Paralititan

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Paralititan
Temporal range: Late Cretaceous, Cenomanian
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
Genus:† Paralititan
Smith et al., 2001
Species:
†P. stromeri
Binomial name
†Paralititan stromeri
Smith et al., 2001

Paralititan (meaning "tidal giant") is a genus of 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 a fragmentary skeleton including vertebrae and limb bones. These fossils were unearthed by an American expedition to the Bahariya Oasis in western Egypt in rock layers of the Bahariya Formation. This formation dates to the Cenomanian stage of the Late Cretaceous, which lasted 101 to 94 million years ago. An incomplete dorsal (back) vertebra that had been described by German paleontologist Ernst Stromer in 1932 was also assigned to Paralititan. However, this vertebra had been destroyed during the Bombing of Munich in World War II.

Contents

Like other sauropods, Paralititan was a four-legged herbivore with a long neck ending in a small head. It was at one point considered one of the largest dinosaurs known. 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 sauropod, but not as huge as genera like Argentinosaurus and Puertasaurus. Its robust humerus (upper arm bone) is 1.69 meters (5.5 feet) in length and bears a large deltopectoral crest (a forward directed bony flange). The caudal (tail) vertebrae are wider than tall and lack pleurocoels (large cavities that stored air sacs) on their sides.

Paralititan is a member of the Saltasauridae, a family within the sauropod group Titanosauria. Some saltasaurids, including Saltasaurus itself, preserve dorsal osteoderms (bones formed in the skin), though such bones are unknown for Paralititan. Paralititan coexisted with other dinosaurs such as the sauropod Aegyptosaurus , the theropods Tameryraptor , Spinosaurus , and Bahariasaurus , and an unnamed abelisaurid theropod. During the Cenomanian, the Bahariya Formation was on the margin of the Tethys Sea, and represented a large network of mangrove swamps, rivers, and tidal flats.

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] The fossils that Markgraf found, including those of dinosaurs like Spinosaurus and Tameryraptor , had been shipped to Germany where they were described by German paleontologist Ernst Stromer. [4] [5] Markgraf had evidently removed all more complete skeletons, leaving only limited remains behind. In 2000, an American expedition was mounted to revisit the site but failed to find 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 sacrals (hip vertebrae, probably the 5th and 6th), the first caudal (tail vertebra), another caudal from the front portion of the tail, dorsal and sacral ribs, incomplete scapulae (shoulder blades), a complete right and an incomplete left humerus (upper arm bone), the lower end of a metacarpal, and several additional elements. All of these fossils were 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 101 to 94 million years ago old. [3]

In 2001, an international crew made up of paleontologist Joshua B. Smith and colleagues described the partial skeleton as 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 entire 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 dorsal (vertebra from the front portion of the trunk), cataloged at the Paläontologisches Museum München (Bavarian State Collection of Paleontology) under SNSB-BSPG 1912V11164, to Paralititan stromeri. Stromer, in 1932, assumed that this vertebra belonged 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 assigned by Stromer to the other Bahariya sauropod Aegyptosaurus , [7] but these may belong to Paralititan instead. [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. The limited material, especially the long humeri, suggested that it was one of the most massive dinosaurs ever discovered, with an estimated weight of 59 t (65 short tons) according to a 2011 study. [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 measures 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 titanosaur Notocolossus , which includes 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 a 2006 study. [12]

Life restoration Paralititan stromeri.jpg
Life restoration

American researcher Scott Hartman estimated that the animal was massive, but still smaller than the biggest titanosaurs such as Puertasaurus , Alamosaurus , and Argentinosaurus . [13] In the 2010 edition of his book The Princeton Field Guide to Dinosaurs , 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 2016, using equations that estimate body mass based on the circumference of the humerus and femur of quadrupedal animals, Argentine paleontologist Bernardo González Riga and colleagues gave a weight estimate of ~50 t (55 short tons) for Paralititan. [11] In 2019, Paul estimated Paralititan between 30 and 55 tonnes (33–60.6 short tons) in weight. [15] 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) in their book Dinosaur Facts and Figures: The Sauropods and Other Sauropodomorphs. [16]

Skeleton

The sacral vertebrae of Paralititan lack pleurocoels , large spaces in the sides of the vertebra that stored pneumatic air sacs. [3] Air sacs made vertebrae lighter and were part of the respiratory system. [17] [18] The absence of pleurocoels on the caudal vertebrae is common amongst titanosaurids, supporting its assignment to the group. One of the proximal (towards body) caudal vertebrae has a wider than tall, procoelous centrum with a convex rear articular condyle. In contrast to other titanosaurs like Alamosaurus and Neuquensaurus , the centrum is not convex on both ends. The ventral (bottom) surface of the centrum has weakly pronounced ridges that border a sagittal concavity. The other proximal caudal vertebra found has a strongly procoelus centrum with a well-developed distal condyle. The scapula (shoulder blade) has a medial (inner) 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 (rear-bottom) part of the scapula underneath the glenoid socket (the part of the scapula that articulates with the humerus). This feature is only found in Brachiosaurus and Paralititan, demonstrating that it is a unique genus of titanosaur. [3]

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

Paralititan's humerus is huge at 1.69 meters (5.5 feet) in length, with greatly expanded upper and lower ends. The upper region of the proximal surface of the humerus bears a muscular depression (an area for muscle attachment). The deltopectoral crest is large, stretching for more than 53% of the length of the humerus. On the lower proximal surface of the distal end of the humerus is a large fossa (depression) 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 upper 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. At the condyles (area where the humerus articulates with the ulna and radius) at the distal end of the humerus are supracondylar ridges. These ridges extend over 1/3rd of the humerus' length and flank a large olecranon fossa (area of the humerus that the ulna's olecranon process articulates), like in other titanosaurs. The metacarpal known from Paralititan has a rectangular, flattened surface indicative phalangeal reduction or absence on this digit. A second sauropod, Aegyptosaurus, is known from the Bahariya Formation. 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. [19] [3] [20] 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) [21] and 26 metres (85 ft) long respectively. [15] [22] 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. [23] In studies by American paleontologist Kristina Curry Rogers (2005) [24] and British paleontologists Philip D. Mannion and Paul Upchurch (2011), [25] 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. [23]

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. [26] [27] [28] [29] Some saltasaurids like Saltasaurus had dorsal osteoderms on their backs, [30] though other saltasaurids like Paralititan have not been discovered with these osteoderms. [3] Based on the study, 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 of these features are found in Paralititan. [29]

Below is a cladogram by Villa et al. (2022), from the description of the European saltasaurine Abditosaurus. [29]

Saltasauridae

Palaeoenvironment and taphonomy

Restoration of Paralititan (background, left) with contemporaneous animals of the Bahariya Formation Bahariya Formation McAfee.jpg
Restoration of Paralititan (background, left) with contemporaneous animals of the Bahariya Formation

The autochthonous (a fossil deposit with remains that were not moved by biological or physical processes), associated holotype skeleton was preserved in tidal flat deposits with a mangrove vegetation of seed ferns, Weichselia reticulata, according to the occurrence of plant fossils and root marks in the sediments. The 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 skeleton was autochthonous. An isolated tooth assigned to Carcharodontosaurus , [3] a genus formerly thought to be present in the Bahariya Formation but now assigned 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 shed 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. [31] 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. [32] [33] 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. [34] 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 [35] and Paranogmius , [36] sawskates Onchopristis and Schizorhiza , [37] sharks like Squalicorax and Cretolamna, and a broad selection of invertebrates. [38] Additionally, several crocodylomorphs like the stomatosuchid Stomatosuchus [39] and the eunotosuchian Libycosuchus are known from the formation. [40] [41] [33]

References

  1. 1 2 Stromer, E. (1915). "Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 3. Das Original des Theropoden Spinosaurus aegyptiacus nov. gen., nov. spec". Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften, Mathematisch-physikalische Klasse (in German). 28 (3): 1–32.[ permanent dead link ]
  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.
  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.
  9. ↑ Burness, G.P. and Flannery, T. (2001). "Dinosaurs, Dragonslayer, and Dwarfs: The Evolution of Maximal Body Size." Proceedings of the National Academy of Sciences, 98(25): 14518-14523.
  10. ↑ Roach, John (May 31, 2001). "'Tidal Giant' Roamed Coastal Swamps of Ancient Africa". National Geographic News. Washington, D.C.: National Geographic Society. Archived from the original on June 5, 2001. Retrieved December 31, 2012.
  11. 1 2 González Riga BJ, Lamanna MC, Ortiz David LD, Calvo JO, Coria JP (January 2016). "A gigantic new dinosaur from Argentina and the evolution of the sauropod hind foot". Scientific Reports. 6 (1) 19165. Bibcode:2016NatSR...619165G. doi:10.1038/srep19165. PMC   4725985 . PMID   26777391.
  12. ↑ Carpenter K (2006). Foster JR, Lucas SG (eds.). "Biggest of the Big: a Critical Re-evaluation of the Mega-sauropod Amphicoelias fragillimus Paleontology and Geology of the Upper Jurassic Morrison Formation" (PDF). New Mexico Museum of Natural History and Science Bulletin. 36: 131–138. Archived from the original (PDF) on March 8, 2016.
  13. ↑ Hartman, Scott. "The biggest of the big". Skeletal Drawings.
  14. ↑ Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 209
  15. 1 2 Paul, Gregory S. (2019). "Determining the largest known land animal: A critical comparison of differing methods for restoring the volume and mass of extinct animals" (PDF). Annals of the Carnegie Museum. 85 (4): 335–358. Bibcode:2019AnCM...85..335P. doi:10.2992/007.085.0403. S2CID   210840060.
  16. ↑ Molina-Perez, Ruben; Larramendi, Asier (2020). Dinosaur Facts and Figures: The Sauropods and Other Sauropodomorphs. New Jersey: Princeton University Press. p. 267. Bibcode:2020dffs.book.....M.
  17. ↑ O'Connor, Patrick Michael (2009). "Evolution of archosaurian body plans: skeletal adaptations of an air-sac-based breathing apparatus in birds and other archosaurs" . Journal of Experimental Zoology Part A: Ecological Genetics and Physiology. 311A (8): 629–646. Bibcode:2009JEZA..311..629O. doi:10.1002/jez.548. ISSN   1932-5231. PMID   19492308.
  18. ↑ O'Connor, Patrick M. (2006). "Postcranial pneumaticity: An evaluation of soft-tissue influences on the postcranial skeleton and the reconstruction of pulmonary anatomy in archosaurs" . Journal of Morphology. 267 (10): 1199–1226. Bibcode:2006JMorp.267.1199O. doi:10.1002/jmor.10470. ISSN   1097-4687. PMID   16850471.
  19. ↑ Cerda, Ignacio A.; Paulina Carabajal, Ariana; Salgado, Leonardo; Coria, Rodolfo A.; Reguero, Marcelo A.; Tambussi, Claudia P.; Moly, Juan J. (January 1, 2012). "The first record of a sauropod dinosaur from Antarctica". Naturwissenschaften. 99 (1): 83–87. Bibcode:2012NW.....99...83C. doi:10.1007/s00114-011-0869-x. ISSN   1432-1904. PMID   22173579.
  20. ↑ Mocho, Pedro; Escaso, Fernando; Marcos-Fernández, Fátima; Páramo, Adrián; Sanz, José Luis; Vidal, Daniel; Ortega, Francisco (September 4, 2024). "A Spanish saltasauroid titanosaur reveals Europe as a melting pot of endemic and immigrant sauropods in the Late Cretaceous". Communications Biology. 7 (1): 1016. doi:10.1038/s42003-024-06653-0. ISSN   2399-3642. PMC   11375222 . PMID   39232208.
  21. ↑ Paul, G.S. (2024) The Princeton Field Guide to Dinosaurs. 3nd ed. Princeton University Press p. 256
  22. ↑ Lehman, Thomas M.; Coulson, Alan B. (2002). "A juvenile specimen of the sauropod dinosaur Alamosaurus sanjuanensis from the Upper Cretaceous of Big Bend National Park, Texas" . Journal of Paleontology. 76 (1): 156–172. doi:10.1666/0022-3360(2002)076<0156:AJSOTS>2.0.CO;2. ISSN   0022-3360.
  23. 1 2 Gorscak, Eric; O'Connor, Patrick M.; Stevens, Nancy J.; Roberts, Eric M. (July 29, 2014). "The basal titanosaurian Rukwatitan bisepultus (Dinosauria, Sauropoda) from the middle Cretaceous Galula Formation, Rukwa Rift Basin, southwestern Tanzania". Journal of Vertebrate Paleontology. 34 (5): 1133–1154. Bibcode:2014JVPal..34.1133G. doi:10.1080/02724634.2014.845568. ISSN   0272-4634.
  24. ↑ Curry Rogers, K. A. (2005). Titanosauria: a phylogenetic overview; pp. 50–103 in K. A. Curry Rogers and J. A. Wilson (eds.), The Sauropods: Evolution and Paleobiology. University of California Press, Berkeley, California.
  25. ↑ Mannion, Philip D.; Upchurch, Paul (January 15, 2011). "A re-evaluation of the 'mid-Cretaceous sauropod hiatus' and the impact of uneven sampling of the fossil record on patterns of regional dinosaur extinction" . Palaeogeography, Palaeoclimatology, Palaeoecology. 299 (3): 529–540. Bibcode:2011PPP...299..529M. doi:10.1016/j.palaeo.2010.12.003. ISSN   0031-0182.
  26. ↑ Averianov, Alexander; Sues, Hans-Dieter (January 1, 2017). "Review of Cretaceous sauropod dinosaurs from Central Asia" . Cretaceous Research. 69: 184–197. Bibcode:2017CrRes..69..184A. doi:10.1016/j.cretres.2016.09.006. ISSN   0195-6671.
  27. ↑ 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.
  28. ↑ 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.
  29. 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.
  30. ↑ 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. hdl: 11336/73560 . ISSN   0567-7920.
  31. ↑ 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.
  32. ↑ 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.
  33. 1 2 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.
  34. ↑ 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.
  35. ↑ 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.
  36. ↑ 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.
  37. ↑ 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.
  38. ↑ 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.
  39. ↑ 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.
  40. ↑ 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.
  41. ↑ 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.
  42. Holtz, Thomas R. Jr. (2012) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2011 Appendix.
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