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'''Crocodylomorpha''' is a group of [[Pseudosuchia|pseudosuchian]] [[archosaur]]s that includes the [[crocodilia]]ns and their extinct relatives. They were the only members of Pseudosuchia to survive the [[Triassic–Jurassic extinction event|end-Triassic extinction]].
'''Crocodylomorpha''' is a group of [[Pseudosuchia|pseudosuchian]] [[archosaur]]s that includes the [[crocodilia]]ns and their extinct relatives. They were the only members of Pseudosuchia to survive the [[Triassic–Jurassic extinction event|end-Triassic extinction]].


During [[Mesozoic]] and early Cenozoic times, crocodylomorphs were far more diverse than at Present. Triassic forms were small, lightly built, active terrestrial animals. The earliest and most primitive crocodylomorphs are represented by "[[Sphenosuchia|sphenosuchians]]", a [[paraphyletic]] assemblage containing small-bodied forms with elongated limbs that walked upright, which represents the ancestral morphology of Crocodylomorpha. These forms persisted until the end of the Jurassic.<ref>{{Cite journal |last1=Ruebenstahl |first1=Alexander A. |last2=Klein |first2=Michael D. |last3=Yi |first3=Hongyu |last4=Xu |first4=Xing |last5=Clark |first5=James M. |date=2022-06-14 |title=Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros |journal=The Anatomical Record |volume=305 |issue=10 |language=en |pages=2463–2556 |doi=10.1002/ar.24949 |pmid=35699105 |pmc=9541040 |issn=1932-8486}}</ref> During the Jurassic, Crocodylomorphs morphologically diversified into numerous niches, with the subgroups [[Neosuchia]] (which includes modern crocodilians) and the extinct [[Thalattosuchia]] adapting to aquatic life.
During [[Mesozoic]] and early Cenozoic times, crocodylomorphs were far more diverse than at Present. Triassic forms were small, lightly built, active terrestrial animals. The earliest and most primitive crocodylomorphs are represented by "[[Sphenosuchia|sphenosuchians]]", a [[paraphyletic]] assemblage containing small-bodied forms with elongated limbs that walked upright, which represents the ancestral morphology of Crocodylomorpha. These forms persisted until the end of the Jurassic.<ref>{{Cite journal |last1=Ruebenstahl |first1=Alexander A. |last2=Klein |first2=Michael D. |last3=Yi |first3=Hongyu |last4=Xu |first4=Xing |last5=Clark |first5=James M. |date=2022-06-14 |title=Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros |journal=The Anatomical Record |volume=305 |issue=10 |language=en |pages=2463–2556 |doi=10.1002/ar.24949 |pmid=35699105 |pmc=9541040 |issn=1932-8486}}</ref> During the Jurassic, crocodylomorphs morphologically diversified into numerous niches, with the subgroups [[Neosuchia]] (which includes modern crocodilians) and the extinct [[Thalattosuchia]] adapting to aquatic life, while some terrestrial groups adopted herbivorous and omnivorous lifestyles. Terrestrial crocodylomorphs would continue to co-exist alongside aquatic forms until becoming extinct during the [[Miocene]].


== Evolutionary history ==
== Evolutionary history ==
[[File:Hesperosuchus BW.jpg|left|thumb|Life restoration of ''[[Hesperosuchus]]'']]
[[File:Hesperosuchus BW.jpg|left|thumb|Life restoration of ''[[Hesperosuchus]]'']]
The earliest lineages of Crocodylomorpha are placed into the paraphyletic "[[Sphenosuchia]]", which are charactersed by slender bodies with elongate legs. The oldest known crocodylmorph is ''[[Trialestes]]'', known from the Late Triassic (Carnian-Norian) of Argentina, around 231–225 million years ago.<ref>{{Cite journal |last=Irmis |first=Randall B. |last2=Nesbitt |first2=Sterling J. |last3=Sues |first3=Hans-Dieter |date=2013-01 |title=Early Crocodylomorpha |url=https://www.lyellcollection.org/doi/10.1144/SP379.24 |journal=Geological Society, London, Special Publications |language=en |volume=379 |issue=1 |pages=275–302 |doi=10.1144/SP379.24 |issn=0305-8719}}</ref> During the Jurassic, crocodylomorphs diversified, including the emergence of herbivorous and omnivorous forms,<ref>{{Cite journal |last=Melstrom |first=Keegan M. |last2=Irmis |first2=Randall B. |date=2019-07 |title=Repeated Evolution of Herbivorous Crocodyliforms during the Age of Dinosaurs |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982219306906 |journal=Current Biology |language=en |volume=29 |issue=14 |pages=2389–2395.e3 |doi=10.1016/j.cub.2019.05.076}}</ref> as well as the aquatically adapted [[Neosuchia]] and [[Thalattosuchia]], with Thalattosuchia and several groups of neosuchians becoming adapted to a marine lifestyle over the Jurassic and Cretaceous<ref name=":0">{{Cite journal |last=Wilberg |first=Eric W. |last2=Turner |first2=Alan H. |last3=Brochu |first3=Christopher A. |date=2019-01-24 |title=Evolutionary structure and timing of major habitat shifts in Crocodylomorpha |url=https://www.nature.com/articles/s41598-018-36795-1 |journal=Scientific Reports |language=en |volume=9 |issue=1 |doi=10.1038/s41598-018-36795-1 |issn=2045-2322 |pmc=PMC6346023 |pmid=30679529}}</ref> During the Cretaceous, the [[Notosuchia]] were a diverse group across the Southern Hemisphere occupying many diverse ecologies.<ref name=":1">{{Cite journal |last=Stubbs |first=Thomas L. |last2=Pierce |first2=Stephanie E. |last3=Elsler |first3=Armin |last4=Anderson |first4=Philip S. L. |last5=Rayfield |first5=Emily J. |last6=Benton |first6=Michael J. |date=2021-03-31 |title=Ecological opportunity and the rise and fall of crocodylomorph evolutionary innovation |url=https://royalsocietypublishing.org/doi/10.1098/rspb.2021.0069 |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=288 |issue=1947 |doi=10.1098/rspb.2021.0069 |issn=0962-8452 |pmc=PMC8059953 |pmid=33757349}}</ref> Modern [[Crocodilia|crocodilians]], a subgroup of Neosuchia, emerged during the Late Cretaceous.<ref name=":0" /> Crocodylmorph diversity was severely reduced by the [[end-Cretaceous extinction event]].<ref name=":1" /> The last group of terrestrially adapted crocodylomorphs was the [[Sebecidae]], a group of large predatory notosuchians which persisted in South America until the middle [[Miocene]] around 12 million years ago.<ref>{{Cite journal |last=Martin |first=Jeremy E. |last2=Pochat-Cottilloux |first2=Yohan |last3=Laurent |first3=Yves |last4=Perrier |first4=Vincent |last5=Robert |first5=Emmanuel |last6=Antoine |first6=Pierre-Olivier |date=2022-10-28 |title=Anatomy and phylogeny of an exceptionally large sebecid (Crocodylomorpha) from the middle Eocene of southern France |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2023.2193828 |journal=Journal of Vertebrate Paleontology |language=en |volume=42 |issue=4 |doi=10.1080/02724634.2023.2193828 |issn=0272-4634}}</ref><ref name=":0" />
When their extinct species and [[stem group]] are examined, the crocodylian lineage (clade [[Pseudosuchia]], formerly [[Crurotarsi]]) proves to have been a very diverse and adaptive group of reptiles. Not only are they an ancient group of animals – at least as old as the [[dinosaur]]s – they also evolved into a great variety of forms and sizes. Some terrestrial [[notosuchia]]ns were noticeably small animals, less than {{cvt|1.2|m}} in length, while aquatic or semiaquatic crocodylomorphs, especially [[tethysuchia]]ns, [[thalattosuchia]]ns, and some [[crocodilia]]ns reached around {{cvt|8|m}} in length.<ref>{{cite journal|last1=Godoy |first1=P. L. |last2=Benson |first2=R. B. J. |last3=Bronzati |first3=M. |last4=Butler |first4=R. J. |year=2019 |title=The multi-peak adaptive landscape of crocodylomorph body size evolution |journal=[[BMC Evolutionary Biology]] |volume=19 |issue=167 |page=167 |doi=10.1186/s12862-019-1466-4 |pmid=31390981 |pmc=6686447 |url=https://bmcecolevol.biomedcentral.com/counter/pdf/10.1186/s12862-019-1466-4.pdf |doi-access=free }}</ref>

The earliest forms, the [[sphenosuchia]]ns, evolved during the Late [[Triassic]], and were highly [[Gracility|gracile]] terrestrial forms built like greyhounds. During the [[Jurassic]] and the Cretaceous, marine forms in the family [[Metriorhynchidae]], such as ''[[Metriorhynchus]]'', evolved forelimbs that were paddle-like and had a tail similar to modern fish. ''[[Dakosaurus andiniensis]]'', a species closely related to ''Metriorhynchus'', had a skull that was adapted to eat large marine reptiles. Several terrestrial species during the [[Cretaceous]] were herbivorous, such as ''[[Simosuchus clarki]]'' and ''[[Chimaerasuchus paradoxus]]''. A number of lineages during the Cenozoic became wholly terrestrial predators.


==Taxonomy and phylogeny==
==Taxonomy and phylogeny==

Revision as of 23:11, 10 April 2024

Crocodylomorphs
Temporal range: Late Triassic–Present, 235–0 Ma[1]
Skeleton of Terrestrisuchus, an early saltoposuchid crocodylomorph
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Paracrocodylomorpha
Clade: Loricata
Clade: Crocodylomorpha
Hay, 1930
Subgroups

Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction.

During Mesozoic and early Cenozoic times, crocodylomorphs were far more diverse than at Present. Triassic forms were small, lightly built, active terrestrial animals. The earliest and most primitive crocodylomorphs are represented by "sphenosuchians", a paraphyletic assemblage containing small-bodied forms with elongated limbs that walked upright, which represents the ancestral morphology of Crocodylomorpha. These forms persisted until the end of the Jurassic.[4] During the Jurassic, crocodylomorphs morphologically diversified into numerous niches, with the subgroups Neosuchia (which includes modern crocodilians) and the extinct Thalattosuchia adapting to aquatic life, while some terrestrial groups adopted herbivorous and omnivorous lifestyles. Terrestrial crocodylomorphs would continue to co-exist alongside aquatic forms until becoming extinct during the Miocene.

Evolutionary history

Life restoration of Hesperosuchus

The earliest lineages of Crocodylomorpha are placed into the paraphyletic "Sphenosuchia", which are charactersed by slender bodies with elongate legs. The oldest known crocodylmorph is Trialestes, known from the Late Triassic (Carnian-Norian) of Argentina, around 231–225 million years ago.[5] During the Jurassic, crocodylomorphs diversified, including the emergence of herbivorous and omnivorous forms,[6] as well as the aquatically adapted Neosuchia and Thalattosuchia, with Thalattosuchia and several groups of neosuchians becoming adapted to a marine lifestyle over the Jurassic and Cretaceous[7] During the Cretaceous, the Notosuchia were a diverse group across the Southern Hemisphere occupying many diverse ecologies.[8] Modern crocodilians, a subgroup of Neosuchia, emerged during the Late Cretaceous.[7] Crocodylmorph diversity was severely reduced by the end-Cretaceous extinction event.[8] The last group of terrestrially adapted crocodylomorphs was the Sebecidae, a group of large predatory notosuchians which persisted in South America until the middle Miocene around 12 million years ago.[9][7]

Taxonomy and phylogeny

Historically, all known living and extinct crocodiles were indiscriminately lumped into the order Crocodilia. However, beginning in the late 1980s, many scientists began restricting the order Crocodilia to the living species and close extinct relatives such as Mekosuchus. The various other groups that had previously been known as Crocodilia were moved to Crocodylomorpha and the slightly more restrictive Crocodyliformes.[10] Crocodylomorpha has been given the rank of superorder in some 20th and 21st century studies.[11]

The old Crocodilia was subdivided into the suborders:

Mesosuchia is a paraphyletic group as it does not include eusuchians (which nest within Mesosuchia). Mesoeucrocodylia was the name given to the clade that contains mesosuchians and eusuchians (Whetstone and Whybrow, 1983).

Phylogeny

Below is a cladogram modified from Nesbitt (2011)[12] and Bronzati (2012).[13]

Crocodylomorpha 

The previous definitions of Crocodilia and Eusuchia did not accurately convey evolutionary relationships within the group. The only order-level taxon that is currently considered valid is Crocodilia in its present definition. Prehistoric crocodiles are represented by many taxa, but since few major groups of the ancient forms are distinguishable, a conclusion on how to define new order-level clades is not yet possible. (Benson & Clark, 1988).

Biology

The Crocodylomorpha comprise a variety of forms, shapes, and sizes, which occupied a range of habitats. As with most amniotes, Crocodylomorphs were and are oviparous, laying eggs in a nest or mound, known from strata as old as the Late Jurassic.[14] Adult size varies widely, from about 55 cm long in Knoetschkesuchus to much larger dimensions, as in Sarcosuchus. Most crocodylomorphs were carnivores, but many lineages evolved to be obligate piscivores, such as the extant gharials.

References

  1. ^ Irmis, R. B.; Nesbitt, S. J.; Sues, H. -D. (2013). "Early Crocodylomorpha". Geological Society, London, Special Publications. 379 (1): 275–302. Bibcode:2013GSLSP.379..275I. doi:10.1144/SP379.24. S2CID 219190410.
  2. ^ Spiekman, S. N. F. (2023). "A revision and histological investigation of Saltoposuchus connectens (Archosauria: Crocodylomorpha) from the Norian (Late Triassic) of south-western Germany". Zoological Journal of the Linnean Society. 199 (2): 354–391. doi:10.1093/zoolinnean/zlad035.
  3. ^ Ruebenstahl, A. A.; Klein, M. D.; Yi, H.; Xu, X.; Clark, J. M. (2022). "Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros". The Anatomical Record. 305 (10): 2463–2556. doi:10.1002/ar.24949. PMC 9541040. PMID 35699105.
  4. ^ Ruebenstahl, Alexander A.; Klein, Michael D.; Yi, Hongyu; Xu, Xing; Clark, James M. (14 June 2022). "Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros". The Anatomical Record. 305 (10): 2463–2556. doi:10.1002/ar.24949. ISSN 1932-8486. PMC 9541040. PMID 35699105.
  5. ^ Irmis, Randall B.; Nesbitt, Sterling J.; Sues, Hans-Dieter (2013-01). "Early Crocodylomorpha". Geological Society, London, Special Publications. 379 (1): 275–302. doi:10.1144/SP379.24. ISSN 0305-8719. {{cite journal}}: Check date values in: |date= (help)
  6. ^ Melstrom, Keegan M.; Irmis, Randall B. (2019-07). "Repeated Evolution of Herbivorous Crocodyliforms during the Age of Dinosaurs". Current Biology. 29 (14): 2389–2395.e3. doi:10.1016/j.cub.2019.05.076. {{cite journal}}: Check date values in: |date= (help)
  7. ^ a b c Wilberg, Eric W.; Turner, Alan H.; Brochu, Christopher A. (24 January 2019). "Evolutionary structure and timing of major habitat shifts in Crocodylomorpha". Scientific Reports. 9 (1). doi:10.1038/s41598-018-36795-1. ISSN 2045-2322. PMC 6346023. PMID 30679529.{{cite journal}}: CS1 maint: PMC format (link)
  8. ^ a b Stubbs, Thomas L.; Pierce, Stephanie E.; Elsler, Armin; Anderson, Philip S. L.; Rayfield, Emily J.; Benton, Michael J. (31 March 2021). "Ecological opportunity and the rise and fall of crocodylomorph evolutionary innovation". Proceedings of the Royal Society B: Biological Sciences. 288 (1947). doi:10.1098/rspb.2021.0069. ISSN 0962-8452. PMC 8059953. PMID 33757349.{{cite journal}}: CS1 maint: PMC format (link)
  9. ^ Martin, Jeremy E.; Pochat-Cottilloux, Yohan; Laurent, Yves; Perrier, Vincent; Robert, Emmanuel; Antoine, Pierre-Olivier (28 October 2022). "Anatomy and phylogeny of an exceptionally large sebecid (Crocodylomorpha) from the middle Eocene of southern France". Journal of Vertebrate Paleontology. 42 (4). doi:10.1080/02724634.2023.2193828. ISSN 0272-4634.
  10. ^ Martin, J.E.; Benton, M.J. (2008). "Crown Clades in Vertebrate Nomenclature: Correcting the Definition of Crocodylia". Systematic Biology. 57 (1): 173–181. doi:10.1080/10635150801910469. PMID 18300130.
  11. ^ Parrilla-Bel, J.; Young, M. T.; Moreno-Azanza, M.; Canudo, J. I. (2013). Butler, Richard J (ed.). "The First Metriorhynchid Crocodylomorph from the Middle Jurassic of Spain, with Implications for Evolution of the Subclade Rhacheosaurini". PLOS ONE. 8 (1): e54275. Bibcode:2013PLoSO...854275P. doi:10.1371/journal.pone.0054275. PMC 3553084. PMID 23372699.
  12. ^ Nesbitt, S.J. (2011). "The early evolution of archosaurs: relationships and the origin of major clades". Bulletin of the American Museum of Natural History. 352: 1–292. doi:10.1206/352.1. hdl:2246/6112. S2CID 83493714.
  13. ^ Bronzati, M.; Montefeltro, F. C.; Langer, M. C. (2012). "A species-level supertree of Crocodyliformes". Historical Biology. 24 (6): 598–606. doi:10.1080/08912963.2012.662680. S2CID 53412111.
  14. ^ Russo, J.; Mateus, O.; Marzola, M.; Balbino, A. (2017). "Two new ootaxa from the late Jurassic: The oldest record of crocodylomorph eggs, from the Lourinhã Formation, Portugal". PLOS One. 12 (3): 1–23. Bibcode:2017PLoSO..1271919R. doi:10.1371/journal.pone.0171919. PMC 5342183. PMID 28273086.

Sources