Cosmopolitodus

Cosmopolitodus; Temporal range: Rupelian-Calabrian, 30–1.07 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	Tooth series of C. hastalis
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Chondrichthyes
Subclass:	Elasmobranchii
Subdivision:	Selachimorpha
Order:	Lamniformes
Family:	Lamnidae
Genus:	† Cosmopolitodus ; Glikman, 1964
Type species
	†Cosmopolitodus hastalis; Agassiz, 1843
Species
	†C. hastalis(Agassiz, 1843); †C. planusAgassiz, 1856;
Synonyms
	Carcharodon hastalis(Agassiz, 1843); Isurus hastalis(Agassiz, 1843); Isurus xiphodon(Agassiz, 1843); Oxyrhina hastalis(Agassiz, 1843); Oxyrhina xiphodon(Agassiz, 1843); Oxyrhina plana(Agassiz, 1856); Isurus planus(Jordan, 1907);

Last updated October 16, 2024

Cosmopolitodus is an extinct genus of mackerel shark that lived between thirty and one million years ago during the late Oligocene to the Early Pleistocene epochs. Its type species is Cosmopolitodus hastalis, the broad-tooth mako (other common names include the extinct giant mako and broad-tooth white shark). In 2021, Isurus planus was reassigned to the genus, and thus became the second species C. planus.^[3] However, some researchers still consider both species of Cosmopolitodus as species of Carcharodon .^[4]

Taxonomy

Etymology

Cosmopolitodus is derived from the Ancient Greek κοσμοπολίτης "kosmopolítēs" meaning "citizen of the world" and ὀδών "odṓn" meaning "tooth". The specific name hastalis may be derived from the Latin word hasta meaning "spear". The disputed species xiphodon is derived from the Ancient Greek ξίφος "xíphos" meaning "sword" and ὀδών "odṓn" meaning "tooth".

Taxonomic history

The taxonomy of C. hastalis, especially the status of its genus, has long been subject to debate. The initial binomial name was first given as Oxyrhina hastalis and Oxyrhina xiphodon for the narrow and broad-form variations respectively by Swiss naturalist Louis Agassiz in his 1843 paper Recherches sur les poissons fossiles, although some indications show that he had coined the taxon as early as 1838.^[8] Throughout the early and mid 20th century, different genera and species of other lamniformes would be clumped into the two taxa as the genus Oxyrhina began to be used as a wastebasket taxon. Sharks previously identified as variations of Oxyrhina hastalis or Oxyrhina xiphodon included two species of extinct ancestral makos, Isurus desori and Isurus retroflexus , the serrated mako ( Carcharomodus escheri ), and the false-toothed mako ( Parotodus benedenii ).^[9] Eventually, later studies would begin to show that much of the sharks within the genus and two species are distinct from each other, discarding the genus Oxyrhina altogether and creating another issue on what new taxa the sharks should be placed in. As of now, the genus is still uncertain and debated.^[8]

Shortly after the discarding of the genus Oxyrhina, a review by Holec et al. (1995) placed the species hastalis and xiphodon as a species of mako under the genus Isurus, citing the similarities between the teeth of the two and that of modern mako sharks.^[9] In 2001, a study by Ward & Bonavi (2001) considered Isurus hastalis and Isurus xiphodon as conspecific and declared the latter a nomen dubium .^[10] Although this conclusion is widely accepted, some scientists disagree, with a study by Whitenack and Gottfried (2010) demonstrating geometrically morphological differences between I. hastalis and I. xiphodon.^[11] The scientific name Isurus hastalis is considered as the "traditional view" in the debate regarding the shark's taxonomy.^[8]

Isurus schoutedeni

Isurus praecursor

Isurus desori

Cosmopolitodus hastalis

Narrow form

	Cosmopolitodus planus

	Carcharomodus escheri

Broad form

	Carcharodon hubbelli

	Carcharodon carcharias

Relationship between C. hastalis and its close relatives.

Ward & Bonavi (2001) reexamined the teeth of I. hastalis and noted a strong morphological similarity between it and the extant great white shark. The study concluded that I. hastalis is a direct ancestor of the great white and is more related to it than other makos. They moved the species into the genus Cosmopolitodus, which was a move that was first proposed by Glikman (1964) but was long rejected beforehand.^[10] Later discoveries supports Ward & Bonavi's conclusion of its ancestry to the great white. An analysis of a newly discovered Carcharodon hubbelli by Ehret et al. (2012) further cemented the theory of C. hastalis's ancestry to the great white, however, they also proposed that Cosmopolitodus should be moved to Carcharodon, remarking that the two genera were separated solely due to the lack of serrations in C. hastalis and the lack of lateral cusplets in C. carcharias and pointing out examples of Late Miocene C. hastalis teeth showing basal serrations.^[12] Cione et al. (2012) also noted a possibility of moving all species in the genus Cosmopolitodus into the genus Carcharodon to avoid a possible paraphyly that would occur if one of them, which was traditionally identified as C. xiphodon, is a putative sister species of C. carcharias. However, the study also noted that this would only be done if the putative sister species is conspecific with C. hastalis. The study concluded that there is indeed a putative sister species of C. carcharias distinct from C. hastalis and proposed the taxon Carcharodon plicatilis for it, resolving the paraphyly issue.^[13]

However, Yun argued that the tooth fossil remains of Cosmopolitodus and the great white "have been documented from the same deposits, hence the former cannot be a chronospecific ancestor of the latter." He also criticized the fact that the "Cosmopolitodus morphotype has never been tested through phylogenetic analyses," and denoted that as of 2021, the argument that the Carcharodon lineage with narrow, serrated teeth evolved from Cosmopolitodus with broad, unserrated teeth is uncertain.^[3]^[14]

Description

C. hastalis teeth can grow up to 8.9 cm (3.5 in) in length, suggesting a very large shark. Its body was probably very similar to that of modern great whites. It is also believed to have a cosmopolitan distribution, with C. hastalis teeth being found worldwide.^[15] The species is divided into two forms based on tooth morphology, each with a unique evolutionary line. The maximum adult length is estimated between 5 and 7 m (16 and 23 ft).^[16]^[17] Smaller individuals were about 2.6–4.5 metres (8.5–14.8 ft) long.^[18]

Teeth

C. planus teeth are somewhat similar to those of other mako sharks, especially the extinct species C. hastalis and Isurus desori . Adult upper teeth are generally 2–4 cm long with an unserrated cutting edge and no lateral cusplets. True to its name, each tooth is "hooked", the point of which is shifted away from the middle axis, in the direction of the corners of the mouth. In this, the teeth of C. planus and I. desori differ from C. hastalis. I. desori teeth are much more heterodontic than C. planus and their roots are more uneven in thickness and shape, with more narrow builds and rounder lobes, somewhat making the appearance of a heart. But the crowns of the lower teeth of this species are described to be more straight and identical to the lower teeth of C. hastalis.^[19]^[20] The hooked teeth, which probably would be efficient for gripping prey, would suggest a diet of smaller and medium-sized animals.

Broad-form

A broad form C. hastalis tooth

A narrow form C. hastalis tooth

The broad form is characterized by broad-shaped teeth often described as being identical to that of modern great whites besides the lack of serrations.^[15] Fossil evidence shows that the broad-form is the direct ancestor of the genus Carcharodon , and specimens from the Pisco Formation in Peru show an evolutionary mosaic between them.^[12]

This form is also commonly labeled by its species synonym xiphodon to reflect its difference from the narrow-form, although this is scientifically invalid.^[21]

Narrow-form

The narrow form has more slender teeth than its broad-form counterpart, which resemble broadened Isurus oxyrhinchus teeth. Unlike the broad-form, the narrow form is believed to be the ancestor of two extinct sharks, the hooked-tooth "mako" (Cosmopolitodus/Isurus planus), and the serrated "mako" (Carcharomodus escheri)^[15]

Paleobiology

Hunting behavior

C. hastalis was a confirmed hunter of marine mammals. It most likely would have been one of the major predators in its ecosystem, preying upon small whales and other mammals. Trace fossils in the form of tooth marks on the bones of a Pliocene dolphin of the species Astadelphis gastaldii reveal that C. hastalis attacked its prey from below and behind, much like the modern great white shark does. The deepest bite marks on the dolphin's ribs indicate the shark aimed for the abdomen of its prey to inflict a fatal bite quickly and incapacitate its prey, and that when the dolphin was attacked a second time, it was bitten near the dorsal fin, suggesting that the dolphin rolled over while injured. The size of the bites indicates further that the shark responsible was estimated to be 4 m (13 ft) long.^[22]

A well-preserved skeleton of a juvenile C. hastalis from Peru described in 2017 showcases that the shark also included fish in its diet. Stomach contents preserved with the fossil show this shark also fed on pilchards such as ancient sardines (genus Sardinops ), suggesting bony fish featured prominently in the shark's diet while it was young.^[23]

Related Research Articles

The great white shark, also known as the white shark, white pointer, or simply great white, is a species of large mackerel shark which can be found in the coastal surface waters of all the major oceans. It is the only known surviving species of its genus Carcharodon. The great white shark is notable for its size, with the largest preserved female specimen measuring 5.83 m (19.1 ft) in length and around 2,000 kg (4,400 lb) in weight at maturity. However, most are smaller; males measure 3.4 to 4.0 m, and females measure 4.6 to 4.9 m on average. According to a 2014 study, the lifespan of great white sharks is estimated to be as long as 70 years or more, well above previous estimates, making it one of the longest lived cartilaginous fishes currently known. According to the same study, male great white sharks take 26 years to reach sexual maturity, while the females take 33 years to be ready to produce offspring. Great white sharks can swim at speeds of 25 km/h (16 mph) for short bursts and to depths of 1,200 m (3,900 ft).

The Lamniformes are an order of sharks commonly known as mackerel sharks. It includes some of the most familiar species of sharks, such as the great white as well as less familiar ones, such as the goblin shark and megamouth shark.

Carcharodon is a genus of sharks within the family Lamnidae, colloquially called the "white sharks." The only extant member is the great white shark. The extant species was preceded by a number of fossil (extinct) species including C. hubbelli and C. hastalis. The first appearance of the genus may have been as early as the Early Miocene or Late Oligocene. Carcharocles megalodon is still argued by some paleontologists to be a close relative of Carcharodon carcharias - as well as being in the same genus. The megalodon's scientific name was originally "Carcharodon" megalodon, but more recently, the giant shark has been assigned by most scientists to either the genus Carcharocles or Otodus.

Otodus megalodon, commonly known as megalodon, is an extinct species of giant mackerel shark that lived approximately 23 to 3.6 million years ago (Mya), from the Early Miocene to the Pliocene epochs. O. megalodon was formerly thought to be a member of the family Lamnidae and a close relative of the great white shark, but has been reclassified into the extinct family Otodontidae, which diverged from the great white shark during the Early Cretaceous.

The Lamnidae are the family of mackerel sharks known as white sharks. They are large, fast-swimming predatory fish found in oceans worldwide, though they prefer environments with colder water. The name of the family is formed from the Greek word lamna, which means "fish of prey", and was derived from the Greek legendary creature, the Lamia.

Cretoxyrhina is an extinct genus of large mackerel shark that lived about 107 to 73 million years ago during the late Albian to late Campanian of the Late Cretaceous. The type species, C. mantelli, is more commonly referred to as the Ginsu shark, first popularized in reference to the Ginsu knife, as its theoretical feeding mechanism is often compared with the "slicing and dicing" when one uses the knife. Cretoxyrhina is traditionally classified as the likely sole member of the family Cretoxyrhinidae but other taxonomic placements have been proposed, such as within the Alopiidae and Lamnidae.

The longfin mako shark is a species of mackerel shark in the family Lamnidae, with a probable worldwide distribution in temperate and tropical waters. An uncommon species, it is typically lumped together under the name "mako" with its better-known relative, the shortfin mako shark. The longfin mako is a pelagic species found in moderately deep water, having been reported to a depth of 220 m (720 ft). Growing to a maximum length of 4.3 m (14 ft), the slimmer build and long, broad pectoral fins of this shark suggest that it is a slower and less active swimmer than the shortfin mako.

Otodus is an extinct, cosmopolitan genus of mackerel shark which lived from the Paleocene to the Pliocene epoch. The name Otodus comes from Ancient Greek ὠτ- and ὀδούς – thus, "ear-shaped tooth".

Isurus is a genus of mackerel sharks in the family Lamnidae, commonly known as the mako sharks. They are largely pelagic, and are fast, predatory fish capable of swimming at speeds of up to 50 km/h (31 mph).

Otodus angustidens is an extinct species of prehistoric megatoothed sharks in the genus Otodus, which lived during the Late Eocene and Miocene epochs about 34 to 21 million years ago. The largest individuals were about 11–12 metres (36–39 ft) long. This shark is related to another extinct megatoothed shark, the famous Otodus megalodon.

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<i>Carcharodon hubbelli</i> Extinct species of shark

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References

↑ Klug, S.; Kriwet, J., "Skeletal remains of the Miocene lamniform shark, Cosmopolitodus hastalis, from Peru" (PDF), Erlanger Geologische Abhandlungen, 6: 99
↑ Ebersole, J.A.; Ebersole, S.M.; Cicimurri, D.J. (2017). "The occurrence of early Pleistocene marine fish remains from the Gulf Coast of Mobile County, Alabama, USA". Palaeodiversity. 10 (1): 97–115. doi: 10.18476/pale.v10.a6 . S2CID 134476316.
1 2 3 Yun, C. (2021). "A tooth of the extinct lamnid shark, Cosmopolitodus planus comb. nov. (Chondrichthyes, Elasmobranchii) from the Miocene of Pohang City, South Korea" (PDF). Acta Palaeontologica Romaniae. 18 (1): 9–16. doi:10.35463/j.apr.2022.01.02. S2CID 242113412.
↑ Zevallos-Valdivia, L.; Martínez-Pérez, C.; García-Flores, V.; Chávez-Valencia, A.; Botella, H. (2023). "First record of palaeozoic vertebrates from Peru". Spanish Journal of Palaeontology. 38 (1): 95–100. doi: 10.7203/sjp.25691 . S2CID 256585278.
↑ "Carcharodon hastalis Agassiz 1843 (white shark)". Fossilworks. Retrieved December 17, 2021.
↑ Crumpton, Nick (November 14, 2012). "Great whites 'not from megashark'". BBC News.
↑ Dana J. Ehret; Bruce J. Macfadden; Douglas S. Jones; Thomas J. Devries; David A. Foster; Rodolfo Salas-Gismondi (2012). "Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru". Palaeontology. 55 (6): 1139–1153. https://doi.org/10.1111/j.1475-4983.2012.01201.x.
1 2 3 "A Well Preserved Skeleton of the Fossil Shark Cosmopolitodus Hastalis from the Late Miocene of Peru, Featuring Fish Remains as Fossilized Stomach Contents". Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy). 123 (1). 2017. doi:10.13130/2039-4942/8005. Archived from the original on June 1, 2022. Retrieved August 8, 2019.
1 2 Kocsis, László (February 2007). "Central Paratethyan shark fauna (Ipolytarnóc, Hungary)". Geologica Carpathica. 58: 27–40.
1 2 "Additions to and a review of the Miocene Shark and Ray fauna of Malta". ResearchGate.
↑ Whitenack †, Lisa B.; Gottfried, Michael D. (2010). "A morphometric approach for addressing toothbased species delimitation in fossil mako sharks, Isurus (Elasmobranchii: Lamniformes)". Journal of Vertebrate Paleontology. 30 (1): 17–25. Bibcode:2010JVPal..30...17W. doi: 10.1080/02724630903409055 . S2CID 85821523.
1 2 Ehret, Dana J.; MacFadden, Bruce J.; Jones, Douglas S.; Devries, Thomas J.; Foster, David A.; Salas-Gismondi, Rodolfo (2012). "Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru". Palaeontology. 55 (6): 1139–1153. Bibcode:2012Palgy..55.1139E. doi: 10.1111/j.1475-4983.2012.01201.x . S2CID 128666594.
↑ Cione, Alberto Luis; Cabrera, Daniel Alfredo; Barla, María Julia (2012). "Oldest record of the Great White Shark (Lamnidae, Carcharodon; Miocene) in the Southern Atlantic". Geobios. 45 (2): 167–172. Bibcode:2012Geobi..45..167C. doi:10.1016/j.geobios.2011.06.002. hdl: 11336/139163 .
↑ Yun, Chan-gyu (August 7, 2020). "New example of Cosmopolitodus hastalis (Lamniformes, Lamnidae) from the Miocene of South Korea | Zoodiversity". Zoodiversity. 54 (5): 433–438. doi: 10.15407/zoo2020.05.433 .
1 2 3 "Great White Sharks and Prehistoric White Sharks".
↑ Patrick L. Jambura; Julia Türtscher; Faviel A. López-Romero; Catalina Pimiento; Jürgen Kriwet (2019). On the origin of the great white shark Carcharodon carcharias.
↑ Pimiento, C.; Cantalapiedra, J.L.; Shimada, K.; Field, D.J.; Smaers, J.B. (2019). "Evolutionary pathways toward gigantism in sharks and rays" (PDF). Evolution. 73 (3): 588–599. doi:10.1111/evo.13680. ISSN 1558-5646. PMID 30675721. S2CID 59224442.
↑ McCormack, Jeremy; Griffiths, Michael L.; Kim, Sora L.; Shimada, Kenshu; Karnes, Molly; Maisch, Harry; Pederzani, Sarah; Bourgon, Nicolas; Jaouen, Klervia; Becker, Martin A.; Jöns, Niels (May 31, 2022). "Trophic position of Otodus megalodon and great white sharks through time revealed by zinc isotopes". Nature Communications. 13 (1): 2980. Bibcode:2022NatCo..13.2980M. doi:10.1038/s41467-022-30528-9. ISSN 2041-1723. PMC 9156768 . PMID 35641494. S2CID 249235478.
↑ Cocke, Joe (2002). Fossil Shark Teeth of the World: A collector's guide.
↑ Ciampaglio, C., Ricketts,D. Miocene Madness: Fossil Finder ID and Information Guide. Bakersfield, California. Buena Vista Museum of Natural History.
↑ "Great White Lineage". elasmo.com. Retrieved April 20, 2022.
↑ "Jaws -- 4 million BC: How an extinct shark attacked its prey".
↑ Collareta, Alberto; Landini, Walter; Chacaltana Budiel, César Augusto; Valdivia Vera, Waldir; Altamirano Sierra, Ali J.; Urbina Schmitt, Mario; Bianucci, Giovanni (March 2017). "A well preserved skeleton of the fossil shark Cosmopolitodus hastalis from the late Miocene of Peru, featuring fish remains as fossilized stomach contents". Repositorio Institucional Ingemmet.

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[Klug-1] Klug, S.; Kriwet, J., "Skeletal remains of the Miocene lamniform shark, Cosmopolitodus hastalis, from Peru" (PDF), Erlanger Geologische Abhandlungen, 6: 99

[Ebersoleetal-2] Ebersole, J.A.; Ebersole, S.M.; Cicimurri, D.J. (2017). "The occurrence of early Pleistocene marine fish remains from the Gulf Coast of Mobile County, Alabama, USA". Palaeodiversity. 10 (1): 97–115. doi: 10.18476/pale.v10.a6 . S2CID 134476316.

[Yun-3] 1 2 3 Yun, C. (2021). "A tooth of the extinct lamnid shark, Cosmopolitodus planus comb. nov. (Chondrichthyes, Elasmobranchii) from the Miocene of Pohang City, South Korea" (PDF). Acta Palaeontologica Romaniae. 18 (1): 9–16. doi:10.35463/j.apr.2022.01.02. S2CID 242113412.

[4] Zevallos-Valdivia, L.; Martínez-Pérez, C.; García-Flores, V.; Chávez-Valencia, A.; Botella, H. (2023). "First record of palaeozoic vertebrates from Peru". Spanish Journal of Palaeontology. 38 (1): 95–100. doi: 10.7203/sjp.25691 . S2CID 256585278.

[fossilworks-5] "Carcharodon hastalis Agassiz 1843 (white shark)". Fossilworks. Retrieved December 17, 2021.

[6] Crumpton, Nick (November 14, 2012). "Great whites 'not from megashark'". BBC News.

[7] Dana J. Ehret; Bruce J. Macfadden; Douglas S. Jones; Thomas J. Devries; David A. Foster; Rodolfo Salas-Gismondi (2012). "Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru". Palaeontology. 55 (6): 1139–1153. https://doi.org/10.1111/j.1475-4983.2012.01201.x.

[COLLARETA-8] 1 2 3 "A Well Preserved Skeleton of the Fossil Shark Cosmopolitodus Hastalis from the Late Miocene of Peru, Featuring Fish Remains as Fossilized Stomach Contents". Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy). 123 (1). 2017. doi:10.13130/2039-4942/8005. Archived from the original on June 1, 2022. Retrieved August 8, 2019.

[Hungary-9] 1 2 Kocsis, László (February 2007). "Central Paratethyan shark fauna (Ipolytarnóc, Hungary)". Geologica Carpathica. 58: 27–40.

[Ward-10] 1 2 "Additions to and a review of the Miocene Shark and Ray fauna of Malta". ResearchGate.

[Gottfreid-11] Whitenack †, Lisa B.; Gottfried, Michael D. (2010). "A morphometric approach for addressing toothbased species delimitation in fossil mako sharks, Isurus (Elasmobranchii: Lamniformes)". Journal of Vertebrate Paleontology. 30 (1): 17–25. Bibcode:2010JVPal..30...17W. doi: 10.1080/02724630903409055 . S2CID 85821523.

[Ehret-12] 1 2 Ehret, Dana J.; MacFadden, Bruce J.; Jones, Douglas S.; Devries, Thomas J.; Foster, David A.; Salas-Gismondi, Rodolfo (2012). "Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru". Palaeontology. 55 (6): 1139–1153. Bibcode:2012Palgy..55.1139E. doi: 10.1111/j.1475-4983.2012.01201.x . S2CID 128666594.

[Coine-13] Cione, Alberto Luis; Cabrera, Daniel Alfredo; Barla, María Julia (2012). "Oldest record of the Great White Shark (Lamnidae, Carcharodon; Miocene) in the Southern Atlantic". Geobios. 45 (2): 167–172. Bibcode:2012Geobi..45..167C. doi:10.1016/j.geobios.2011.06.002. hdl: 11336/139163 .

[14] Yun, Chan-gyu (August 7, 2020). "New example of Cosmopolitodus hastalis (Lamniformes, Lamnidae) from the Miocene of South Korea | Zoodiversity". Zoodiversity. 54 (5): 433–438. doi: 10.15407/zoo2020.05.433 .

[Fossilguy-15] 1 2 3 "Great White Sharks and Prehistoric White Sharks".

[16] Patrick L. Jambura; Julia Türtscher; Faviel A. López-Romero; Catalina Pimiento; Jürgen Kriwet (2019). On the origin of the great white shark Carcharodon carcharias.

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Taxon identifiers
Cosmopolitodus	Wikidata: Q20686304 GBIF: 4824543 IRMNG: 1223316 Paleobiology Database: 118027
Carcharodon hastalis	Wikidata: Q3803933 GBIF: 5816397 IRMNG: 11083619 Paleobiology Database: 265174

Cosmopolitodus Temporal range: Rupelian-Calabrian, 30–1.07 Ma PreꞒ Ꞓ O S D C P T J K Pg N ^[1]^[2]

Tooth series of C. hastalis
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Chondrichthyes
Subclass:	Elasmobranchii
Subdivision:	Selachimorpha
Order:	Lamniformes
Family:	Lamnidae
Genus:	† Cosmopolitodus Glikman, 1964
Type species
†Cosmopolitodus hastalis Agassiz, 1843
Species
†C. hastalis(Agassiz, 1843) †C. planusAgassiz, 1856
Synonyms
Carcharodon hastalis(Agassiz, 1843) Isurus hastalis(Agassiz, 1843) Isurus xiphodon(Agassiz, 1843) Oxyrhina hastalis(Agassiz, 1843) Oxyrhina xiphodon(Agassiz, 1843) Oxyrhina plana(Agassiz, 1856) Isurus planus(Jordan, 1907)