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Gryposuchus

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Gryposuchus
Temporal range: Miocene,
16–5.3 Ma[1]
Fossils of the skull and mandible of G. colombianus, Museo Geológico José Royo y Gómez, Bogotá.
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Order: Crocodilia
Family: Gavialidae
Subfamily: Gryposuchinae
Genus: Gryposuchus
Gurich, 1912
Type species
Gryposuchus jessei
Gurich, 1912
Other species
  • G. neogaeus
    (Burmeister, 1885 [originally Rhamphostomopsis neogaeus])
  • G. colombianus
    Langston, 1965
  • G. croizati
    Riff & Aguilera, 2008
  • G. pachakamue
    Salas-Gismondi et al., 2016

Gryposuchus is an extinct genus of gavialid crocodilian. Fossils have been found from Argentina, Colombia, Venezuela, Brazil and the Peruvian Amazon. The genus existed during the Miocene epoch (Colhuehuapian to Huayquerian).[2] One recently described species, G. croizati, grew to an estimated length of 10 metres (33 ft). Gryposuchus is the type genus of the subfamily Gryposuchinae, although a 2018 study indicates that Gryposuchinae and Gryposuchus might be paraphyletic and rather an evolutionary grade towards the gharial.

Species

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The type species of Gryposuchus is G. jessei, named based on a well-preserved rostrum collected along the Pauini River of Brazil in 1912. The specimen was probably destroyed during World War II by the 1943 bombing of Hamburg.[3] Another specimen named UFAC 1272, consisting of a premaxilla and maxilla, was discovered in the nearby Sena Madureia locality of the late Miocene Solimões Formation,[2] in and referred to the species in 1997.[4] Gryposuchus jessei is also referred to from the Urumaco Formation of northwestern Venezuela.[2] A second species, G. neogaeus, was referred to the genus in 1982; specimens from this species were first described from the late Miocene Ituzaingó Formation of Argentina in 1885,[2][5] although it was referred to Rhamphostomopsis at the time.[3][6][7]

Another species, G. colombianus, has been recovered from deposits from the Middle Miocene Honda Group of Colombia, and the late Miocene Urumaco Formation in Venezuela.[2] This species, named in 1965, was originally referred to Gavialis.[8] Fragmentary material of Gryposuchus from the Fitzcarrald Arch in the Peruvian Amazon dating back to the late middle Miocene bear a close resemblance to G. colombianus, but differ in rostrum proportions.[2][9] G. neogaeus and G. colombianus have been proposed as synonyms of G. jessei,[10] but this is unlikely due to the number of anatomical differences between them.[3][4]

Scale diagram showing the size of G. croizati (light blue)
Gryposuchus is located in South America
Gryposuchus
Gryposuchus
Gryposuchus
Gryposuchus
Gryposuchus
Gryposuchus fossil localities

A species described in 2008, G. croizati, also found from the upper Miocene Urumaco Formation in Venezuela,[2] can be distinguished from other species of Gryposuchus on the basis of a reduced number of maxillary teeth, a slender parietal interfenestral bar, and widely separated and reduced palatine fenestrae, and other characters. Based on measurements of the orbital cranial skeleton, the length of the animal has been estimated at around 10.15 metres (33.3 ft) in length, with a total mass of about 1,745 kilograms (3,847 lb). Measuring the entire length of the skull from the end of the rostrum to the supraoccipital would result in a much larger size estimate, up to three times as great. However, because there is considerable variation seen in rostral proportions among crocodilians, the latter measurements are probably not an accurate way of estimating body mass and length.[11] Despite this, the species is still one of the largest crocodilians known to have existed, and it may have been the second largest gavialoid to have ever existed if a recent revision in the estimated size of the large gharial relative Rhamphosuchus is correct (the genus was once considered to be 15 metres (49 ft) in length; the new estimate puts it at approximately 11 metres (36 ft)).[12]

Some skull material also recovered from Peruvian Amazon (Iquitos) in the Pebas Formation of the Middle Miocene,[2] was named as Gryposuchus pachakamue in 2016 by Rodolfo Salas-Gismondi et al. It includes the holotype MUSM 987, a well preserved skull that lacks of temporal and occipital bones; it measures 623.2 millimeters in length, and a series of referred specimens, including possible juveniles. The species was named after the Quechuan word for a primordial god and "storyteller".[13] This new species is characterized by have 22 teeth in the mandible and the maxilla, a snout comparable in relative length to the modern Gavialis gangeticus, and is notable since that its orbits were wider than long and not so upturned as another species of gavialids, including the gryposuchines, which implies that G. pachakamue doesn't had the "telescoped" orbits (protruding eyes) condition fully developed. Since that it species, that inhabited the proto-Amazon fluvial system 13 million years ago, is the oldest record of gavialids in this area and it had a primitive telescoped eyes condition, it shows that the development of such condition was a case of convergent evolution with the species of Gavialis also found in fluvial environments.[13]

Indeterminate finds of Gryposuchus were noted from the early Miocene Castillo Formation of Venezuela, middle Miocene Pebas Formation of Peru, middle/late Miocene Tranquitas Formation of Argentina and from the late Miocene formations Urumaco of Venezuela and Solimões in both Brazil and Peru.[2] Additionally, indeterminate finds of gavialoids (all in either coastal or marine sediments) are present in the early Miocene Jimol Formation and the early/middle Miocene Castilletes Formation in Colombia,[2][14] and from the Oligo-Miocene boundary Pirabas Formation of coastal Brazil.[15]

Phylogeny

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A phylogenetic analysis conducted in a 2007 study found Gryposuchinae to include the genera Aktiogavialis, Gryposuchus, Ikanogavialis, Piscogavialis, and Siquisiquesuchus. Below is a cladogram from the 2007 analysis showing the phylogenetic relationships of gryposuchines among gavialoids:[16]

Gavialoidea 

Alternatively, a 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data indicated that the members of Gryposuchinae and the genus Gryposuchus may in fact be paraphyletic and rather an evolutionary grade towards Gavialis and the gharial, as shown in the cladogram below:[17]

Paleoecology

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The Miocene epoch represents the only history of gavialoids (solely of the subfamily Gryposuchinae) in South America, from a Caribbean launchpad (Aktiogavialis from the Middle Oligocene of Puerto Rico,[16] and Dadagavialis from the Early Miocene of Panama).[18] Although there were six other confirmed genera of gryposuchine, Gryposuchus was almost certainly the most successful, with an existence potentially encompassing almost all of the Miocene, and a range from Venezuela to Argentina in the Middle to Late Miocene. This dominance was likely due to the fact that Gryposuchus was one of only two freshwater adapted gryposuchines (other than Hesperogavialis),[19] whereas the others (such as Siquisiquesuchus and Piscogavialis) were either primarily estuarine, coastal or marine based predators.[20][21] This would certainly have been useful in taking advantage of the extensive continental waterways and swamps of what would become the Amazon basin. Gryposuchus can be observed far and wide, from coastally adjacent and inclusive formations, such as the Urumaco Formation of Venezuela,[22][23] to even beyond the northern drainage basins, into Argentina.[5] This is in contrast with almost all the other species within the subfamily, which are limited to certain time periods near or on coast, with only Hesperogavialis penetrating into Brazil in the Late Miocene.

Although Gryposuchus had already reached Argentina by the Middle Miocene,[2] known species diversity reached its peak by the Late Miocene, with four of the five species present, three of which were also overlapping in the Urumaco Formation. Gryposuchinae diversity also reached its peak, at five genera across South America. However, at the Miocene/Pliocene boundary, all Gavialoidea and Crocodyloidea (another superfamily colonising in the Miocene) were likely extirpated from South America, with the endemic Caimaninae undergoing a severe reduction in size and diversity as well. This was likely due to the continuing elevation of the northern sections of the Andes chain creating the future Amazon basin, re-rerouting drainage flowing towards the Caribbean to the much cooler Atlantic, and transforming the mega-wetlands responsible for the hyper-diversity of crocodilians into a fully developed riverine drainage system. The co-current aridification of the continental interior, and filling of peripheral wetland basins, further restricted the space and food resources of these large, food-intensive specialist crocodilians, and was probably the primary cause of their extinction.[2][14][24]

References

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  1. ^ Rio, Jonathan P.; Mannion, Philip D. (6 September 2021). "Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem". PeerJ. 9: e12094. doi:10.7717/peerj.12094. PMC 8428266. PMID 34567843.
  2. ^ a b c d e f g h i j k l Cidade, Giovanne; Fortier, Daniel; Hsiou, Annie (2018-12-01). "The crocodylomorph fauna of the cenozoic of South America and its evolutionary history: A review". Journal of South American Earth Sciences. 90: 392–411. doi:10.1016/j.jsames.2018.12.026. S2CID 134902094.
  3. ^ a b c de Souza, R.G.; Riff, D.; de Souza-Filho, J.P.; Kellner, A.W.A. (2018). "Revisiting Gryposuchus jessei Gürich, 1912 (Crocodylia: Gavialoidea): specimen description and comments on the genus". Zootaxa. 4457 (1): 167–178. doi:10.11646/zootaxa.4457.1.9. PMID 30314186. S2CID 52976475.
  4. ^ a b Langston, W.; Gasparini, Z. (1997). "Crocodilians, Gryposuchus, and the South Americans gavials". In Kay, R.F.; Madden, R.H.; Cifelli, R.L.; Flynn, J.J. (eds.). Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta, Colombia. Washington: Smithsonian Institution. pp. 113–154.
  5. ^ a b Bona, Paula; Riff, Douglas; Gasparini, Zulma (2013-09-23). "Late Miocene crocodylians from northeast Argentina: New approaches about the austral components of the Neogene South American crocodylian fauna". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 29 (3–4): 551–570. doi:10.1017/S175569101300042X. hdl:11336/23291. S2CID 131646050.
  6. ^ Burmeister, G. (1885). "Examen crítico de los mamíferos y los reptiles denominados pot Don Augusto Bravard". Anales del Museo Púbtico de Buenos Aires. 3: 95–173.
  7. ^ Cozzuol, M. A. (2006). "The Acre vertebrate fauna: Age, diversity, and geography". Journal of South American Earth Sciences. 21 (3): 185–203. Bibcode:2006JSAES..21..185C. doi:10.1016/j.jsames.2006.03.005.
  8. ^ Langston, W. (1965). "Fossil crocodilians from Colombia and the Cenozoic history of the Crocodilia in South America". University of California Publications in Geological Sciences. 52: 1–152.
  9. ^ Salas-Gismondi, R.; Antoine, P. O.; Baby, P.; Brusset, S.; Benammi, M.; Espurt, N.; de Franceschi, D.; Pujos, F.; Tejada, J.; Urbina, M (2007). "Middle Miocene crocodiles from the Fitzcarrald Arch, Amazonian Peru". In Díaz-Martínez, E.; Rábano, I. (eds.). 4th European Meeting on the Palaeontology and Stratigraphy of Latin America. Cuadernos del Museo Geominero. Vol. 8. Madrid: Instituto Geológico y Minero de España.
  10. ^ Buffetaut, E. (1982). "Systematique, origine et évolution des Gavialidae Sud Américains". Geobios. 16 (1S): 127–140. Bibcode:1982Geobi..15..127B. doi:10.1016/S0016-6995(82)80107-1.
  11. ^ Riff, D.; Aguilera, O. A. (2008). "The world's largest gharials Gryposuchus: description of G. croizati n. sp. (Crocodylia, Gavialidae) from the Upper Miocene Urumaco Formation, Venezuela". Paläontologische Zeitschrift. 82 (2): 178–195. Bibcode:2008PalZ...82..178R. doi:10.1007/bf02988408. S2CID 85172486.
  12. ^ Head, J. J. (2001). "Systematics and body size of the gigantic, enigmatic crocodyloid Rhamphosuchus crassidens, and the faunal history of Siwalik Group (Miocene) crocodylians". Journal of Vertebrate Paleontology. 21 (Supplement to No. 3): 59A.
  13. ^ a b Salas-Gismondi, Rodolfo; Flynn, John J.; Baby, Patrice; Tejada-Lara, Julia V.; Claude, Julien; Antoine, Pierre-Olivier (2016). "A New 13 Million Year Old Gavialoid Crocodylian from Proto-Amazonian Mega-Wetlands Reveals Parallel Evolutionary Trends in Skull Shape Linked to Longirostry". PLOS ONE. 11 (4): e0152453. Bibcode:2016PLoSO..1152453S. doi:10.1371/journal.pone.0152453. PMC 4838223. PMID 27097031.
  14. ^ a b Moreno-Bernal, Jorge W.; Head, Jason; Jaramillo, Carlos A. (2016-05-03). "Fossil Crocodilians from the High Guajira Peninsula of Colombia: Neogene faunal change in northernmost South America". Journal of Vertebrate Paleontology. 36 (3): e1110586. Bibcode:2016JVPal..36E0586M. doi:10.1080/02724634.2016.1110586. ISSN 0272-4634. S2CID 130332367.
  15. ^ Moraes-Santos, Heloisa; Villanueva, Jean Bocquentin; Toledo, Peter Mann (2011-12-01). "New remains of a gavialoid crocodilian from the late Oligocene−early Miocene of the Pirabas Formation, Brazil". Zoological Journal of the Linnean Society. 163 (suppl_1): S132–S139. doi:10.1111/j.1096-3642.2011.00710.x. ISSN 0024-4082.
  16. ^ a b Vélez-Juarbe, Jorge; Brochu, C.A.; Santos, H. (2007). "A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile". Proceedings of the Royal Society B. 274 (1615): 1245–1254. doi:10.1098/rspb.2006.0455. PMC 2176176. PMID 17341454.
  17. ^ Michael S. Y. Lee; Adam M. Yates (27 June 2018). "Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil". Proceedings of the Royal Society B. 285 (1881). doi:10.1098/rspb.2018.1071. PMC 6030529. PMID 30051855.
  18. ^ Salas-Gismondi, Rodolfo; Moreno-Bernal, Jorge W.; Scheyer, Torsten M.; Sánchez-Villagra, Marcelo R.; Jaramillo, Carlos (2019-06-18). "New Miocene Caribbean gavialoids and patterns of longirostry in crocodylians". Journal of Systematic Palaeontology. 17 (12): 1049–1075. Bibcode:2019JSPal..17.1049S. doi:10.1080/14772019.2018.1495275. ISSN 1477-2019. S2CID 91495532.
  19. ^ Kay, R. F. and Madden, R. H. (1997). Paleogeography and paleoecology. In: Kay, R. F., Madden, R. H, Cifelli, R. L., and Flynn, J. J., eds., Vertebrate paleontology in the neotropics: the Miocene fauna of La Venta, Colombia. Smithsonian Institution Press; Washington, DC. pp. 520–550.
  20. ^ Kraus, R (1998). "The cranium of Piscogavialis jugaliperforatus n. gen., n. sp. (Gavialidae, Crocodylia) from the Miocene of Peru". Paläontologische Zeitschrift. 72 (3–4): 389–406. Bibcode:1998PalZ...72..389K. doi:10.1007/bf02988368. S2CID 84214781.
  21. ^ Brochu, C. A.; Rincon, A. D. (2004). "A gavialoid crocodylian from the Lower Miocene of Venezuela". Special Papers in Palaeontology. 71: 61–78.
  22. ^ Linares, O. J. (2004). "Bioestratigrafia de la fauna de mamiferos de las Formaciones Socorro, Urumaco y Codore (Mioceno Medio–Plioceno Temprano) de la region de Urumaco, Falcon, Venezuela". Paleobiologia Neotropical. 1: 1–26.
  23. ^ Sánchez-Villagra, M. R.; Aguilera, O. A. (2006). "Neogene vertebrates from Urumaco, Falcón State, Venezuela: diversity and significance". Journal of Systematic Palaeontology. 4 (3): 213–220. Bibcode:2006JSPal...4..213S. doi:10.1017/s1477201906001829. S2CID 84357359.
  24. ^ "Fourteen closely related crocodiles existed around 5 million years ago". ScienceDaily. Retrieved 2020-04-19.
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