Spinosaurus

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Spinosaurus
Skeletal reconstruction of Spinosaurus aegyptiacus in a floating position at the National Geographic Society Museum in Washington, DC

Skeletal reconstruction of Spinosaurus aegyptiacus in a floating position at the National Geographic Society Museum in Washington, DC

Temporal occurrence
Lower to Upper Cretaceous (possibly Albium to Cenomanium )
112.9 to 93.9 million years
Locations
Systematics
Dinosaur (dinosauria)
Lizard dinosaur (Saurischia)
Theropoda
Spinosauroidea
Spinosauridae
Spinosaurus
Scientific name
Spinosaurus
Stromer , 1915

Spinosaurus ("thorn lizard") is a genus of theropod dinosaur thatlivedin what is now North Africa during the late Lower and early Upper Cretaceous .

The first bones of this dinosaur were discovered in Egypt in 1912 and described by the German paleontologist Ernst Stromer von Reichenbach in 1915 . These remains were destroyed in World War II - however, other remains have been discovered in recent years. The type and only recognized species today is Spinosaurus aegyptiacus from Egypt and Morocco . Moroccan finds have also been described as Spinosaurus maroccanus . Another synonym is Sigilmassasaurus brevicollis .

Spinosaurus shows conspicuous spinous processes of the vertebrae , which were at least 1.69 meters long and were probably connected with skin to form a sail-like structure. Some authors also suggest that they were surrounded by muscles and formed an elongated hump. The function of this structure remains unclear - among other things, functions for thermoregulation and communication with conspecifics, e.g. B. Discussed the impressive behavior .

Recent estimates suggest that Spinosaurus grew 16 to 18 meters long and weighed 7 to 9 tons when fully grown . It would be the largest known carnivorous dinosaur, ahead of other large theropods such as Tyrannosaurus and Giganotosaurus .

features

Skeletal reconstruction

Because of its size, the eye-catching sail and the crocodile-like skull, Spinosaurus is one of the most popular dinosaurs. In contrast, this genus was mainly known from fossils destroyed during World War II, with the exception of some more recently discovered teeth and skull bones, as well as from a relatively complete find from eastern Morocco that was identified as a neotype .

New jaw and skull material described in 2005 suggests that the skull was about 1.75 meters long - Spinosaurus thus had one of the longest skulls of carnivorous dinosaurs ever. The elongated skull had a narrow muzzle, which was studded with straight, conical and unsawed teeth. In the intermaxillary bone at the front end of the upper jaw there were six to seven teeth on each side, followed by 12 teeth on each side of the maxillary . The second and third teeth on each side were significantly larger than the rest of the intermaxillary teeth, creating a gap between the former and the large teeth of the anterior maxillary - this gap was filled by large teeth in the anterior mandible. The tip of the muzzle that carried these large teeth was enlarged, and a small comb sat on the muzzle in front of the eyes.

The sail (or muscle crest) of Spinosaurus was supported by very high spinous processes (processus spinosi) of the vertebrae. These appendages were seven to eleven times higher than the vertebrae they carried.

Numerous features indicate a semi-aquatic way of life. This includes the small pelvis. The feet resembled those of today's waders. They may also be webbed. The long neck and the relatively long trunk move the body's center of gravity forward. Although Spinosaurus' hind legs were not particularly long, they were sufficient to allow a bipedal (two-legged) walk on land. Most likely Spinosaurus hunted fish like Onchopristis , a large representative of the pseudo- sawfish , especially in shallow water . The strong forelimbs with large, curved claws were used to hold on to slippery prey. The relatively small nostrils were in the middle of the skull and could have enabled Spinosaurus to breathe almost completely submerged, much like crocodiles can. Openings for nerves at the tip of the snout indicate receptors for movement stimuli, which are also found in crocodiles. The tail of Spinosaurus was equipped with high spinous processes and chevron bones and was therefore not round in cross-section, but flattened laterally like the tails of crocodiles and newts. With the help of a robotic hammer in the form of a replica Spinosaurus tail, it was demonstrated that the organ was capable of extensive lateral deflections and was able to generate propulsion while swimming.

Systematics and types

Spinosauridae cladogram
    Spinosauridae     

  ? Suchosaurus


   Baryonychinae     

  Baryonyx


   

  ? Cristatusaurus


   

  Suchomimus


Template: Klade / Maintenance / 3

   Spinosaurinae     

  Irritator


   

  ? Siamosaurus


   

  Oxalaia


   

  Spinosaurus


Template: Klade / Maintenance / 3Template: Klade / Maintenance / 4

Template: Klade / Maintenance / 3

Template: Klade / Maintenance / Style

Spinosaurus is the eponymous representative of the Spinosauridae . Other representatives of this group include Baryonyx from southern England , Irritator and Angaturama (which is probably identical with Irritator ) from Brazil , Suchomimus from Niger , possibly Siamosaurus , which is known from fragmentary bones from Thailand , Oxalaia , another great representative of the Spinosauridae from Brazil and Ichthyovenator from Laos . Spinosaurus is most closely related to Irritator , with which it shares unsawed straight teeth. These two genera are summarized as Spinosaurinae . In 2003, Oliver Rauhut suspected that Stromer's Spinosaurus -type specimen was a chimera composed of the vertebrae of a carcharodontosaurid similar to Acrocanthosaurus and the dentary of a large theropod similar to Baryonyx . However, recent studies reject this hypothesis.

Spinosaurus compared to other large theropods.
Drawing life reconstruction from 2020

So far, two species of Spinosaurus have been described: Spinosaurus aegyptiacus ("Egyptian thorn lizard", from Latin spina "sting, thorn" and Gr. Sauros "lizard") and Spinosaurus maroccanus ("Moroccan thorn lizard"). The species Spinosaurus maroccanus established by Dale Russell differs from Spinosaurus aegyptiacus in the length of the cervical vertebrae. The status of this species is controversial; While some authors consider the length of the vertebrae to be variable between individuals and therefore regard Spinosaurus maroccanus as invalid or as a synonym of Spinosaurus aegyptiacus , other authors maintain the validity of this species.

The spinosaurids Sigilmassasaurus brevicollis and Oxalaia quilombensis are considered synonyms of Spinosaurus by some scientists . Since Spinosaurus was named before the other genera, Sigilmassasaurus and Oxalaia would lose their validity.

Paleoecology and Nutrition

Live plastic reconstructions of Spinosaurus and Mawsonia prey fish in front of the Museu Blau building

The habitat inhabited by Spinosaurus in today's North Africa is only partially understood. The Bahariyya Oasis in Egypt, for example, was probably a mudflat landscape with mangrove forests. Spinosaurus lived there together with the similarly sized predators Bahariasaurus and Carcharodontosaurus , the gigantic titanosaur Paralititan , the smaller titanosaur Aegyptosaurus , the 10 meter long crocodile Stomatosuchus and the 3.5 meter large coelacanth Mawsonia .

It is controversial whether Spinosaurus was a primarily terrestrial (land-based) predator or a fish eater. The fish eater hypothesis is supported by anatomical features such as the elongated jaws, the conical teeth and the nostrils lying above. As early as 1997, Charig and Milner saw a specialized fisherman in the related European genus Baryonyx . Direct evidence for the diet of spinosaurids has been discovered in Baryonyx as well as South American species. In the digestive tract of Baryonyx is etched fish scales and bones found juvenile acids Iguanodon , while a tooth in a pterosaur infected -Bone from South America, indicating that Spinosauriden occasionally these flying archosaurs reenacted. Gregory Paul suspects that Spinosaurus was a Cretaceous equivalent of today's grizzly bear - an opportunistic predator who was adapted to fishing, but still preyed on various small and medium-sized land-based prey and did not spurn carrion.

According to a recent study, Spinosaurus was semi-aquatic, i.e. partially aquatic. The researchers led by Romain Amiot examined the mineral apatite from the teeth of spinosaurids for the ratio between two isotopes of oxygen , oxygen-16 and oxygen-18. The analysis shows a ratio of the two isotopes, as is typically found in animals living in water. The isotope ratio is different in terrestrial and aquatic animals because the body of land animals loses water through evaporation, with the heavier oxygen-18 isotope building up in the body.

A 2018 study by Donald Henderson tested the hypothesis of a semi-aquatic lifestyle using a 3D digital model. The model was found to be unsinkable and unstable when swimming. In addition, the reconstruction indicates a bipedal posture, which is suitable for a terrestrial way of life. On the basis of the isotope analysis mentioned above, the author nevertheless assumes that Spinosaurus spent a lot of time near coasts and in shallow water and mainly fed on fish.

Paleobiology

size

Comparison of large theropods, Spinosaurus in red

Since its discovery, Spinosaurus has been considered one of the largest known theropod dinosaurs. Friedrich von Huene (1926) and Donald F. Glut (1982) estimated this genus to be 6 tons in weight and 15 to 17 meters in length. Gregory S. Paul (1988) also considers Spinosaurus to be the longest theropod, 15 meters in length, but gave a lower weight estimate. More recent estimates by Dal Sasso and colleagues (2005) based on new finds assume a length of 16 to 18 meters and a weight of 7 to 9 tons.

François Therrien and Donald Henderson (2007) compared the skull length with that of the better known theropods and estimate Spinosaurus to be shorter from 12.6 to 14.3 meters and a greater weight of 12 to 20.9 tons. This study was criticized because of the skull reconstruction used and because of the large theropods used for comparison - the latter are mostly tyrannosaurids and carnosaurs , which, however, showed a different physique than spinosaurids.

sail

Spinosaurus had greatly elongated spinous processes of the vertebrae, similar to some other dinosaurs such as the ornithopod Ouranosaurus and the sauropod Amargasaurus . Possibly these spinous processes were connected with skin and thus formed a sail, analogous to the dorsal sails of some pelycosaurs such as Dimetrodon and Edaphosaurus . The function of such a sail on Spinosaurus is unclear. If the sail was crisscrossed with many blood vessels, it could have served - directed across the sun - to absorb heat in the cool morning hours. However, this would mean that the animal was at most partially warm-blooded. A function for cooling is also possible, whereby the sail had to be held away from the sun or at a 90 ° angle to a cold wind. Another hypothesis is a function of the display behavior. Like the tail of the peacock , the sail could have been used for courtship advertising. Stromer speculated that the sail could have been of different sizes in male and female animals.

Jack Bowman Bailey suspects that the appendages did not carry a sail, but a hump, similar to that of today's bison . The spinous processes of Spinosaurus and other dinosaurs are, unlike pelycosaurs, relatively thick. Bailey notes that a sail would have been an efficient heat collector on large theropods, but inefficient as a cooling mechanism - thus a sail would be a disadvantage in the tropical Spinosaurus habitat . According to this researcher, a hump could have served as a heat buffer to protect against heat and as an energy store, which could have allowed long migrations to nesting sites.

Ibrahim and colleagues initially proposed a semi-aquatic way of life for Spinosaurus , with a "display" function for the back sail, which would have always remained above the surface of the water. In 2015, however, Gimsa and colleagues published the hypothesis that Spinosaurus hunted underwater, for which the authors assumed a corresponding hydrodynamic stability. Since it would have warned prey of their attackers, a sail protruding out of the water would have been nonsensical from a behavioral point of view in order to ambush animals on land from their safe hiding place in the water, similar to crocodiles .

For the function of the dorsal sail under water, a function analogous to the dorsal fin of the sailfish is obvious, which, similar to the sword of a sailing ship, stabilizes the body against lateral movements. The shape and relative size of the dorsal sail are similar to those of a fanfish, suggesting that its stabilizing effect improved the precision and strength of lateral tail and spear strokes. As a result, the back sail would have allowed faster and more precise neck and head movements when hunting and improved the stability of the entire body in the water when "wrestling" with larger prey. Gimsa et al. also assume that the stabilizing effect of the back sail under water allowed larger tail amplitudes without destabilizing the body, which would have allowed higher accelerations. In addition to a whole series of adaptations, its skull shape also suggests that Spinosaurus was piscivor .

posture

Although Spinosaurus is traditionally depicted as biped (two-legged), there have been speculations since the mid-1970s that Spinosaurus was at least intermittently quadruped (four-legged). These conjectures were supported with the discovery of Baryonyx , a related genus with sturdy arms. Bailey (1997) considered quadruped locomotion possible because of the extra weight of the hump he hypothesized, which led to corresponding new reconstructions with Spinosaurus as quadruped. In describing the neotype, it was noted that Spinosaurus had shorter hind legs than other theropods and that its center of gravity was in front of the pelvis, so a quadruple was likely to be mandatory. However, recent reconstructions show that the center of gravity was possibly much closer to the pelvis than previously thought.

History of discovery and finds

Fossil of Spinosaurus aegyptiacus

The first skeleton was discovered in 1912 in the Bahariyya oasis in Egypt by Richard Markgraf and scientifically described in 1915 by the German paleontologist Ernst Stromer von Reichenbach (1871–1952) as Spinosaurus aegyptiacus . In 1934, Stromer attributed further fragmentary remains from Bahariyya - vertebrae and hind leg bones - to another species, which he temporarily named " Spinosaurus B"; today, however, these bones are ascribed to either Carcharodontosaurus or Sigilmassasaurus . Some of the Spinosaurus fossils were damaged during the transport from Egypt to the German Museum in Munich - all remaining bones were lost in the Museum Alte Akademie in 1944 in an Allied air raid on Munich .

To date, seven Spinosaurus finds have been described. The remains are dated to an age of around 113 to 94 million years ( Albium to Cenomanium ).

IPHG 1912 VIII 19 , the holotype example described by Stromer in 1915, was destroyed in World War II - but detailed drawings remained. It came from a subadult (not yet fully grown) individual. Estimates assume a length of 14 meters and a weight of 6.7 tons, the skull is estimated at 1.45 meters in length. The skeleton consisted of an upper jaw fragment (maxilla), an incomplete dental (lower jaw), 19 teeth, two incomplete cervical vertebrae, seven vertebrae, dorsal ribs, abdominal ribs (gastralia) and eight caudal vertebrae centers. The longest surviving vertebral process was 1.69 meters long.

Jawbone of Spinosaurus marocannus in the Muséum national d'histoire naturelle , Paris

CMN 50791 comes from the Kem Kem Beds , a formation in southeastern Morocco, and was described by Russell in 1996. Russell attributed the find to a second species, Spinosaurus marocannus . The remains include a 19.5 cm long central cervical vertebra as well as the anterior and central parts of the dentary.

MNHN SAM 124 from Algeria was described by Taquet and Russell in 1998 and consists of a fragmentary intermaxillary bone (premaxillary), a fragmentary upper jaw bone, ploughshare bones and a fragment of the dental bone . Estimates assume a length of 14 meters and a weight of 6.7 tons for this individual; the total length of the skull is estimated to be 1.42 meters.

BM231 is an anterior dental from the Chenini formation from southern Tunisia . Buffetaut and Ouaja attributed this find to Spinosaurus aegyptiacus in 2002 , as the bone shows strong similarities with the corresponding bone of the type material of this species. The find is currently held in the Tunisian Office for Mining (Office National des Mines).

MSNM V4047 from S. aegyptiacus

MSNM V4047 is a skull find that consists of the intermaxillary bone , part of the maxillary and a fragmentary paired nasal bone , which together measure 98.8 cm in length. The find comes from the Kem Kem Beds and was described in 2005 by Cristiano Dal Sasso from the Museo Civico di Storia Naturale di Milano and other scientists.

UCPC-2 is a skull comb described by the same researchers.

FSAC-KK 11888 is a relatively complete find that includes most of the skull, spine, spinous processes supporting the sail, pelvis, and limbs. The find comes from the Kem Kem Beds near Erfoud , was described in 2014 by Paul C. Sereno , Cristiano Dal Sasso and colleagues and established as a neotype .

Other finds consist mainly of very fragmentary remains and isolated teeth. For example, teeth from the Echkar formation from Niger were ascribed to Spinosaurus aegyptiacus . Possible Spinosaurus fossils have been reported from the Turkana Grits of Kenya and from the Cabao Formation of Libya . The latter find comes from the Hauterivian about 130 million years ago, whereby the assignment to Spinosaurus is provisional.

The relatively complete skeletons of the first Spinosaurus found by Ernst Stromer von Reichenbach and the specimen excavated a century later in the Kem Kem Beds complement each other very well. They were summarized in a digital skeleton model and completed with the relatively few missing parts, with the assistance of Nizar Ibrahim , Paul C. Sereno and Cristiano Dal Sasso. This results in a real synthetic resin model. The bizarre appearance of the Spinosaurus can be explained by the fact that the largest predatory dinosaur of all time was the only dinosaur that could also live in water and feed on large fish. So he didn't have to compete on land with the other large dinosaurs for the few herbivores found.

Popular culture

Spinosaurus has long been featured in popular science books about dinosaurs, although enough information for an accurate reconstruction has only recently become available. Older popular illustrations are often based on the influential skeletal reconstruction by Lapparent and Lavocat (1955) and show Spinosaurus as an upright theropod, with a skull similar to that of other large theropods, a sail on the back, and hands with four fingers.

Spinosaurus is the animal antagonist in the movie Jurassic Park III , where he is portrayed as bigger and stronger than Tyrannosaurus . There he is wrongly called "Spinosaurus aegypticus".

Individual evidence

  1. ^ A b Matthew T. Carrano, Roger BJ Benson, Scott D. Sampson: The phylogeny of Tetanurae (Dinosauria: Theropoda). In: Journal of Systematic Palaeontology. Vol. 10, No. 2, 2012, ISSN  1477-2019 , pp. 211-300, doi : 10.1080 / 14772019.2011.630927 .
  2. ↑ a b c d e f g Cristiano Dal Sasso , Simone Maganuco, Eric Buffetaut , Marco A. Mendez: The phylogeny of Tetanurae (Dinosauria: Theropoda). In: Journal of Vertebrate Paleontology. Vol. 10, No. 2, 2012, ISSN  0272-4634 , pp. 888-896, doi : 10.1671 / 0272-4634 (2005) 025 [0888: NIOTSO] 2.0.CO; 2 .
  3. ↑ a b Robert SH Smyth; Nizar Ibrahim; David M. Martill: Sigilmassasaurus is Spinosaurus : a reappraisal of African spinosaurines . In: Cretaceous Research . 2020. doi: 10.1016 / j.cretres.2020.104520 .
  4. ↑ a b c d e f g h Nizar Ibrahim, Paul C. Sereno, Cristiano Dal Sasso, Simone Maganuco, Matteo Fabbri, David M. Martill, Samir Zouhri, Nathan Myhrvold & Dawid A. Iurino. Semiaquatic adaptations in a giant predatory dinosaur. Science , 11 September 2014 DOI: 10.1126 / science.1258750
  5. ↑ Ralph E. Molnar, Sergei M. Kurzanov, Zhiming Dong : Carnosauria. In: David B. Weishampel , Peter Dodson , Halszka Osmólska (eds.): The Dinosauria . University of California Press, Berkeley CA et al. 1990, ISBN 0-520-06727-4 , pp. 169-209.
  6. ↑ Ibrahim, N., Maganuco, S., Dal Sasso, C. et al .: Tail-propelled aquatic locomotion in a theropod dinosaur . In: Nature . April 29, 2020, doi : 10.1038 / s41586-020-2190-3 (English).
  7. ↑ a b c Thomas R. Holtz Jr. , Ralph E. Molnar, Philip J. Currie : Basal Tetanurae. In: David B. Weishampel, Peter Dodson, Halszka Osmólska (eds.): The Dinosauria. 2nd edition. University of California Press, Berkeley CA et al. 2004, ISBN 0-520-24209-2 , pp. 71-110.
  8. ↑ Oliver WM Rauhut : The interrelationships and evolution of basal theropod dinosaurs (= Special Papers in Palaeontology. Vol. 6). The Palaeontological Association, London 2003, ISBN 0-901702-79-X .
  9. ^ A b Dale A. Russell : Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco. In: Bulletin du Muséum National d'Histoire Naturelle. Section C: Sciences de la Terre, paleontology, geology, mineralogy. 4. Série, Vol. 18, No. 2/3, 1996, ISSN  0181-0642 , pp. 349-402.
  10. ^ A b Paul C. Sereno , Allison L. Beck, Didier B. Dutheil, Boubacar Gado, Hans CE Larsson, Gabrielle H. Lyon, Jonathan D. Marcot, Oliver WM Rauhut, Rudyard W. Sadleir, Christian A. Sidor, David D. Varricchio, Gregory P. Wilson, Jeffrey A. Wilson: A Long-Snouted Predatory Dinosaur from Africa and the Evolution of Spinosaurids. In: Science . Vol. 282, No. 5392, 1998, pp. 1298-1302, doi : 10.1126 / science.282.5392.1298 , PMID 9812890 .
  11. ↑ Joshua B. Smith, Matthew C. Lamanna, Kenneth J. Lacovara, Peter Dodson, Jennifer R. Smith, Jason C. Poole, Robert Giegengack, Yousry Attia: A Giant Sauropod Dinosaur from an Upper Cretaceous Mangrove Deposit in Egypt. In: Science. Vol. 292, No. 5522, 2001, pp. 1704-1706, doi : 10.1126 / science.1060561 .
  12. ^ Alan J. Charig , Angela C. Milner : Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. In: Bulletin of the Natural History Museum. Geology Series. Vol. 53, No. 1, 1997, ISSN  0968-0462 , pp. 11-70, digitized .
  13. ↑ Eric Buffetaut, David Martill, François Escuillié: Pterosaurs as part of a spinosaur diet. In: Nature . Vol. 430, No. 6995, 2004, p. 33, doi : 10.1038 / 430033a .
  14. ↑ Gregory S. Paul : Family Spinosauridae. In: Gregory S. Paul: Predatory Dinosaurs of the World. Simon & Schuster, New York NY et al. 1988, ISBN 0-671-61946-2 , pp. 271-274.
  15. ↑ Romain Amiot, Eric Buffetaut, Christophe Lécuyer, Xu Wang, Larbi Boudad, Zhongli Ding, François Fourel, Steven Hutt, François Martineau, Manuel Alfredo Medeiros, Jinyou Mo, Laurent Simon, Varavudh Suteethorn, Steven Sweetman, Haiyan Tong, Fusong Zhang, Zhonghe Zhou : Oxygen isotope evidence for semi-aquatic habits among spinosaurid theropods. In: Geology. Vol. 38, No. 2, 2010, ISSN  0091-7613 , pp. 139-142, doi : 10.1130 / G30402.1 .
  16. ↑ a b Henderson, DM (2018). A buoyancy, balance and stability challenge to the hypothesis of a semi-aquatic Spinosaurus Stromer, 1915 (Dinosauria: Theropoda) . PeerJ. 6: e5409. doi: 10.7717 / peerj.5409
  17. ^ Friedrich von Huene : The carnivorous saurischia in the Jura and Cretaceous formations principally in Europe. In: Revista del Museo de La Plata. Vol. 29, 1926, ISSN  0375-1147 , pp. 35-167.
  18. ^ Donald F. Glut: The New Dinosaur Dictionary. Citadel Press, Secaucus NJ 1982, ISBN 0-8065-0782-9 , pp. 226-228.
  19. ^ Gregory S. Paul: Predatory Dinosaurs of the World. Simon & Schuster, New York NY et al. 1988, ISBN 0-671-61946-2 .
  20. ^ François Therrien, Donald M. Henderson: My theropod is bigger than yours ... or not: estimating body size from skull length in theropods. In: Journal of Vertebrate Paleontology. Vol. 27, No. 1, 2007, pp. 108-115, doi : 10.1671 / 0272-4634 (2007) 27 [108: MTIBTY] 2.0.CO; 2 .
  21. ↑ Mickey Mortimer: Comments on Therrien and Henderson's new paper. Dinosaur Mailing List, March 25, 2007, accessed July 4, 2007 .
  22. ^ Lambert B. Halstead: The Evolution and Ecology of the Dinosaurs. Eurobook Limited, London 1975, ISBN 0-85654-018-8 .
  23. ^ A b c Ernst Stromer : Results of Prof. E. Stromer's research trips in the deserts of Egypt. Volume 2: Vertebrate remains of the Baharîje stage (lowest Cenoman). Part 3: The original of the theropod Spinosaurus aegyptiacus nov. gen., nov. spec. (= Treatises of the Bavarian Academy of Sciences. Mathematical-Physical Class. Vol. 28, 3, ISSN  0176-7038 = Memoranda of the Bavarian Academy of Sciences. Vol. 89, 3). Publishing house of the Bavarian Academy of Sciences, Munich 1915.
  24. ↑ a b Jack Bowman Bailey: Neural spine elongation in dinosaurs: sailbacks or buffalo-backs? In: Journal of Paleontology. Vol. 71, No. 6, 1997, ISSN  0022-3360 , pp. 1124-1146, abstract .
  25. ↑ a b Jan Gimsa, Robert Sleigh, Ulrike Gimsa: The riddle of Spinosaurus aegyptiacus' dorsal sail . In: Geological Magazine . tape 153 , no. 3 , November 2015, ISSN  0016-7568 , p. 544-547 , doi : 10.1017 / S0016756815000801 ( cambridge.org [accessed May 18, 2020]).
  26. ↑ a b c Donald F. Glut: Spinosaurus. In: Donald F. Glut: Dinosaurs. The Encyclopedia. 1st supplement. McFarland & Company, Jefferson NC 2000, ISBN 0-7864-0591-0 , pp. 329-333.
  27. ^ Ernst Stromer: Results of Prof. E. Stromer's research trips in the deserts of Egypt. Volume 2: Vertebrate remains of the Baharîje stage (lowest Cenoman). Part 13: Dinosauria (= treatises of the Bavarian Academy of Sciences. Mathematical and Natural Science Department. NF Vol. 22, ISSN  0005-6995 ). Publishing house of the Bavarian Academy of Sciences, Munich 1934.
  28. ↑ Mickey Mortimer: Megalosauroidea. (No longer available online.) In: The Theropod Database. University of Washington, 2004, archived from the original December 19, 2013 ; Retrieved August 14, 2008 . Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. @1@ 2Template: Webachiv / IABot / archosaur.us
  29. ↑ Eric Buffetaut, Mohamed Ouaja: A new specimen of Spinosaurus (Dinosauria, Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the evolutionary history of the Spinosauridae. In: Bulletin de la Société Géologique de France. Vol. 173, No. 5, 2002, ISSN  0037-9409 , pp. 415-421, doi : 10.2113 / 173.5.415 .
  30. ↑ Stephen Brusatte, Paul Sereno: A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and its implications for allosauroid phylogeny. Journal of Vertebrate Paleontology. Vol. 25, Supplement 3, 2005, p. 40A, digitized version (PDF; 14.68 MB) ( Memento of the original from July 26, 2014 in the Internet Archive ) Info: The archive link was inserted automatically and not yet checked. Please check the original and archive link according to the instructions and then remove this notice. . @1@ 2Template: Webachiv / IABot / vertpaleo.org
  31. ^ David B. Weishampel: Dinosaur distribution. In: David B. Weishampel, Peter Dodson, Halszka Osmólska (eds.): The Dinosauria. University of California Press, Berkeley CA et al. 1990, ISBN 0-520-06727-4 , pp. 63-139.
  32. ^ Fabio M. Dalla Vecchia: Second record of a site with dinosaur skeletal remains in Libya (northern Africa). In: Natura Nascosta. Vol. 11, 1995, ISSN  1590-9522 , pp. 16-21.
  33. ↑ http://www.zdf.de/ZDFmediathek#/beitrag/video/2257788/Expedition-Supersaurier
  34. ^ Graham Chandler: A bite-size guide to the dinosaurs of the new movie Jurassic Park III. In: National Geographic World. No. 312, August 2001, ZDB -ID 2109805-0 , p. 25.

Web links

Commons : Spinosaurus  - collection of images, videos and audio files