The skull of Spinosaurus was long and narrow, similar to that of a modern crocodilian. Spinosaurus is known to have eaten fish, and most scientists believe that it hunted both terrestrial and aquatic prey; evidence suggests that it lived both on land and in water as a modern crocodilian does. The distinctive spines of Spinosaurus, which were long extensions of the vertebrae, grew to at least 1.65 meters (5.4 ft) long and were likely to have had skin connecting them, forming a sail-like structure, although some authors have suggested that the spines were covered in fat and formed a hump. Multiple functions have been put forward for this structure, including thermoregulation and display.

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