Cretalamna

Cretalamna is a genus of extinct otodontid shark lived about 103 to 46 million years ago from the Late Cretaceous to Eocene epochs. It is considered by many to be the ancestor of the largest sharks to have ever lived, Carcharocles angustidens, and Carcharocles megalodon.

Cretalamna Temporal range: Late Albian-Lutetian, 103.13–46.25 Ma PreꞒ Ꞓ O S D C P T J K Pg N [1][2][3][4][5][lower-alpha 1]
Jaws and teeth of the C. hattini holotype (LACM 128126) from the Niobrara Formation of Kansas
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Chondrichthyes
Order:	Lamniformes
Family:	†Otodontidae
Genus:	†Cretalamna Glükman, 1958
Type species
†Cretalamna appendiculata Agassiz, 1835
Species
List of species †C. appendiculata Agassiz, 1835 (type) †C. lata Agassiz, 1843 †C. borealis Priem, 1897 †C. biauriculata Wanner, 1902 †C. maroccana Arambourg, 1935 †C. nigeriana Cappetta, 1972 †C. arambourgi Cappetta & Case, 1975 †C. catoxodon Siversson et al., 2015 †C. deschutteri Siversson et al., 2015 †C. ewelli Siversson et al., 2015 †C. gertericorum Siversson et al., 2015 †C. hattini Siversson et al., 2015 †C. sarcoportheta Siversson et al., 2015 †C. bryanti Ebersole and Ehret, 2018 Disputed or uncertain † 'C.' aschersoni Stromer, 1905 † 'C'. twiggsensis Case, 1981
Synonyms[1][6][7][8][9]
List of synonyms Squalus mustelus Mantell, 1822 Squalus cornubicus Geinitz, 1839 Odontaspis raphiodon Geinitz, 1839 Lamna appendiculata Agassiz, 1835 Otodus appendiculatus Agassiz, 1843 Otodus latus Agassiz, 1843 Otodus basalis Stoliczka, 1873 Lamna (Otodus) appendiculata Zittel, 1895 Lamna borealis Priem, 1897 Lamna (Otodus) appendicularis Toula, 1900 Odontaspis gigas Dalinkevicus, 1935 Lamna lata Gyen, 1937 Plicatolamna arcuata Edwards, 1976 Cretolamna appendiculata var. pachyrhiza Herman, 1977 Cretolamna appendiculata pachyrhiza Lauginiger and Hartstein, 1983 Cretoxyrhina cf. mantelli Kemp, 1991 Cretolamna woodwardi Welton and Farish, 1993 Cretolamna pachyrhyza Herman and Van Waes, 2012

Taxonomy

Research History

Original syntypes of Otodus appendiculatus (Fig. 1-25) and holotype of C. lata (Fig. 26) in the third volume of Recherches sur les poissons fossiles; the tooth in Fig. 10 is the sole lectotype of C. appendiculata

Cretalamna was first described by Swiss naturalist Louis Agassiz using five teeth previously identified as the common smooth-hound and collected by English paleontologist Gideon Mantell from the Southerham Grey Pit near Lewes, East Sussex. In his 1835 publication Rapport sur les poissons fossiles découverts en Angleterre, he reidentified them as a new species of porbeagle shark under the taxon Lamna appendiculata.[9] In 1843, Agassiz published Recherches sur les poissons fossiles, which reexamined Mantell's five teeth. Using them, eight additional teeth collected by Mantell, and twenty more teeth collected by various paleontologists in various locations (One tooth found by the 3rd Earl of Enniskillen from the Speeton Clay in Yorkshire; one tooth of the collection of a Strasbourg Museum from an unspecified location; one tooth of the collection of the Hancock Museum from the Marly Chalk near Cambridge; and six teeth of the collection of German paleontologist Heinrich Georg Bronn from chalk around Aachen), he described a species whose teeth had thick bulged roots, lateral cusplets, and extreme variability. Agassiz remarked that some of the examined teeth may be variable enough to belong to a separate species, but ultimately unified them under a new taxon Otodus appendiculatus.[10] The species would later be found in 1958 by Soviet paleontologist Leonid Glickman to belong to a distinct new genus- Cretalamna.[11]

Despite Agassiz's remarks on variability,[10] his ultimately broad interpretation of O. appendiculatus subsequently led the species to become a wastebasket taxon culminating to an interpretation of C. appendiculata as a variable cosmopolitan species with a 50 million year range. This changed when paleontologist Mikael Siversson found that the twenty-five syntypes actually represented a mix of at least six or more different species including three additional genera Dwardius, Cretoxyrhina, and Cretodus. To remedy the taxonomic issue, he redesignated one of the syntypes as the sole lectotype of C. appendiculata in 1999.[1][12] In 2015, he led a study which revisited the taxonomic situation and established a renewed description of the species, which led to the erection of six additional Cretalamna species- C. catoxodon, C. deschutteri, C. ewelli, C. gertericorum, C. hattini, and C. sarcoporthea.[1]

Before Siversson, other Cretalamna species have been described. Another species described by Agassiz under the taxon Otodus latus was demoted to a variation of C. appendiculata in 1908,[13] promoted into a subspecies in 1977 by French paleontologist Jaques Herman,[14] and finally elevated to the species level as Cretolamna lata by Herman and paleontologist Van Waes Hilde in 2012.[15] In 1897, French paleontologist Fernand Priem described a single tooth from the Köpinge Sandstone in Scania, Sweden under the taxon Lamna borealis. This would be revised to 'Cretolamna borealis' by Glickman in a 1980 paper.[1] In 1902, German paleontologist Johannes Wanner described teeth from Egyptian Cretaceous deposits near the Dakhla Oasis and Farafra. He noted that the teeth are almost identical with that of the Otodus appendiculatus teeth, except that the Egyptian teeth also contained two clear pairs of lateral cusplets (a feature not seen in Otodus appendiculatus). Wanner concluded that the teeth were of a closely related new species and placed it under the taxon Otodus biauriculatus.[16] In 1935, French Paleontologist Camille Arambourg described a new subspecies of C. biauriculata from teeth found in Moroccan phosphates under the taxon Lamna biauriculata maroccana,[17] which was elevated into its own species in 1997.[18] In 1972, French ichthyologist Henri Cappetta described teeth from Maastrichtian deposits near the Mentès well in Tahoua, Niger, which he assigned to the subspecies Lamna biauriculata nigeriana.[19] This subspecies would also be elevated to its own species in 1991.[20] In 1975, Cappetta and American paleontologist Gerard Case examined Cretalamna teeth described by Arambourg in 1952 from Danian deposits in Morocco and proposed that it represents a new subspecies of the type species and assigned it the taxon Cretolamna appendiculata arambourgi,[21] which Siversson et al. (2015) elevated into its own species.[1] In 2018, American paleontologists Jun Ebersole and Dana Ehret described a new species of Cretalamna from various teeth from the Eutaw Formation and Mooreville Chalk in Alabama, which they named C. bryanti.[22]

Etymology

The genus Cretalamna is a portmanteau of creta, the Latin word for "chalk", prefixed to the genus Lamna, which is a romanization of the Ancient Greek λάμνα (lámna, meaning "kind of fierce shark"). When put together they mean "chalk-shark", which refers to chalk deposits from which the species' type specimens were found in.[11] The type species name appendiculata is a feminine form of the Latin word appendiculātus (having an appendage), a reference to the thick bulged roots found in C. appendiculata teeth.[10] The species name lata is derived from the feminine form of the Latin lātus (wide); a reference to the notably wide teeth of the species.[10] The species name borealis is derived from the Latin boreālis (northern); this is a reference to its discovery from fossil deposits in Sweden, a boreal locality.[1] The specific epithet of C. maroccana is a feminine form of the Latin word maroccānus (Moroccan), a reference to its type locality in Morocco.[23] C. biauriculata's specific epithet is a portmanteau derived from the Latin prefix bi- (two) prefixed onto the Latin auriculātā (eared), together meaning "having two ears". This is a reference to the species' large lateral cusplets, which somewhat resemble a pair of ears.[16] The species name nigeriana is derived from the country name Niger prefixed to the suffix -iana, a feminine variation of the Latin suffix -ānus (pertaining to), together meaning "pertaining to Niger". This is a reference to the species' type locality in Niger.[19] The species name sarcoportheta is derived from the Ancient Greek σαρκός (sarkos, meaning "flesh") prefixed to the Ancient Greek πορθητής (porthitís, meaning "destroyer"), together meaning "destroyer of flesh". The species name catoxodon is derived from the Ancient Greek κατοξυς (katoxys, meaning "very sharp") prefixed to the Ancient Greek ὀδών (odon, meaning "tooth". Together they mean "very sharp tooth", referring to the unusually sharp cutting edges of some C. catoxodon teeth.[1]

Six of the Cretalamna species have specific epithets that are named in honor of specific people, either for their contributions to the research of their associated species or for notable work they undertook. Of these six, five share a similar word structure that has a person's last name prefixed onto the Latin suffix -i (from). These species are C. arambourgi, which honors paleontologist Camille Arambourg for his discovery of the C. arambourgi type specimens and his contributions to North African paleontology;[21] C. bryanti, which honors the Bryant family who helped enhance the reputation and missions of the University of Alabama, Alabama Museum of Natural History, and McWane Science Center through their commitment to education and support;[22] C. deschutteri, which honors paleontologist Pieter De Scutter for his efforts to make Cretalamna teeth from a Bettrechies quarry available to Siversson et al. (2015) and for his work on Belgian Cenozoic sharks; C. ewelli, which honors paleontologist Keith Ewell who collected most of the C. ewelli type specimens in 2004; and C. hattini, which honors the late geologist Donald E. Hattin "for his work on the stratigraphy of the Niobrara Formation, western Kansas". The specific epithet of C. gertericorum is structured differently; it is derived from the names "Gert", "Eric", and the Latin suffix -orum (a masculine plural declension). The derived names "Gert" and "Eric" refer to fossil collectors Gert De Bie and Eric Collier, both of whom collected the majority of Cretalamna teeth examined in Siversson et al. (2015) that were from the Bettrechies quarry.[1]

Spelling

The valid spelling of Cretalamna, specifically between it and 'Cretolamna', has been subject to controversy. Originally, Glickman described the genus with the intention of naming it as 'Cretolamna' , but during publication of the corresponding 1958 paper a typographical error occurred, with the print misspelling it as 'Cretalamna' . Glickman pointed out the spelling as an error and continued to use his intended spelling 'Cretolamna' in later works.[17] This spelling was universally adopted until 1999 when Siversson remarked that this violates ICZN Articles 32 and 33, reinstating 'Cretalamna' as the valid spelling.[12] Since then, the reinstatement of 'Cretalamna' gained prominence and by the 2010s, was accepted by the majority of paleontologists.[1] However, some paleontologists including Cappetta strongly opposed it.[1][17] In an attempt to suppress the usage of 'Cretalamna' , Cappetta appealed to a representative of the ICZN, arguing that the original intentions of Glickman and the prevailing usage of 'Cretolamna' prior to Siversson (1999) secures its priority. The ICZN, who reportedly were impressed by Cappetta's "spirit", subsequently erected Article 33.3.1 of the 2000 Edition of the Code in order to address this situation in the future,[17] which states that "when an unjustified emendation is in prevailing usage and is attributed to the original author and date it is deemed to be a justified emendation". While Cappetta argued in a 2012 handbook that this new provision justifies the priority of 'Cretolamna' due to the spelling's overwhelmingly prevailing usage prior to its replacement by Siversson in 1999, Siversson himself pointed out in a 2015 paper that the provision cannot be worked retroactively, and that the continued prevailing usage of 'Cretalamna' since the provision's establishment ironically secures its priority rather than threaten it.[1] 'Cretalamna' currently remains as the most prevalent spelling and paleontologists have expressed the unlikeliness of a return to the usage of 'Cretolamna' .[17]

Biology

Body

Cretalamna was a medium-sized shark with an estimated total length of 2.3 – 3m. It is presumed to have had a fusiform-like body comparable to those of modern lamniformes. This is evidenced by the circular vertebra centra of Cretalamna.[24]

Given the size of Cretalamna teeth relative to its body size, it is likely that its caudal fin was not used for stunning prey (as is the case for thresher sharks). Instead, the caudal fin of Cretalamna was more similar to that of the great white shark.[24]

Dentition

Fossil teeth of C. biauriculata from Khouribga (Morocco)

Cretalamna teeth are distinguished by a broad triangular cusp and two lateral cusplets. The cutting edges of the teeth are razor-like, while the sides have a smooth surface. Teeth symmetry is variable; some have exact bilateral symmetry whereas others have high asymmetry. Adjacent teeth do not overlap.[24]

The exact dentition of Cretalamna is uncertain due to poor fossil representation. Traditionally, most reconstructions of its dentition were constructed from individual shed teeth.[1] Based on a specimen of C. hattini (LACM 128126), the dentition of the shark follows a lamnoid pattern with at least fifteen upper tooth rows and eight lower tooth rows on each side of the jaw. The upper tooth rows contain, from front to back: two symphysial, two anterior, one intermediate, and ten lateral tooth rows. The lower tooth rows contain: two anterior, one intermediate, and five lateral tooth rows. This is given in the dental formula S2?.A2.I1.L10(+?)s?.a2.i1.l4, constructed in a 2007 study of LACM 128126 by paleontologist Kenshu Shimada. It is possible that Cretalamna contained more than two symphysial tooth rows, as the related Cretoxyrhina mantelli possessed four upper symphysial tooth rows.[24]

Jaw

In C. hattini, the upper and lower jaws are similar to that of Cretoxyrhina mantelli. The jaws also resemble those of modern alopiids (thresher sharks) and lamnids. Limited fossil evidence suggests that the upper jaws extended over the lower jaws, giving Cretalamna a subterminal mouth.[24]

Paleoecology

Distribution

Cretalamna was a widespread genus found in North Africa (Morocco), the Near East (Jordan) [25] and North America both on the East Coast and in the Midwest. Deposits in Morocco are usually Eocene in age; deposits in Jordan are of Cretaceous and Eocene in age; and most deposits in the U.S. are of Cretaceous and Paleocene age.[26] C. maroccana is more prevalent in Morocco and Jordan, while C. appendiculata is more prevalent in the United States. Both species overlapped at one point in time.

Habitat

Cretalamna (13, lower right) with contemporaneous aquatic animals

Fossil evidence of Cretalamna is found in deposits representing a diverse set of marine environments, indicating that it was able to adapt to a wide range of habitats. This may have attributed to its ability to exist through a long temporal range.[24] The fusiform body of Cretalamna suggests it was a pelagic shark.[27]

The Cretaceous waters inhabited by Cretalamna were also home to a diverse range of cartilaginous fishes, bony fishes, turtles, squamates, plesiosaurs, pterosaurs, birds, and even some non-avian dinosaurs.[24]

Diet

The teeth morphology of Cretalamna implies that it was a generalist.[24] It was a predator and preyed upon large bony fish, turtles, mosasaurs, squids, and other sharks.[28] Some tooth specimens of Cretalamna exhibit heavy wear—likely the result of drastic diet changes.[1]

Extinction

A possible factor to the extinction of Cretalamna is increased competition with newer generalist sharks during the Cenozoic.[24] It is widely believed that Otodus (and thus Carcharocles) is derived from Cretalamna due to its strong similarity to certain species within the genus.[22]

Notes

1. Cappetta (2012) reported 'C'. aschersoni from the Priabonian of Egypt.

See also

• Cretaceous sharks
• Prehistoric fish
• Squalicorax

References

1. Mikael Siversson; Johan Lindgren; Michael G. Newbrey; Peter Cederström; Todd D. Cook (2015). "Cenomanian–Campanian (Late Cretaceous) mid-palaeolatitude sharks of Cretalamna appendiculata type" (PDF). Acta Palaeontologica Polonica. 60 (2): 339–384. doi:10.4202/app.2012.0137.
2. Mikael Siversson; Marcin Machalski (2017). "Late late Albian (Early Cretaceous) shark teeth from Annopol, Poland". Alcheringa: An Australasian Journal of Palaeontology. 41 (4): 433–463. doi:10.1080/03115518.2017.1282981.
3. J.G. Ogg and L.A. Hinnov (2012). Cretaceous. The Geologic Time Scale 2012. pp. 793–853. doi:10.1016/B978-0-444-59425-9.00027-5. ISBN 9780444594259.
4. Agnete Weinreich Carlsen; Gilles Cuny (2014). "A study of the sharks and rays from the Lillebælt Clay (Early–Middle Eocene) of Denmark, and their palaeoecology" (PDF). Bulletin of the Geological Society of Denmark. 62 (1): 39–88.
5. N. Vandenberghe; F. J. Hilgen; R. P. Speijer (2012). "The Paleogene Period". The Geologic Time Scale 2012: 855–921. doi:10.1016/B978-0-444-59425-9.00028-7. ISBN 9780444594259.
6. Vasja Mikuz (2003). "The elasmobranch Cretolamna appendiculata also in the Upper Cretaceous - Gosau beds near Stranice, Eastern Slovenia". Geologija (in Slovenian). 46 (1): 83–87. doi:10.5474/geologija.2003.006.
7. Arthur Smith Woodward (1889). Catalogue of the fossil fishes in the British Museum (Natural History). 1. British Museum. p. 393. doi:10.5962/bhl.title.61854.
8. Kenshu Shimada (2006). "Fossil marine vertebrates from the Lowermost Greenhorn Limestone (Upper Cretaceous: Middle Cenomanian) in southeastern Colorado". Journal of Paleontology. 80: 1–45. doi:10.1666/0022-3360(2006)80[1:FMVFTL]2.0.CO;2. ISSN 0022-3360.
9. Louis Agassiz (1835). Rapport sur les poissons fossiles découverts en Angleterre (in French). Imprimerie de Petitpierre et Prince. p. 49. doi:10.5962/bhl.title.5745.
10. Louis Agassiz (1833–1843) [1838]. Recherches sur les poissons fossiles (in French). 3. Museum of Comparative Zoology.
11. Leonid Glickman (1958). "Rates of evolution in Lamoid sharks" (PDF). Doklady Akademii Nauk SSSR (in Russian). 123: 568–571.
12. Mikael Siversson (1999). "A new large lamniform shark from the uppermost Gearle Siltstone (Cenomanian, Late Cretaceous) of Western Australia". Transactions of the Royal Society of Edinburgh: Earth Sciences. 90 (1): 49–66. doi:10.1017/S0263593300002509.
13. Fernand Priem (1908). Etude des Poissons Fossiles du Bassin Parisien (PDF) (in French). Publications des Annales de Paleontologie. pp. 69–70.
14. Jaques Herman (1977). Les sélaciens des terrains néocrétacés & paléocènes de Belgique & des contrées limitrophes eléments d'une biostratigraphie intercontinentale (in French). Geological Survey of Belgium. pp. 210–216. ISSN 0408-9510.
15. Jaques Herman; Van Waes Hilde (2012). Observations concernant l'Évolution et la Systématique de quelques Euselachii, Neoselachii et Batoidei (Pisces - Elasmobranchii), actuels et fossiles (PDF) (in French). Géominpal Belgica 2. pp. 55–56. ISSN 2033-6365.
16. Johannes Wanner (1902). "Die Fauna der obersten weissen Kreide der libyschen Wüste". Palaeontographica (in German). 30 (2): 91–152.
17. Jim Bourdon (2011). "Cretalamna". elasmo.com. Retrieved 31 March 2019.
18. Gerard Case; Henri Cappetta (1997). "A new selachian fauna from the Late Maastrichtian of Texas (Upper Cretaceous/Navarro Group; Kemp Formation)". Münchner Geowissenschaftliche Abhandlungen Reihe A: Geologie und Paläontologie. 34 (1): 134–189. ISSN 0177-0950.
19. Henri Cappetta (1972). "Les poissons crétacés et tertiaires du bassin des Iullemmeden (République du Niger)". Palaeovertebrata (in French). 5 (5): 179–251. ISSN 2274-0333.
20. R.T.J. Moody; P.J.C. Sutcliffe (1991). "The Cretaceous deposits of the Iullemmeden Basin of Niger, central West Africa". Cretaceous Research. 12 (2): 137–157. doi:10.1016/S0195-6671(05)80021-7.
21. Henri Cappetta; Gerard Case (1976). "Contribution a l'étude des sélaciens du groupe Monmouth (Campanien-Maestrichtien) du New Jersey". Palaeontographica Abteilung A (in French). 151 (1–3): 1–46.
22. Jun A. Ebersole; Dana J. Ehret (2018). "A new species of Cretalamna sensu stricto (Lamniformes, Otodontidae) from the Late Cretaceous (Santonian-Campanian) of Alabama, USA". PeerJ. 6 (e4229): e4229. doi:10.7717/peerj.4229. PMC 5764036. PMID 29333348.
23. Camille Arambourg (1952). "Les vertébrés fossiles des gisements de phosphates (Maroc-Algérie-Tunisie)". Notes et Mémoires du Service Géologique du Maroc (in French). 92 (1): 1–372.
24. Shimada, Kenshu (2007). "Skeletal and Dental Anatomy of Lamniform Shark, Cretalamna Appendiculata, from Upper Cretaceous Niobrara Chalk of Kansas". Journal of Vertebrate Paleontology. 27 (3): 584–602. doi:10.1671/0272-4634(2007)27[584:SADAOL]2.0.CO;2.
25. Kaddumi H. F. 2009. Fossils of the Harrana Fauna and the Adjacent Areas. Publications of the Eternal River Museum of Natural History, Amman, pp 324
26. Yoost, Derek. "Potomac River Fossils". Retrieved 14 May 2012.
27. Ferrón, Humberto G. (2017-09-22). "Regional endothermy as a trigger for gigantism in some extinct macropredatory sharks". PLoS ONE. 12 (9): e0185185. Bibcode:2017PLoSO..1285185F. doi:10.1371/journal.pone.0185185. ISSN 1932-6203. PMC 5609766. PMID 28938002.
28. Kent, Bretton W. (1987). Fossil Sharks of the Chesapeake Bay Region. Egan Rees & Boyer Inc.