User:Macrophyseter/sandbox11

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Classification[edit]

Prescientific and early research history[edit]

Left image: Chesapeake megalodon tooth (Fig. 7) excavated from a Hopewell burial mound. Right image: Colonna's 1616 comparison of a megalodon (top left) and great white tooth (right).

Megalodon teeth have been excavated and used since ancient times. They were a valued artifact amongst pre-Columbian cultures in the Americas for their large sizes and serrated blades, from which they were modified into projectile points, knives, jewelry, and funeral accessories.[1][2] At least some, such as the Panamanian Sitio Conte societies, seemed to have used them for primarily ceremonial purposes.[2] Mining[3] of megalodon teeth by the Algonquin peoples in the Chesapeake Bay and their selective trade with the Adena culture in Ohio occurred since as early as 430 BC.[1] The earliest written account of megalodon teeth was by Pliny the Elder in an AD 73 volume of Historia Naturalis, who described them as resembling petrified human tongues that Roman folklorists believed to have fallen from the sky during lunar eclipses and called them glossopetrae ("tongue stones").[4] The purported tongues were later thought in a 12th-century Maltese tradition to have belonged to serpents that Paul the Apostle turned to stone while shipwrecked there, and were given antivenom powers by the saint.[5] Glossopetrae reappeared throughout Europe in late 13th to 16th century literature, ascribed with more supernatural properties that cured a wider variety of poisons. Use of megalodon teeth for this purpose became widespread among medieval and Renaissance nobility, who fashioned them into protective amulets and tableware to purportedly detoxify poisoned liquids or bodies that touched the stones. By the 16th century, teeth were directly consumed as ingredients of European-made Goa stones.[4]

An illustration of a shark head (sideview). Visible are wrinkles and an exaggerated nose and eyes, and at the bottom are two individual drawings of shark teeth
Mercati's depictions of a great white's head and teeth (left) and fossil megalodon and great white teeth (right), reused by Steensen in 1667

The true nature of the glossopetrae as shark's teeth was held by some since at least 1554, when cosmographer André Thevet described hearsay about the belief, although he did not believe them. The earliest scientific argument for this view was made by Italian naturalist Fabio Colonna, who in 1616 published an illustration of a Maltese megalodon tooth alongside a great white shark's and noted their striking similarities. He argued that the former and its likenesses were not petrified serpent's tongues but actually the teeth of similar sharks that washed up on shore. Colonna supported this thesis through an experiment of burning glossopetrae samples, from which he observed carbon residue he interpreted as proving an organic origin.[5] However, interpretations of the stones as shark's teeth remained widely unaccepted at the time. This was in part due the inability to explain how some of them are found far from the sea.[6] The shark tooth argument was academically raised again during the late 17th century by English scientists Robert Hooke, John Ray, and Danish naturalist Niels Steensen (Latinized Nicholas Steno).[7] Steensen's argument in particular is most recognized today as it was inferred from his dissection of the head of a great white caught in 1666. His 1667 report depicted engravings of a shark's head and megalodon teeth that became especially iconic. However, the illustrated head was not actually the head that Steensen dissected, nor were the fossil teeth illustrated by him. Both engravings were originally commissioned in the 1590s by Papal physician Michele Mercati, who also had in possession the head of a great white, for his book Metallotheca. The work remained unpublished in Steensen's time due to Mercati's premature death, and the former reused the two illustrations per suggestion by Carlo Roberto Dati, who thought a depiction of the actual dissected shark was unsuitable for readers.[8] Steensen also stood out in pioneering a stratigraphic explanation for how similar stones appeared further inland. He observed that rock layers bearing megalodon teeth contained marine sediments and hypothesized that these layers correlated to a period of flood that was later covered by terrestrial layers and uplifted by geologic activity.[6]

Holotype tooth (Fig. 2-3) in Agassiz (1835), now cataloged as TE-PLI 18

Swiss naturalist Louis Agassiz gave megalodon its scientific name in his seminal 1833-1843 work Recherches sur les poissons fossiles (Research on fossil fish). He initially named it Carcharias megalodon in an 1835 illustration of the holotype and additional teeth, congeneric with the modern sand tiger shark.[9][10] The specific name is a portmanteau of the Ancient Greek words μεγάλος (megálos, meaning "big") and ὀδών (odṓn, meaning "tooth"),[11][12] combined meaning "big tooth." Agassiz referenced the name as early as 1832, but because specimens were not referenced they are not taxonomically recognized uses.[10] Formal description of the species was published in an 1843 volume, where Agassiz revised the name to Carcharodon megalodon as its teeth were far too large for the former genus and more alike to the great white shark.[9] He also erroneously identified several megalodon teeth as belonging to additional species eventually named Carcharodon rectidens, Carcharodon subauriculatus, Carcharodon productus, and Carcharodon polygurus.[9][13] Because Carcharodon megalodon appeared first in the 1835 illustration, the remaining names are considered junior synonyms under the principle of priority.[10][13]

Evolution[edit]

Megalodon and its serrated predecessors are classified within the Otodontidae, their placement inspiring the family's vernacular name "megatoothed sharks." This is based on the consensus that the lineage arose from Otodus obliquus, another large shark that lived during the Paleocene and Eocene. The generic name for Megalodon has likewise coalesced around this ancestor's genus in recent years, making the full scientific name Otodus megalodon. However, the exact sequence in which Megalodon evolved from O. obliquus remains controversial, which muddles a sensitive part of evolution-based classification that affects which genus Megalodon should actually be assigned to. A review of the situation by Kent (2018) summarized three traditional models of evolution:

  • The archaic Carcharodon model held that Megalodon is actually a member of the white sharks (Lamnidae) and closely related to the modern great white, either as a direct ancestor or sister species. Once prominent until the 2010s, this model is now universally rejected by paleontologists. This is following the 2012 description of Carcharodon hubbelli, a transitional species that demonstrated the great white's descent from a separate lineage that diverged from the otodontids during the Early Cretaceous.
  • The Carcharocles model upholds Megalodon's classification as an otodontid.
  • The Otodus model is an extension of the Carcharocles model first proposed in 1997. It specifies that the lineage between O. obliquus and Megalodon evolved via anagenesis, representing a single unbranching line of chronospecies that gradually evolved over 50 million years. Because of the inherent difficulty of delineating genera and species within a gradually changing lineage, classification of Megalodon under this model underwent several variations. Cappetta (2006) thought that the appearance of serrations is insufficient to warrant a distinct genus and instead moved the splitting line to the disappearance of strong lateral cusplets, which required Megalodon to be reclassified under the genus Megaselachus as Carcharocles was no longer available in this case. Cappetta (2012) revised this, moving Megalodon directly into Otodus but retaining Megaselachus as a subgenus. Shimada et al. (2017) found that classifying Megalodon into a separate genus breaks a classification rule in cladistics that requires genera to contain all members of an evolutionary clade; Otodus would break this rule by containing O. obliquus and some of its ancestors, but not its descendants, unless all former Carcharocles species are reassigned into it.


It assumes that O. obliquus is an intermediate between older species of Otodus described in Russia and Central Asia

Junk[edit]

Naming[edit]

According to Renaissance accounts, gigantic triangular fossil teeth often found embedded in rocky formations were once believed to be the petrified tongues, or glossopetrae, of dragons and snakes. This interpretation was corrected in 1667 by Danish naturalist Nicolas Steno, who recognized them as shark teeth, and famously produced a depiction of a shark's head bearing such teeth. He described his findings in the book The Head of a Shark Dissected, which also contained an illustration of a megalodon tooth.[14][15][16]

Swiss naturalist Louis Agassiz gave this shark its initial scientific name, Carcharodon megalodon, in his 1843 work Recherches sur les poissons fossiles, based on tooth remains.[17][18] English paleontologist Edward Charlesworth in his 1837 paper used the name Carcharias megalodon, while citing Agassiz as the author, indicating that Agassiz described the species prior to 1843. English paleontologist Charles Davies Sherborn in 1928 listed an 1835 series of articles by Agassiz as the first scientific description of the shark.[19] The specific name megalodon translates to "big tooth", from Ancient Greek: μέγας, romanized: mégas, lit. 'big, mighty' and ὀδούς, odoús, 'tooth'.[20][21] The teeth of megalodon are morphologically similar to those of the great white shark (Carcharodon carcharias), and on the basis of this observation, Agassiz assigned megalodon to the genus Carcharodon.[18]

There was one apparent description of the shark in 1881 classifying it as Selache manzonii.[22]

Evolution[edit]

Diagram of the chronospecies evolution of megalodon

While the earliest megalodon remains have been reported from the Late Oligocene, around 28 million years ago (Mya),[23][24] there is disagreement as to when it appeared, with dates ranging to as young as 16 mya.[25] It has been thought that megalodon became extinct around the end of the Pliocene, about 2.6 Mya;[25][26] claims of Pleistocene megalodon teeth, younger than 2.6 million years old, are considered unreliable.[26] A 2019 assessment moves the extinction date back to earlier in the Pliocene, 3.6 Mya.[27][28]

Megalodon is now considered to be a member of the family Otodontidae, genus Otodus, as opposed to its previous classification into Lamnidae, genus Carcharodon.[25][29][26][30][31] Megalodon's classification into Carcharodon was due to dental similarity with the great white shark, but most authors currently believe that this is due to convergent evolution. In this model, the great white shark is more closely related to the extinct broad-toothed mako (Isurus hastalis) than to megalodon, as evidenced by more similar dentition in those two sharks; megalodon teeth have much finer serrations than great white shark teeth. The great white shark is more closely related to the mako shark (Isurus spp.), with a common ancestor around 4 Mya.[18][32] Proponents of the former model, wherein megalodon and the great white shark are more closely related, argue that the differences between their dentition are minute and obscure.[33]: 23–25 

A black megalodon tooth and two white great white shark teeth above a centimeter scale, the megalodon tooth extends between the zero and thirteen-and-a-half centimeter marks. One great white tooth extends between the eleven and thirteen centimeter marks, and the other extends between from the thirteen and sixteen centimeter marks.
Megalodon tooth with two great white shark teeth

The genus Carcharocles currently contains four species: C. auriculatus, C. angustidens, C. chubutensis, and C. megalodon.[34]: 30–31  The evolution of this lineage is characterized by the increase of serrations, the widening of the crown, the development of a more triangular shape, and the disappearance of the lateral cusps.[34]: 28–31 [35] The evolution in tooth morphology reflects a shift in predation tactics from a tearing-grasping bite to a cutting bite, likely reflecting a shift in prey choice from fish to cetaceans.[36] Lateral cusplets were finally lost in a gradual process that took roughly 12 million years during the transition between C. chubutensis and C. megalodon.[36] The genus was proposed by D. S. Jordan and H. Hannibal in 1923 to contain C. auriculatus. In the 1980s, megalodon was assigned to Carcharocles.[18][34]: 30  Before this, in 1960, the genus Procarcharodon was erected by French ichthyologist Edgard Casier, which included those four sharks and was considered separate from the great white shark. It is now considered a junior synonym of Carcharocles.[34]: 30  The genus Palaeocarcharodon was erected alongside Procarcharodon to represent the beginning of the lineage, and, in the model wherein megalodon and the great white shark are closely related, their last common ancestor. It is believed to be an evolutionary dead-end and unrelated to the Carcharocles sharks by authors who reject that model.[33]: 70 

A great white shark swimming a few meters below the surface, above a school of much smaller fish.
The great white shark (Carcharodon carcharias) and megalodon were previously thought to be close relatives.[18][32]

Another model of the evolution of this genus, also proposed by Casier in 1960, is that the direct ancestor of the Carcharocles is the shark Otodus obliquus, which lived from the Paleocene through the Miocene epochs, 60 to 13 Mya.[32][35] The genus Otodus is ultimately derived from Cretolamna, a shark from the Cretaceous period.[30][37] In this model, O. obliquus evolved into O. aksuaticus, which evolved into C. auriculatus, and then into C. angustidens, and then into C. chubutensis, and then finally into C. megalodon.

Another model of the evolution of Carcharocles, proposed in 2001 by paleontologist Michael Benton, is that the three other species are actually a single species of shark that gradually changed over time between the Paleocene and the Pliocene, making it a chronospecies.[34]: 17 [24][38] Some authors suggest that C. auriculatus, C. angustidens, and C. chubutensis should be classified as a single species in the genus Otodus, leaving C. megalodon the sole member of Carcharocles.[24][39]

The genus Carcharocles may be invalid, and the shark may actually belong in the genus Otodus, making it Otodus megalodon.[40] A 1974 study on Paleogene sharks by Henri Cappetta erected the subgenus Megaselachus, classifying the shark as Otodus (Megaselachus) megalodon, along with O. (M.) chubutensis. A 2006 review of Chondrichthyes elevated Megaselachus to genus, and classified the sharks as Megaselachus megalodon and M. chubutensis.[40] The discovery of fossils assigned to the genus Megalolamna in 2016 led to a re-evaluation of Otodus, which concluded that it is paraphyletic, that is, it consists of a last common ancestor but it does not include all of its descendants. The inclusion of the Carcharocles sharks in Otodus would make it monophyletic, with the sister clade being Megalolamna.[30]

The cladogram below represents the hypothetical relationships between megalodon and other sharks, including the great white shark. Modified from Shimada et al. (2016),[30] Ehret et al., (2009),[32] and the findings of Siversson et al. (2013).[41][42][43]

Lamniformes
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  2. ^ a b Farrell, A.D. (2021). A Use-Wear and Functional Analysis of Precontact Shark Teeth Assemblages from Florida (MS). Florida State University.
  3. ^ Jacobs, J. "Hopewell Culture National Historical Park Sharks Teeth HOCU - 2832 and 4222". National Park Service.
  4. ^ a b Duffin, C.J. (2017). "Fossil Sharks' Teeth as Alexipharmics". In Philip Wexler (ed.). Toxicology of the Middle Ages and Renaissance. pp. 125–133. doi:10.1016/B978-0-12-809554-6.00012-3.
  5. ^ a b Forli, M.; Guerrini, A. (2022). "Quaestio de Fossilibus: Glossopetres, Snake Tongues and Ceraunids". The History of Fossils Over Centuries. pp. 41–83. doi:10.1007/978-3-031-04687-2_6.
  6. ^ a b Bressan, D. (2016-01-14). "How the Dissection Of A Shark's Head Revealed The True Nature Of Fossils". Forbes.
  7. ^ "Nicholas Steno". University of California Museum of Paleontology.
  8. ^ Davidson, J.P. (2000). "Historical Point of View: Fish Tales: Attributing the First Illustration of a Fossil Shark's Tooth to Richard Verstegan (1605) and Nicolas Steno (1667)". Proceedings of the Academy of Natural Sciences of Philadelphia. 150: 329–344.
  9. ^ a b c Agassiz, L. (1843). Recherches sur les poissons fossiles (in French). Vol. 3. doi:10.5962/bhl.title.4275.
  10. ^ a b c Brignon, A. (2021). "Historical and nomenclatural remarks on some megatoothed shark teeth (Elasmobranchii, Otodontidae) from the Cenozoic of New Jersey (U.S.A.)". Rivista Italiana di Paleontologia e Stratigrafia. 127 (3): 595–625. doi:10.13130/2039-4942/16440.
  11. ^ μεγάλος. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
  12. ^ ὀδών. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
  13. ^ a b Woodward, A.S. (1899). Catalogue of the fossil fishes in the British Museum (Natural History), Part I. Containing the Elasmobranchii. British Museum (Natural History). pp. 415–420. doi:10.5962/bhl.title.61854.
  14. ^ Haven, Kendall (1997). 100 Greatest Science Discoveries of All Time. Westport, Connecticut: Libraries Unlimited. pp. 25–26. ISBN 978-1-59158-265-6. OCLC 230807846.
  15. ^ Hsu, Kuang-Tai (2009). "The Path to Steno's Synthesis on the Animal Origin of Glossopetrae". In Rosenburg, G. D. (ed.). The Revolution in Geology from the Renaissance to the Enlightenment. Vol. 203. Boulder, Colorado: Geological Society of America. ISBN 978-0-8137-1203-1. OCLC 608657795.
  16. ^ Eilperin, J. (2012). Demon Fish. Pantheon Books. p. 43. ISBN 978-0-7156-4352-5.
  17. ^ Agassiz, Louis (1843). Recherches sur les poissons fossiles [Research on the fossil fishes] (in French). Neuchatel: Petitpierre. p. 41.
  18. ^ a b c d e Nyberg, K. G.; Ciampaglio C. N.; Wray G. A. (2006). "Tracing the ancestry of the great white shark, Carcharodon carcharias, using morphometric analyses of fossil teeth". Journal of Vertebrate Paleontology. 26 (4): 806–814. doi:10.1671/0272-4634(2006)26[806:TTAOTG]2.0.CO;2. S2CID 53640614.
  19. ^ Keyes, I. W. (2012). "New records of the Elasmobranch C. megalodon (Agassiz) and a review of the genus Carcharodon in the New Zealand fossil record". New Zealand Journal of Geology and Geophysics. 15 (2): 229. doi:10.1080/00288306.1972.10421956.
  20. ^ μέγας. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
  21. ^ ὀδούς. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
  22. ^ Lawley, R. (1881). "Selache manzonii n. sp. – Dente Fossile délia Molassa Miocenica del Monte Titano (Repubblica di San Marino)" [Fossil tooth from Miocene Molasse from Monte Titano (Republic of San Marino)]. Atti della Società Toscana di Scienze Naturali (in Italian). 5: 167–172.
  23. ^ Yabe, H.; Goto, M.; Kaneko, N. (2004). "Age of Carcharocles megalodon (Lamniformes: Otodontidae): A review of the stratigraphic records". The Palaeontological Society of Japan. 75: 7–15.
  24. ^ a b c Gottfried, M. D.; Fordyce, R. E. (2001). "An associated specimen of Carcharodon angustidens (Chondrichthyes, Lamnidae) from the Late Oligocene of New Zealand, with comments on Carcharodon interrelationships". Journal of Vertebrate Paleontology. 21 (4): 730–739. doi:10.1671/0272-4634(2001)021[0730:AASOCA]2.0.CO;2. S2CID 86092645. Archived from the original on 17 December 2019. Retrieved 6 November 2017.
  25. ^ a b c Pimiento, C.; MacFadden, B. J.; Clements, C. F.; Varela, S.; Jaramillo, C.; Velez-Juarbe, J.; Silliman, B. R. (2016). "Geographical distribution patterns of Carcharocles megalodon over time reveal clues about extinction mechanisms". Journal of Biogeography. 43 (8): 1645–1655. doi:10.1111/jbi.12754. S2CID 55776834.
  26. ^ a b c Pimiento, C.; Clements, C. F. (2014). "When Did Carcharocles megalodon Become Extinct? A New Analysis of the Fossil Record". PLOS ONE. 9 (10): e111086. Bibcode:2014PLoSO...9k1086P. doi:10.1371/journal.pone.0111086. PMC 4206505. PMID 25338197.
  27. ^ Cite error: The named reference Boessenecker2019 was invoked but never defined (see the help page).
  28. ^ "Giant 'megalodon' shark extinct earlier than previously thought". Science Daily. 13 February 2019.
  29. ^ Cite error: The named reference Pimiento&Balk2015 was invoked but never defined (see the help page).
  30. ^ a b c d Shimada, K.; Chandler, R. E.; Lam, O. L. T.; Tanaka, T.; Ward, D. J. (2016). "A new elusive otodontid shark (Lamniformes: Otodontidae) from the lower Miocene, and comments on the taxonomy of otodontid genera, including the 'megatoothed' clade". Historical Biology. 29 (5): 1–11. doi:10.1080/08912963.2016.1236795. S2CID 89080495.
  31. ^ Shimada, Kenshu (2019). "The size of the megatooth shark, Otodus megalodon (Lamniformes: Otodontidae), revisited". Historical Biology. 33 (7): 1–8. doi:10.1080/08912963.2019.1666840. ISSN 0891-2963. S2CID 208570844.
  32. ^ a b c d Ehret D. J.; Hubbell G.; Macfadden B. J. (2009). "Exceptional preservation of the white shark Carcharodon from the early Pliocene of Peru". Journal of Vertebrate Paleontology. 29 (1): 1–13. doi:10.1671/039.029.0113. JSTOR 20491064. S2CID 129585445.
  33. ^ a b Klimley, Peter; Ainley, David (1996). "Evolution". Great White Sharks: The Biology of Carcharodon carcharias. San Diego, California: Academic Press. ISBN 978-0-12-415031-7. OCLC 212425118.
  34. ^ a b c d e Cite error: The named reference H was invoked but never defined (see the help page).
  35. ^ a b Cite error: The named reference C was invoked but never defined (see the help page).
  36. ^ a b Perez, V. J.; Godfrey, S. J.; Kent, B. W.; Weems, R. E.; Nance, J. R. (2019). "The transition between Carcharocles chubutensis and Carcharocles megalodon (Otodontidae, Chondrichthyes): lateral cusplet loss through time". Journal of Vertebrate Paleontology. 38 (6): e1546732. doi:10.1080/02724634.2018.1546732.
  37. ^ Siverson, M.; Lindgren, J.; Newbrey, M.G.; Cederström, P.; Cook, T.D. (2013). "Late Cretaceous (Cenomanian-Campanian) mid-palaeolatitude sharks of Cretalamna appendiculata type" (PDF). Acta Palaeontologica Polonica: 2. doi:10.4202/app.2012.0137. S2CID 58906204. Archived from the original (PDF) on 19 October 2013.
  38. ^ Benton, M. J.; Pearson, P. N. (2001). "Speciation in the fossil record". Trends in Ecology and Evolution. 16 (7): 405–411. doi:10.1016/s0169-5347(01)02149-8. PMID 11403874.
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  40. ^ a b Cappetta, H. (1987). "Mesozoic and Cenozoic Elasmobranchii". Handbook of Paleoichthyology. Vol. 3B. München, Germany: Friedrich Pfeil. ISBN 978-3-89937-046-1. OCLC 829906016.
  41. ^ Siversson, M.; Lindgren, J.; Newbrey, M.G.; Cederström, P.; Cook, T.D. (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. S2CID 58906204.
  42. ^ Vivian, G. (2013). "Research debunks Great White lineage".
  43. ^ "Palaeontological detective work unravels evolution of megatooth sharks". 2013.
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