Paralititan

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Paralititan
Temporal range: Late Cretaceous, 99.6–93.5  Ma
Paralititan stromeri by Hatem Moushir 9.JPG
Humeri (upper arm bones) of Paralititan at the Egyptian Geological Museum
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Dinosauria
Clade: Saurischia
Clade:† Sauropodomorpha
Clade:† Sauropoda
Clade:† Macronaria
Clade:† Titanosauria
Family:† Saltasauridae
Subfamily:† Saltasaurinae
Genus:† Paralititan
Smith et al., 2001
Type species
†Paralititan stromeri
Smith et al., 2001

Paralititan (meaning "tidal giant") is an extinct genus of titanosaurian sauropod dinosaur that lived in present-day Egypt during the Late Cretaceous period. It was described by American paleontologist Joshua B. Smith and colleagues in 2001. The genus contains a single species, Paralititan stromeri, named based on an incomplete postcranial skeleton, including vertebrae and limb bones. These fossils were unearthed by an American expedition to the Bahariya Oasis in western Egypt in rock layers coming from the Bahariya Formation. This formation dates to the Cenomanian stage of the Late Cretaceous period, between 99.6 and 93.5 million years ago. Additionally, an incomplete dorsal (back) vertebra that had been described by German paleontologist Ernst Stromer in 1932 was assigned to Paralititan. However, this vertebra was destroyed during the Bombing of Munich in World War II.

Contents

Paralititan was at one point considered one of the largest dinosaurs known, however more recent estimates place it at 27 meters (89 feet) in length and 30 tonnes (33 short tons) in mass. This would make it a large titanosaur, but not as huge as genera like Argentinosaurus and Puertasaurus. Its humerus (upper arm bone) is large, robust, and bears a large deltopectoral crest, with the total length 1.69 meters (5.5 feet). The caudal (tail) vertebrae are wider than tall and lack pleurocoels (large cavities that stored air sacs) on the sides of their centra (bodies of the vertebrae).

Paralititan is a member of the family Saltasauridae, a group of titanosaurs with long necks, large bodies, and small heads. Some saltasaurids like Saltasaurus itself preserve dorsal osteoderms (scutes), however these have not yet been found in Paralititan. Being a titanosaur, Paralititan was likely a four-legged herbivore. Paralititan coexisted with many other dinosaurs including the other sauropod Aegyptosaurus , the theropods Tameryraptor , Spinosaurus , and Bahariasaurus , and an unnamed abelisaurid theropod. During the Cenomanian, the Bahariya Formation was on the border of the Tethys Sea, causing it to be a large network of mangrove swamps, rivers, and tidal flats. In these waters existed a host of giant fishes, like the bichir Bawitius , sawskate Onchopristis , and the plethodid Paranogmius , and crocodylomorphs, like the stomatosuchid Stomatosuchus .

Discovery

In 1999, researcher Joshua Smith rediscovered a site in the Bahariya Oasis of Egypt, known as Gebel el Dist, that had formerly been explored by Austro-Hungarian paleontologist Richard Markgraf between 1912 and 1914. [1] [2] [3] These fossils, including those of dinosaurs like Spinosaurus and Tameryraptor , had been shipped to Germany where they were described by German paleontologist Ernst Stromer. [4] [5] However, apparently Markgraf had already removed all more complete skeletons, leaving only limited remains behind. In 2000, an American expedition was mounted to revisit the site, however it failed in finding any dinosaur skeletons. At a new site, the nearby Gebel Fagga, the expedition succeeded in locating a partial sauropod skeleton. [6] [3]

Geological map of the Bahariya Oasis, where fossils of Paralititan were unearthed Geological map of the Bahariya Oasis.PNG
Geological map of the Bahariya Oasis, where fossils of Paralititan were unearthed

This incomplete skeleton consists of two sacral (probably the 5th and 6th) vertebrae, the first caudal (tail) vertebra, a proximal caudal vertebra, dorsal and sacral ribs, incomplete scapulae, a complete right and incomplete left humerus, the distal (away from body) end of a metacarpal, and several additional elements. All of these fossils were then transported to the Egyptian Geological Museum, Cairo and deposited under catalog number CGM 81119. Gebel Fagga and Gebel el Dist are sandstone outcrops of the Bahariya Formation, dating to the Cenomanian stage of the Late Cretaceous period. This makes these fossils around 93.5 to 99 million years old. [3]

In 2001, an international crew made up of paleontologists Joshua B. Smith, Matthew C. Lamanna, Kenneth J. Lacovara, Peter Dodson, Jennifer R. Smith, Jason Charles Poole, Robert Giegengack and Yousri Attia described the partial skeleton as belonging to a new genus and species of sauropod dinosaur, named Paralititan stromeri. They established this skeleton, CGM 81119, as the holotype (name-bearing) specimen. The generic name, Paralititan, derives from the Greek para, meaning "near", halos "sea", and titan, as in the Greek Titans. The specific name is in honor of Stromer, who first established the presence of dinosaur fossils in the Bahariya Oasis during a 1911 expedition. [1] [2] The whole name therefore translates to "Stromer's tidal titan" or "Stromer's tidal giant", in reference to the paralic tidal flats the dinosaur lived on. [3]

Additionally, Smith and colleagues assigned a large anterior (front) dorsal (back) vertebra, cataloged at the Paläontologisches Museum München (Bavarian State Collection of Paleontology) under SNSB-BSPG 1912V11164, to Paralititan stromeri. This dorsal vertebra was originally described by Stromer in 1932 as belonging to an undetermined "Giant Sauropod", [7] but it was destroyed alongside other fossils from the Bahariya Formation during the Bombing of Munich in World War II. [2] [3] Several isolated sauropod fossils, mainly consisting of caudal vertebrae, were referred by Stromer to the other Bahariya sauropod Aegyptosaurus , [7] but these may belong to Paralititan instead. Paralititan represents the first tetrapod reported from the Bahariya Formation since American paleontologist Alfred Romer's publication of 1935. [3]

Description

Hypothetical scale diagram comparing Paralititan to some humans, with some of the known material in white. Paralititan-Scale-Diagram-SVG-Steveoc86.svg
Hypothetical scale diagram comparing Paralititan to some humans, with some of the known material in white.

Size

Only 5.1% of Paralititan's skeleton is known, [8] so its exact size is difficult to estimate. However, the limited material, especially the long humeri, suggested that it is one of the most massive dinosaurs ever discovered, with an estimated weight of 59 t (65 short tons). [9] Joshua Smith, who informally led the research team that found Paralititan, told an interviewer, "It was a truly enormous dinosaur by any reckoning." [10] The complete right humerus measured 1.69 meters (5.5 feet) long which at the time of discovery was the longest known in a Cretaceous sauropod; this was surpassed in 2016 with the discovery of the other titanosaur Notocolossus which had a 1.76 m (5 ft 9 in) humerus. [11] Using Saltasaurus as a guide, American paleontologist Kenneth Carpenter estimated its length at around 26 m (85 ft) in 2006. [12]

Life restoration Paralititan stromeri.jpg
Life restoration

American researcher Scott Hartman estimates an animal that is massive, but still smaller than the biggest titanosaurs such as Puertasaurus , Alamosaurus , and Argentinosaurus . [13] In 2010, American researcher Gregory S. Paul estimated its length at 20+ meters (66+ ft), and its weight at 20 tonnes (24.2 short tons). [14] In 2012 Holtz gave a length of 32 meters (105 feet) and an estimated weight of 65.3–72.5 tonnes (72–80 short tons). [15] In 2016, using equations that estimate body mass based on the circumference of the humerus and femur of quadrupedal animals, it was given an estimated weight of ~50 t (55 short tons). [11] In 2019, Paul estimated Paralititan between 30 and 55 tonnes (33–60.6 short tons). [16] In 2020, researchers Rubén Molina-Pérez and Asier Larramendi estimated the size of the animal at 27 meters (89 feet) and 30 tonnes (33 short tons). [17]

Postcrania

The sacral vertebrae of Paralititan lack pleurocoels , large spaces in vertebrae that stored pneumatic air sacs. [3] These air sacs made vertebrae lighter and were part of a larger air system in dinosaur bodies. [18] [19] The absence of pleurocoels on the caudal vertebrae is common amongst titanosaurids, supporting its classification in the group. The first caudal vertebra has a wider than tall, procoelous centrum with a convex distal (away from body) articular condyle. In contrast to other titanosaurs like Alamosaurus and Neuquensaurus , the centrum is not biconvex in Paralititan. The ventral (bottom) surface of the centrum has weakly pronounced ridges that border a sagittal concavity. The second proximal caudal vertebra found has a strongly procoelus centrum with a well-developed distal condyle. As for the appendicular bones, the scapula has a medial (towards right) concavity that is bordered by a prominent rugosity, like in the titanosaurs Aeolosaurus, Lirainosaurus, Neuquensaurus, and Saltasaurus. As in the brachiosaurid Brachiosaurus , Paralititan bears a prominent tabular process on the caudoventral (back bottom) part of the scapula underneath the glenoid process. This

Fossils of Paralititan Paralititan stromeri by Hatem Moushir 3.JPG
Fossils of Paralititan

feature is only found in Brachiosaurus and Paralititan, demonstrating that it is a unique genus of titanosaur. [3]

As for the limb bones, Paralititan's humerus is huge at 1.69 meters (5.5 feet) in height, with greatly expanded proximal (toward body) and distal ends. The proximal region of the cranial (exposed) surface of the humerus bears a muscular depression. The deltopectoral crest is large, stretching for more than 53% of the length of the humerus. On the proximocaudal (towards body-back) surface of the distal end of the humerus is a large fossa that is bordered by two large ridges, a medial and a lateral one. The lateral ridge in particular expands into a tuberosity relatively close to the proximal margin of the humerus, as in Saltasaurus and Lirainosaurus, whereas the medial ridge, one unique to Paralititan, is relatively shorter and does not develop into a tuberosity. Lengthy supracondylar ridges for articulation extend over 1/3rd of the humerus' length and border an extensive olecranon fossa, like in other titanosaurs. The metacarpal known from Paralititan has a rectangular, flattened surface indicative phalangeal reduction or absence on this digit. From the formation another sauropod had already been known, Aegyptosaurus. Paralititan differs from Aegyptosaurus in its larger size, possibly in not having pleurocoels in its front tail vertebrae, and in possessing a relatively longer deltopectoral crest on its humerus. [3]

Classification

Paralititan belonged to the group Lithostrotia, a group of sauropods that existed during the Cretaceous period in every continent except for Australia. [20] [3] [21] This group includes a wide array of sauropods, including some of the largest dinosaurs known to ever live such as Argentinosaurus, Patagotitan, and Puertasaurus. Notocolossus and Alamosaurus were comparable in size to Paralititan, being 24 m (79 ft) [22] and 26 metres (85 ft) long respectively. [16] [23] The position of Paralititan within Lithostrotia has changed; Smith and colleagues (2001) considered Paralititan a member of Titanosauridae, a now unused clade name, alongside Saltasaurus, Malawisaurus, Opisthocoelicaudia, Epachthosaurus, and Alamosaurus. [3] However, many later studies did not analyze the phylogenetic placement of Paralititan, likely on account of its fragmentary nature. [24] In studies by American paleontologist Kristina Curry Rogers (2005) [25] and British paleontologists Philip D. Mannion and Paul Upchurch (2011) [26] , Paralititan was recovered as a basal titanosaurian in their phylogenetic analyses. Paralititan was also found to be distantly related to other African taxa like Malawisaurus and Rukwatitan, making the relationships of Paralititan relative to those of other African titanosaurs convoluted. [24]

In a 2022 phylogenetic analysis, Spanish paleontologist Bernat Villa and colleagues recovered Paralititan in the family Saltasauridae, a group of Asian, African, European, North American, and South American lithostrotian titanosaurs that lived during the Cretaceous period. [27] [28] [29] [30] Saltasaurids, like other sauropods, were herbivorous with small heads, long necks, and large bodies. [31] [32] Some saltasaurids like Saltasaurus had dorsal osteoderms on their backs, [33] though other saltasaurids like Paralititan have not been discovered with these osteoderms. [3] Within Saltasauridae, Paralititan was a member of Saltasaurinae, a subfamily that is divided in two clades: Saltasaurini, endemic to South America, and an Afro-European clade formed by Paralititan and Abditosaurus . The clade containing both the Afro-European and South American clades is backed by two synapomorphies: divided humeral distal condyles and caudal centra that are wider than tall, both are characteristics are found in Paralititan. [30]

Below is a cladogram by Villa et al. (2022), from the description of the European saltasaurine Abditosaurus. [30]

Saltasauridae

Palaeoenvironment and taphonomy

Restoration of Paralititan with contemporaneous animals of the Bahariya Formation Bahariya Formation McAfee.jpg
Restoration of Paralititan with contemporaneous animals of the Bahariya Formation

The autochthonous, scavenged skeleton was preserved in tidal flat deposits containing in the form of fossil leaves and root systems, a mangrove vegetation of seed ferns, Weichselia reticulata. The bone layer of water where the Paralititan fossils were found was likely shallow with low currents based on the presence of plant roots and fine-grained sediments in the matrix. Additionally, the associated nature of the holotype and presence of a shallow, vegetated tidal flat indicates that the holotype individual walked over to the site where the fossils were found before its death. An isolated tooth referred to Carcharodontosaurus , [3] a genus formerly thought to be present in the Bahariya Formation but now referred to Tameryraptor, [4] was found associated with the skeleton. Smith and colleagues (2001) stated that this is evidence of the holotype being scavenged by a carnivorous dinosaur as the tooth is larger than the clasts that could be transported in the tidal flat channels, indicating that the Carcharodontosaurus individual moved towards the site and lost its tooth there. [3]

This mangrove system bordered the Tethys Sea, which transformed the region into a mangrove-dominated coastal environment filled with vast tidal flats and waterways. [34] Paralititan is the first dinosaur demonstrated to have inhabited a mangrove habitat. Alongside the earlier named Aegyptosaurus, Paralititan is one of the few sauropods known from the Bahariya Formation, alongside the coeval theropods Bahariasaurus, Tameryraptor, and Spinosaurus, the latter being also known from the Kem Kem beds. A Paralititan-sized titanosaur is known from the Kem Kem Beds, however it is only represented by fragmentary remains. [35] [36] The faunal composition of both the Bahariya Formation and the Kem Kem Beds were thought to be similar in the past, but the describers of Tameryraptor suggested that such superficial comparisons require further examination. [4] Contemporary abelisaurid dinosaurs from the Bahariya Formation were also terrestrial carnivores, preying on other terrestrial fauna. [37] A diverse fauna of aquatic animals is known from the Bahariya Formation. Underwater life diversity exploded during this period in the mangroves of North Africa, with turtles represented by the pleurodian Apertotemporalis , large bony fish like Mawsonia [38] and Paranogmius , [39] sawskates Onchopristis and Schizorhiza , [40] sharks like Squalicorax and Cretolamna, and a broad selection of invertebrates. [41] Additionally, several crocodylomorphs like the stomatosuchid Stomatosuchus [42] and the eunotosuchian Libycosuchus are known from the formation. [43] [44] [45]

References

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  2. 1 2 3 Smith, J.B.; Lamanna, M.C.; Mayr, H.; and Lacovara, K.J. (2006). "New information regarding the holotype of Spinosaurus aegyptiacus Stromer, 1915". Journal of Paleontology. 80 (2): 400–406. doi:10.1666/0022-3360(2006)080[0400:NIRTHO]2.0.CO;2. S2CID   130989487.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Smith JB, Lamanna MC, Lacovara KJ, Dodson P, Smith JR, Poole JC, Giegengack R, Attia Y (June 2001). "A giant sauropod dinosaur from an Upper Cretaceous mangrove deposit in Egypt" (PDF). Science. 292 (5522): 1704–6. Bibcode:2001Sci...292.1704S. doi:10.1126/science.1060561. PMID   11387472. S2CID   33454060.
  4. 1 2 3 Kellermann, Maximilian; Cuesta, Elena; Rauhut, Oliver W. M. (January 14, 2025). "Re-evaluation of the Bahariya Formation carcharodontosaurid (Dinosauria: Theropoda) and its implications for allosauroid phylogeny". PLOS ONE. 20 (1): e0311096. Bibcode:2025PLoSO..2011096K. doi: 10.1371/journal.pone.0311096 . ISSN   1932-6203. PMC   11731741 . PMID   39808629.{{cite journal}}: CS1 maint: article number as page number (link)
  5. ↑ Stromer, Ernst (1931). "Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharîjestufe (unterstes Cenoman). 10. Ein Skelett-Rest von Carcharodontosaurus nov. gen" [Results of Prof. E. Stromer's research trips in the deserts of Egypt. II. Vertebrate remains from the Baharîje stage (lower Cenomanian). 10. A skeletal remains of Carcharodontosaurus nov. gen.](PDF). Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung. Neue Folge (in German). 9. Translated by Carrano, Matthew: 1–31.
  6. ↑ Nothdurft, William; Joshua Smith; Matt Lamana; Ken Lacovara; Jason Poole & Jen Smith, 2002, The Lost Dinosaurs of Egypt: The Astonishing and Unlikely True Story of One of the Twentieth Century's Greatest Paleontological Discoveries, Random House, 272 pp
  7. 1 2 Stromer, E. (1932a). Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltierreste der Baharîje-Stufe (unterstes Cenoman). 11. Sauropoda. Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung, Neue Folge,10: 1-21.
  8. ↑ Lacovara, Kenneth J.; Lamanna, Matthew C.; Ibiricu, Lucio M.; Poole, Jason C.; Schroeter, Elena R.; Ullmann, Paul V.; Voegele, Kristyn K.; Boles, Zachary M.; Carter, Aja M.; Fowler, Emma K.; Egerton, Victoria M.; Moyer, Alison E.; Coughenour, Christopher L.; Schein, Jason P.; Harris, Jerald D. (September 4, 2014). "A Gigantic, Exceptionally Complete Titanosaurian Sauropod Dinosaur from Southern Patagonia, Argentina". Scientific Reports. 4 (1) 6196. Bibcode:2014NatSR...4.6196L. doi:10.1038/srep06196. ISSN   2045-2322. PMC   5385829 . PMID   25186586.
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  28. ↑ Navarro, Bruno A.; Ghilardi, Aline M.; Aureliano, Tito; Díaz, Verónica Díez; Bandeira, Kamila L. N.; Cattaruzzi, André G. S.; Iori, Fabiano V.; Martine, Ariel M.; Carvalho, Alberto B.; Anelli, Luiz E.; Fernandes, Marcelo A.; Zaher, Hussam (September 15, 2022). "A New Nanoid Titanosaur (Dinosauria: Sauropoda) from the Upper Cretaceous of Brazil". Ameghiniana. 59 (5): 477. Bibcode:2022Amegh..59..477N. doi:10.5710/amgh.25.08.2022.3477. ISSN   0002-7014.
  29. ↑ Carrano, Matthew T.; D’Emic, Michael D. (January 2, 2015). "Osteoderms of the titanosaur sauropod dinosaur Alamosaurus sanjuanensis Gilmore, 1922". Journal of Vertebrate Paleontology. 35 (1): e901334. Bibcode:2015JVPal..35E1334C. doi:10.1080/02724634.2014.901334. ISSN   0272-4634.{{cite journal}}: CS1 maint: article number as page number (link)
  30. 1 2 3 Villa, B.; Sellés, A.; Moreno-Azanza, M.; Razzolini, N.L.; Gil-Delgado, A.; Canudo, J.I.; Galobart, À (2022). "A titanosaurian sauropod with Gondwanan affinities in the latest Cretaceous of Europe". Nature Ecology & Evolution. 92 (3): 288–296. Bibcode:2022NatEE...6..288V. doi:10.1038/s41559-021-01651-5. PMID   35132183. S2CID   246650381.
  31. ↑ Powell, J. E. (1992). Osteología de Saltasaurus Loricatus (Sauropoda-Titanosauridae) del Cretácico Superior argentino. In Los dinosaurios y su entorno biótico: II Curso de Paleontología, 10 a 12 de julio de 1990. Actas (pp. 165-230). Instituto Juan de Valdés.
  32. ↑ Henderson, Donald M. (October 30, 2013). "Sauropod Necks: Are They Really for Heat Loss?". PLOS ONE. 8 (10): e77108. Bibcode:2013PLoSO...877108H. doi: 10.1371/journal.pone.0077108 . ISSN   1932-6203. PMC   3812985 . PMID   24204747.{{cite journal}}: CS1 maint: article number as page number (link)
  33. ↑ Cerda, Ignacio A.; Powell, Jaime E. (2010). "Dermal Armor Histology ofSaltasaurus loricatus,an Upper Cretaceous Sauropod Dinosaur from Northwest Argentina". Acta Palaeontologica Polonica. 55 (3): 389–398. Bibcode:2010AcPaP..55..389C. doi:10.4202/app.2009.1101. ISSN   0567-7920.
  34. ↑ Wanas, Hamdalla A.; Assal, Ehab M. (2021). "Provenance, tectonic setting and source area-paleoweathering of sandstones of the Bahariya Formation in the Bahariya Oasis, Egypt: An implication to paleoclimate and paleogeography of the southern Neo-Tethys region during Early Cenomanian". Sedimentary Geology. 413 105822. Bibcode:2021SedG..41305822W. doi:10.1016/j.sedgeo.2020.105822.
  35. ↑ Ibrahim, Nizar; Dal Sasso, Cristiano; Maganuco, Simone; Fabbri, Matteo; Martill, David M.; Gorscak, Eric; Lamanna, Matthew C. (2016). "Evidence of a derived titanosaurian (Dinosauria, Sauropoda) in the 'Kem Kem beds' of Morocco, with comments on sauropod paleoecology in the Cretaceous of Africa". Cretaceous Period: Biotic Diversity and Biogeography. New Mexico Museum of Natural History and Science Bulletin. 71: 149–159.
  36. ↑ Ibrahim, Nizar; Sereno, Paul C.; Varricchio, David J.; Martill, David M.; Dutheil, Didier B.; Unwin, David M.; Baidder, Lahssen; Larsson, Hans C. E.; Zouhri, Samir; Kaoukaya, Abdelhadi (April 21, 2020). "Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco". ZooKeys (928): 1–216. Bibcode:2020ZooK..928....1I. doi: 10.3897/zookeys.928.47517 . ISSN   1313-2970. PMC   7188693 . PMID   32362741.
  37. ↑ Salem, Belal S.; Lamanna, Matthew C.; O'Connor, Patrick M.; El-Qot, Gamal M.; Shaker, Fatma; Thabet, Wael A.; El-Sayed, Sanaa; Sallam, Hesham M. (2022). "First definitive record of Abelisauridae (Theropoda: Ceratosauria) from the Cretaceous Bahariya Formation, Bahariya Oasis, Western Desert of Egypt". Royal Society Open Science. 9 (6) 220106. Bibcode:2022RSOS....920106S. doi: 10.1098/rsos.220106 . PMC   9174736 . PMID   35706658.
  38. ↑ Allam, Ahmed M. (January 1, 1986). "A regional and paleoenvironmental study on the Upper Cretaceous deposits of the Bahariy Oasis, Libyan Desert, Egypt". Journal of African Earth Sciences (1983). 5 (4): 407–412. Bibcode:1986JAfES...5..407A. doi:10.1016/0899-5362(86)90055-2. ISSN   0731-7247.
  39. ↑ Weiler, Wilhelm (1935). "Ergebnisse der Forschungsreisen Prof. Stromers in den Wusten Aegyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes Cenoman). 16. Neue Untersuchungen an den Fischresten" [Results of Prof. Stromer's research trips to the deserts of Egypt. II. Vertebrate remains of the Baharije stage (lowest Cenomanian). 16. New studies on the fish remains](PDF). Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Abteilung (in German). 32: 1–57.
  40. ↑ Slaughter, Bob H. (1974). "A lower Cenomanian (Cretaceous) ichthyofauna from the Bahariya Formation of Egypt". Annals of the Geological Survey of Egypt. 4: 25–40.
  41. ↑ Salem, Belal S. (2023). Geological and paleontological studies on new pterosaur and crocodyliform fossils from the Upper Cretaceous (Cenomanian) Bahariya Formation, Bahariya Oasis, Egypt (MS). Ohio University.
  42. ↑ Stromer, E. (1925). Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 7. Stomatosuchus inermis Stromer, ein schwach bezahnter Krokodilier und 8. Ein Skelettrest des Pristiden Onchopristis numidus Haug sp. Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung 30(6): 1–22.
  43. ↑ Holliday, Casey M.; Gardner, Nicholas M. (2012). "A new eusuchian crocodyliform with novel cranial integument and its significance for the origin and evolution of Crocodylia". PLOS ONE. 7 (1): e30471. Bibcode:2012PLoSO...730471H. doi: 10.1371/journal.pone.0030471 . ISSN   1932-6203. PMC   3269432 . PMID   22303441.{{cite journal}}: CS1 maint: article number as page number (link)
  44. ↑ Stromer, E. (1933). Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes Cenoman). 12. Die procölen Crocodilia. Abhandlungen der Bayerischen Akademie der Wissenschaften. Mathematisch-naturwissenschaftliche Abteilung, 15, 1-55.
  45. ↑ Ibrahim, Nizar; Sereno, Paul C.; Varricchio, David J.; Martill, David M.; Dutheil, Didier B.; Unwin, David M.; Baidder, Lahssen; Larsson, Hans C. E.; Zouhri, Samir; Kaoukaya, Abdelhadi (2020). "Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco". ZooKeys (928): 1–216. Bibcode:2020ZooK..928....1I. doi: 10.3897/zookeys.928.47517 . ISSN   1313-2989. PMC   7188693 . PMID   32362741.
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