Apertotemporalis

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Apertotemporalis
Temporal range: Late Cretaceous, Cenomanian
Apertotemporalis holotype skull.png
Holotype braincase fragment in dorsal (a) and ventral (b) views
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Testudines
Suborder: Pleurodira
Family:† Bothremydidae
Genus:† Apertotemporalis
Stromer, 1934
Species:
†A. baharijensis
Binomial name
†Apertotemporalis baharijensis
Stromer, 1934

Apertotemporalis (meaning "open temporal fenestra") is a dubious genus of bothremydid turtle that lived in present-day Egypt during the Late Cretaceous period. It was described by German paleontologist Ernst Stromer in 1934. The genus contains a single species, A. baharijensis, named on the basis of a fragmentary skull. This fossil was unearthed by Austro-Hungarian paleontologist Richard Markgraf during an 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. Unfortunately, this skull was destroyed in 1944 during the Bombing of Munich in World War II.

Contents

Like other bothremydid turtles, Apertotemporalis was four-legged with a long neck ending in a small head, though its ecology has not been described. Its holotype skull was very poorly preserved, and only contained much of the braincase and a fragment of the anterior (front) region of the skull. Like that of other bothremydids, the skull was wide and low, measuring 9.6 centimetres (3.8 in) in width and 4 centimetres (1.6 in) in height. The skull was thought to bear diagnostic (distinguishing) characteristics by Stromer, however later paleontologists have considered the skull undiagnostic due to its worn and fragmentary condition, leading it to be declared a nomen dubium.

Apertotemporalis is a dubious member of the Bothremydidae, a family of side-necked turtles (Pleurodira). The genus coexisted with other turtles such as an indeterminate araripemydid, chelonioidean, and bothremydid, along with giant fishes like Mawsonia and Paranogmius , large crocodylomorphs like Stomatosuchus , and a host of dinosaurs like Spinosaurus, Paralititan, and Tameryraptor . 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 and validity

Geologic map of the Bahariya Oasis; Apertotemporalis fossils were found at the G. El Dist site Geological map of the Bahariya Oasis.PNG
Geologic map of the Bahariya Oasis; Apertotemporalis fossils were found at the G. El Dist site

The first fossils of this taxon were unearthed by Austro-Hungarian paleontologist Richard Markgraf in the Bahariya Oasis in the western desert of Egypt, with financial backing from German paleontologist Ernst Stromer and the Paläontologisches Museum München (Bavarian State Collection of Paleontology). The remains, consisting of an incomplete skull in three pieces, were found at the Gebel El Dist locality in a layer of greyish sandy mudstone at the base of the exposure. These mudstones are an outcrop of the Bahariya Formation, which contains fossils dating to the Cenomanian stage of the Late Cretaceous period. [1] [2] [3] In 1934, Stromer scientifically described the skull, and identified it as a new genus and species of turtle, Apertotemporalis baharijensis. The generic name Apertotemporalis comes from the Latin aperto "open" and temporalis (in reference to the temporal fenestra). The specific name baharijensis is in reference to the Bahariya Oasis and the Bahariya Formation where it was discovered. The poorly preserved holotype (name-bearing) specimen consisted of two larger skull fragments that pieced together form the braincase, while the third was a smaller fragment from the anterior portion of the skull. However, this fragment cannot be confidently associated with the other two pieces. [2] [4] [3]

Unfortunately, during the night of 24–25 April 1944, the Paläontologisches Museum München was severely damaged during the British bombing of Munich in World War II. The holotype of Apertotemporalis was destroyed in the attack. [3] [5] Due to personal and political tensions between Stromer and the museum's curator, who was a fervent Nazi, the fossils were not rehoused prior to the bombing. [6] Stromer's finds, including Apertotemporalis, received little academic or public attention. [6] [3] [5] [7] : 117  Since then, no other fossils have been assigned to the genus. [5] In 1946, German paleontologist Oskar Kuhn classified it in its own family within Pleurosternoidea, Apertotemporalidae, [8] which was later thought to include the genus Chitracephalus as well. [9] This clade was thought to be united by their large skulls and open temporal regions, [10] however the name has not seen use in recent decades. [5] [11]

In 1988, French paleontologist France de Lapparent de Broin classified Apertotemporalis as a member of Bothremydidae alongside a number of other Cretaceous North African pleurodirans like Nigeremys . [12] This assessment was followed by Antunes and Broin (1988), who speculated that Apertotemporalis was the African ancestor of later Bothremydids like Rosasia and that it was closely related to Bothremys (now Algorachelys ) parva. This study also stated that Apertotemporalis came from the Albian, [13] however it actually comes from the Cenomanian. [5] In contrary, in their description of Zolhafah, Broin and Werner (1998) assigned Apertotemporalis to a subfamily in Bothremydidae they nicknamed the "Nigeremys group". According to the authors, this clade consisted of Apertotemporalis, Nigeremys, and Sokotochelys, which were united by skull features such as an enlarged snout, an enlarged trochlear process behind the ear, and more. [14] Cladogram from Antunes and Broin (1988), who considered Apertotemporalis to be a basal bothremydid and ancestor of European genera like Rosasia. Their cladogram can be seen below (some unnamed bothremydids are not included): [13]

Pelomedusoides

Pelomedusidae

Podocnemididae

Bothremydidae

Apodichelys

Apertotemporalis

Madagascar taxon A

Madagascar taxon B

In their comprehensive review of pleurodiran turtles, American paleontologist Eugene Gaffney and colleagues (2006) stated that Apertotemporalis should be considered a nomen dubium "dubious name" based on the fact that the skull was destroyed in World War II, it was very poorly preserved due to weathering and damage, and that the skull itself features no diagnostic characteristics that distinguish it from other taxa. Additionally, Gaffney and colleagues (2006) stated that the skull only bears one characteristic of bothremydids, a fully enclosed "incisura columellae auris" (a notch in the quadrate bone of turtles), which is also found in other families. The authors closed by stating that the skull was so poorly preserved that the other Egyptian bothremydid Arenila could be the same taxon as Apertotemporalis, but due to the holotype's inadequate preservation, it is impossible to be sure. Gaffney's opinion that Apertotemporalis is a nomen dubium has been backed by later authors, who typically classify the taxon as a dubious bothremydid. [11] [3]

Description

Life restoration based on Galianemys Apertotemporalis restoration.jpg
Life restoration based on Galianemys

The skull was 9.6 centimetres (3.8 in) wide at the ear regions and only 4 centimetres (1.6 in) tall from the base to the top of the skull roof. Assuming proportions similar to those of Orbitia borneensis , the carapace of Apertotemporalis was 50 centimetres (20 in) long. However, Stromer noted that the holotype of Apertotemporalis was not associated with any carapace fragments and that the proportions of turtles vary widely, making these estimates tentative. From dorsal (top) view, a small crest was visible on the parietal (back cranium) bones that was similar to those of Testudo and Trionyx , though the extent of this crest was impossible to determine. This skull lacks a temporal roof or temporal arches, creating an opening in the temporal for a visible temporal fenestra. Additionally, this crest forms further posteriorly in Apertotemporalis than in other genera like Terrapene carolina . The pterygoids were flat and narrow with a hook-shaped ectopterygoid process on the distal side of the pterygoid. [2]

Palaeoenvironment and taphonomy

Restoration of fauna of the Bahariya Formation Bahariya Formation McAfee.jpg
Restoration of fauna of the Bahariya Formation

The discovery of Apertotemporalis indicates that bothremydids could inhabit marine-littoral environments, such as brackish lagoons and freshwater streams. Bothremydids' adaptations to saltwater environments like these facilitated their spread, with members of the family known from nearly every continent by the climax of the Cretaceous. [5] [15] Bothremydid diets are very diverse, with taxa being omnivorous, piscivorous, or carnivorous, aiding them in achieving their wide distribution. [16] [5] North Africa, during the Cenomanian stage of the Late Cretaceous, bordered the Tethys Sea, which transformed the region into a mangrove-dominated coastal environment filled with vast tidal flats and waterways. [17] [18] Fossils of terrestrial/freshwater animals, including those of Apertotemporalis, [2] were largely found in a lower layer of the Bahariya Formation whereas those of marine animals like plesiosaurs are more common in upper layers of the formation, indicating a change in the environment to a marine depositional one. [4] [3] Apertotemporalis was the first, and only, turtle described from the Bahariya Formation, however indeterminate taxa from the families Araripemydidae, Chelonioidea, Trionychia, Chelyidae, Pelomedusidae, and Bothremydidae have been described.

The faunal composition of both the Bahariya Formation and the Moroccan Kem Kem Beds were thought to be similar in the past, but the describers of the theropod dinosaur Tameryraptor suggested that such superficial comparisons require further examination. [1] A diverse fauna of other aquatic animals is known from the Bahariya Formation. Underwater life diversity exploded during this period in the mangroves of North Africa, represented by genera like the bony fish Mawsonia and Paranogmius , [4] [19] sawskates Onchopristis and Schizorhiza , [20] sharks like Squalicorax and Cretolamna, and a broad selection of invertebrates. [20] [4] As for non-dinosaurian reptiles, the early sea snake Simoliophis, several indeterminate plesiosaurs, crocodylomorphs like the stomatosuchid Stomatosuchus and the eunotosuchian Libycosuchus, [4] [21] [22] and an indeterminate pterosaur, possibly related to Anhanguera , are known from the formation. [23] Dinosaurs themselves are represented by the sauropods Aegyptosaurus , Paralititan , and an indeterminate titanosaur, [24] [18] the theropods Spinosaurus , [4] Tameryraptor, [1] Bahariasaurus , an indeterminate abelisaurid, [25] and possibly Sigilmassasaurus and Deltadromeus . [26] [27] [28]

References

  1. 1 2 3 Kellermann, Maximilian; Cuesta, Elena; Rauhut, Oliver W. M. (2025-01-14). "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.
  2. 1 2 3 4 Stromer, Ernst (1934). "Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltierreste der Baharije-Stufe (unterstes Cenoman). 14. Testudinata". Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch Naturwissenschaftliche Abteilung, Neue Folge. 25: 1–26.
  3. 1 2 3 4 5 6 Augustin, Felix J.; Hartung, Josephina; Kampouridis, Panagiotis (2023), Hamimi, Zakaria; Khozyem, Hassan; Adatte, Thierry; Nader, Fadi H. (eds.), "Dinosaur Faunas of Egypt—The Terrestrial Late Cretaceous Vertebrate Record", The Phanerozoic Geology and Natural Resources of Egypt, Cham: Springer International Publishing, pp. 253–284, doi:10.1007/978-3-030-95637-0_9, ISBN   978-3-030-95637-0
  4. 1 2 3 4 5 6 Stromer, E. (1936). "Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. VII. Baharije-Kessel und -Stufe mit deren Fauna und Flora. Eine ergänzende Zusammenfassung". Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung. Neue Folge (in German). 33: 1–102.
  5. 1 2 3 4 5 6 7 Gaffney, Eugene S; Tong, Haiyan; Meylan, Peter A (2006). "Evolution of the side-necked turtles: the families Bothremydidae, Euraxemydidae, and Araripemydidae". Bulletin of the American Museum of Natural History. 300: 1–698. doi:10.1206/0003-0090(2006)300[1:EOTSTT]2.0.CO;2. ISSN   0003-0090.
  6. 1 2 Smith, Joshua B.; Lamanna, Matthew C.; Mayr, Helmut; Lacovara, Kenneth 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. ISSN   0022-3360.
  7. ↑ Nothdurft, William; Smith, Josh (2002). The Lost Dinosaurs of Egypt. New York: Random House Publishing Group. ISBN   978-1-58836-117-2.
  8. ↑ Kuhn, Oskar (1946). "Das System der fossilen und rezenten Amphibien und Reptilien" (PDF). Bericht der Naturforschenden Gesellschaft in Bamberg. 29: 49–67.
  9. ↑ Pérez-García, Adán (2012). "The European Early Cretaceous Cryptodiran Turtle Chitracephalus dumonii and the Diversity of a Poorly Known Lineage of Turtles". Acta Palaeontologica Polonica. 57 (3): 575–588. Bibcode:2012AcPaP..57..575P. doi:10.4202/app.2011.0065. ISSN   0567-7920.
  10. ↑ Kuhn, Oskar (1960). "Die Familien der fossilen Amphibien und Reptilien". Bericht der Naturforschenden Gesellschaft Bamberg. 37: 20–52.
  11. 1 2 AbdelGawad, Mohamed; Pérez-García, Adán; Hirayama, Ren; Mohesn, Sara; Tantawy, Abdel-Aziz; Abu El-Kheir, Gebely (2023-02-16). "The First Side-Necked Turtle (Pleurodira, Bothremydidae) from the Campanian (Late Cretaceous) of Egypt". Diversity. 15 (2): 284. Bibcode:2023Diver..15..284A. doi: 10.3390/d15020284 . ISSN   1424-2818.
  12. ↑ Lapparent de Broin, France (1988). "Les tortues et le Gondwana. Examen des rapports entre le fractionnement du Gondwana au Cretace et la dispersion geographique des tortues pleurodires a partir du Cretace". Studia Salmanticensia, Studia Palaeocheloniologica. 2 (5): 103–142.
  13. 1 2 Antunes, M. T.; Lapparent de Broin, France (1988). "Le Crétacé terminal de Beira Litoral, Portugal: remarques stratigraphiques et écologiques,étude complémentaire de Rosasia soutoi (Chelonii, Bothremydidae)". Ciencias da Terra. 9: 153–200.
  14. ↑ de Broin, France de Lapparent; Werner, Christa (1998-04-01). "New late Cretaceous turtles from the Western Desert, Egypt". Annales de Paléontologie. 84 (2): 131–214. Bibcode:1998AnPal..84..131D. doi:10.1016/S0753-3969(98)80005-0. ISSN   0753-3969.
  15. ↑ Pérez-García, Adán; Ortega, Francisco; Murelaga, Xabier (2012-03-01). "A new genus of Bothremydidae (Chelonii, Pleurodira) in the Cretaceous of Southwestern Europe". Geobios. 45 (2): 219–229. Bibcode:2012Geobi..45..219P. doi:10.1016/j.geobios.2011.03.001. ISSN   0016-6995.
  16. ↑ AbdelGawad, Mohamed; Pérez-García, Adán; Hirayama, Ren; Mohesn, Sara; Tantawy, Abdel-Aziz; Abu El-Kheir, Gebely (2023-02-16). "The First Side-Necked Turtle (Pleurodira, Bothremydidae) from the Campanian (Late Cretaceous) of Egypt". Diversity. 15 (2): 284. Bibcode:2023Diver..15..284A. doi: 10.3390/d15020284 . ISSN   1424-2818.
  17. ↑ Hamed, Younes; Al-Gamal, Samir Anwar; Ali, Wassim; Nahid, Abederazzak; Dhia, Hamed Ben (March 1, 2014). "Palaeoenvironments of the Continental Intercalaire fossil from the Late Cretaceous (Barremian-Albian) in North Africa: a case study of southern Tunisia". Arabian Journal of Geosciences. 7 (3): 1165–1177. Bibcode:2014ArJG....7.1165H. doi:10.1007/s12517-012-0804-2. S2CID   128755145.
  18. 1 2 Smith, Joshua B.; Lamanna, Matthew C.; Lacovara, Kenneth J.; Dodson, Peter; Smith, Jennifer R.; Poole, Jason C.; Giegengack, Robert; Attia, Yousry (2001). "A Giant Sauropod Dinosaur from an Upper Cretaceous Mangrove Deposit in Egypt". Science. 292 (5522): 1704–1706. Bibcode:2001Sci...292.1704S. doi:10.1126/science.1060561.
  19. ↑ Allam, A. M. (1986-01-01). "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.
  20. 1 2 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.
  21. ↑ 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.
  22. ↑ 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.
  23. ↑ Sallam, Hesham; Salem, Belal; Thabet, Wael; El-Sayed, Sanaa; Lamanna, Matthew (2021). "The first pterosaur from the Upper Cretaceous (lower Cenomanian) Bahariya Formation, Bahariya Oasis, Western Desert of Egypt". Journal of Vertebrate Paleontology, Program and Abstracts: 226–227.
  24. ↑ Mannion, Philip D.; Barrett, Paul M. (2013-10-01). "Additions to the sauropod dinosaur fauna of the Cenomanian (early Late Cretaceous) Kem Kem beds of Morocco: Palaeobiogeographical implications of the mid-Cretaceous African sauropod fossil record". Cretaceous Research. 45: 49–59. Bibcode:2013CrRes..45...49M. doi:10.1016/j.cretres.2013.07.007. ISSN   0195-6671.
  25. ↑ 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. ISSN   2054-5703. PMC   9174736 . PMID   35706658.
  26. ↑ 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. (1996-05-17). "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.
  27. ↑ Cau, Andrea; Paterna, Alessandro (2025). "Beyond the Stromer's Riddle: the impact of lumping and splitting hypotheses on the systematics of the giant predatory dinosaurs from northern Africa". Italian Journal of Geosciences. 144 (2025) f.2 (2): 162–185. doi:10.3301/IJG.2025.10. ISSN   2038-1719.
  28. ↑ 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. ISSN   0008-4077.
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