أشباه التماسيح

(تم التحويل من Crocodylomorpha)

Crocodylomorphs
Temporal range: Late Triassic–Present, 235–0 Ma
Skeleton of Dromicosuchus, a small basal "sphenosuchian" crocodylomorph
التصنيف العلمي e
أصنوفة غير معروفة (أصلحها): Crocopoda
الفرع الحيوي: لوريكاتا
الفرع الحيوي: باثيوتيكا
Superorder: أشباه_التماسيح
Hay, 1930
Type species
Crocodylus niloticus
Laurenti, 1768
Subgroups
Crocodylomorpha incertae sedis
Possible crocodylomorphs

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. Extinct crocodylomorphs were considerably more ecologically diverse than modern crocodilians. The earliest and most primitive crocodylomorphs are represented by "sphenosuchians", a paraphyletic assemblage containing small-bodied, slender forms with elongated limbs that walked upright, which represents the ancestral morphology of Crocodylomorpha. These forms persisted until the end of the Jurassic.[1] 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.

التاريخ التطوري

Life restoration of Hesperosuchus from the Late Triassic (Carnian) of North America, an early branching, so-called "sphenosuchian" crocodylomorph
Life restoration of basal crocodylomorph Galahadosuchus.

The earliest lineages of Crocodylomorpha are placed into the paraphyletic "Sphenosuchia", which are characterized by slender bodies with elongate legs. The oldest known crocodylomorph is Trialestes, known from the Late Triassic (Carnian-Norian) of Argentina, around 231–225 million years ago,[2] the last groups of "sphenosuchians" persisted until the end of the Jurassic.[1] During the Jurassic, crocodylomorphs diversified, including the emergence of herbivorous and omnivorous forms,[3] 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[4] During the Cretaceous, the Notosuchia were a diverse group across the Southern Hemisphere occupying many diverse ecologies.[5] Modern crocodilians, a subgroup of Neosuchia, emerged during the Late Cretaceous.[4] Crocodylomorph diversity was severely reduced by the end-Cretaceous extinction event.[5] 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.[6][4]

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.[7] Crocodylomorpha has been given the rank of superorder in some 20th and 21st century studies.[8]

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).

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). Crocodylomorpha in the modern sense, as defined by Paul Sereno in 2005, is phylogenetically defined as the most inclusive clade containing Crocodylus niloticus (the Nile crocodile), but not Rauisuchus tiradentes, Poposaurus gracilis, Gracilisuchus stipanicicorum, Prestosuchus chiniquensis, or Aetosaurus ferratus.[9]

Phylogeny

Below is a cladogram of most known crocodylomorphs from Stephan F. Speikeman in 2023.[10] The modern consensus is that "sphenosuchians" form a paraphyletic assemblage leading towards the more derived Crocodyliformes. The basal crocodylomorph Saltoposuchidae was defined by Speikman, 2023 as the most inclusive clade containing Saltoposuchus connectens, but not Sphenosuchus acutus, Carnufex carolinensis, and Trialestes romeri. The clade Solidocrania was established by Alexander A. Ruebenstahl and colleagues in 2022 to unite crocodyliforms with their closest "sphenosuchian" relatives who both share similarly reinforced skulls. This clade was defined as the least inclusive clade including Junggarsuchus sloani, Almadasuchus figarii, and Macelognathus vagans.[11] The following cladogram was published by Juan Martín Leardi in 2025.[12]

Crocodylomorpha

Carnufex

CM 73372

Redondavenator

Trialestidae

Pseudohesperosuchus

Trialestes

Hesperosuchus

Dromicosuchus

Kayentasuchus

Saltoposuchus

Terrestrisuchus

Litargosuchus

Sphenosuchus

Hallopus
Dibothrosuchus
Solidocrania

Junggarsuchus

Macelognathus

Almadasuchus

Eopneumatosuchus

Crocodyliformes ملف:Deinosuchus riograndensis.png

Cladogram from Bodenham et al. 2026:[13]

Suchia

Stagonolepis

Gracilisuchus

Saurosuchus

Postosuchus

Crocodylomorpha

CM 73372

Carnufex

Redondavenator

Erpetosuchus

Trialestes

Pseudohesperosuchus

Saltoposuchidae

Saltoposuchus

Litargosuchus

Terrestrisuchus

Galahadosuchus

Hesperosuchus

Dromicosuchus

Kayentasuchus

Sphenosuchus

Dibothrosuchus

Junggarsuchus

Almadasuchus

Macelognathus

Hallopus

Crocodyliformes

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.

In some forms, like Hesperosuchus and Terrestrisuchus, metatarsal V still had one or two phalanges,[15] but in Crocodyliformes all metatarsal V phalanges have been lost.[10]

المراجع

  1. ^ أ ب Ruebenstahl, Alexander A.; Klein, Michael D.; Yi, Hongyu; Xu, Xing; Clark, James M. (14 يونيو 2022). "Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros". The Anatomical Record (in الإنجليزية). 305 (10): 2463–2556. doi:10.1002/ar.24949. ISSN 1932-8486. PMC 9541040. PMID 35699105.
  2. ^ Irmis, Randall B.; Nesbitt, Sterling J.; Sues, Hans-Dieter (يناير 2013). "Early Crocodylomorpha". Geological Society, London, Special Publications (in الإنجليزية). 379 (1): 275–302. Bibcode:2013GSLSP.379..275I. doi:10.1144/SP379.24. ISSN 0305-8719.
  3. ^ Melstrom, Keegan M.; Irmis, Randall B. (يوليو 2019). "Repeated Evolution of Herbivorous Crocodyliforms during the Age of Dinosaurs". Current Biology (in الإنجليزية). 29 (14): 2389–2395.e3. Bibcode:2019CBio...29E2389M. doi:10.1016/j.cub.2019.05.076. PMID 31257139.
  4. ^ أ ب ت Wilberg, Eric W.; Turner, Alan H.; Brochu, Christopher A. (24 يناير 2019). "Evolutionary structure and timing of major habitat shifts in Crocodylomorpha". Scientific Reports (in الإنجليزية). 9 (1): 514. Bibcode:2019NatSR...9..514W. doi:10.1038/s41598-018-36795-1. ISSN 2045-2322. PMC 6346023. PMID 30679529.
  5. ^ أ ب Stubbs, Thomas L.; Pierce, Stephanie E.; Elsler, Armin; Anderson, Philip S. L.; Rayfield, Emily J.; Benton, Michael J. (31 مارس 2021). "Ecological opportunity and the rise and fall of crocodylomorph evolutionary innovation". Proceedings of the Royal Society B: Biological Sciences (in الإنجليزية). 288 (1947). doi:10.1098/rspb.2021.0069. ISSN 0962-8452. PMC 8059953. PMID 33757349.
  6. ^ Martin, Jeremy E.; Pochat-Cottilloux, Yohan; Laurent, Yves; Perrier, Vincent; Robert, Emmanuel; Antoine, Pierre-Olivier (28 أكتوبر 2022). "Anatomy and phylogeny of an exceptionally large sebecid (Crocodylomorpha) from the middle Eocene of southern France". Journal of Vertebrate Paleontology (in الإنجليزية). 42 (4). Bibcode:2022JVPal..42E3828M. doi:10.1080/02724634.2023.2193828. ISSN 0272-4634.
  7. ^ 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.
  8. ^ 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.
  9. ^ 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.
  10. ^ أ ب Spiekman, Stephan 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 – via Oxford Academic.
  11. ^ 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.
  12. ^ Leardi, Juan Martín (2025). "Redescription of Pseudhesperosuchus jachaleri (Archosauria: Crocodylomorpha) from the los Colorados Formation (Norian), Argentina". Journal of Systematic Palaeontology. 23 (1). Bibcode:2025JSPal..2307779L. doi:10.1080/14772019.2025.2507779.
  13. ^ Bodenham, E. H.; Spiekman, S. N. F.; Maidment, S. C. R.; Upchurch, P.; Mannion, P. D. (2026). "A second species of non-crocodyliform crocodylomorph from the Late Triassic fissure deposits of southwestern UK: Implications for locomotory ecological diversity in Saltoposuchidae". The Anatomical Record. doi:10.1002/ar.70162.
  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.
  15. ^ Spiekman, Stephan N. F.; Fernandez, Vincent; Butler, Richard J.; Dollman, Kathleen N.; Maidment, Susannah C. R. (2023). "A taxonomic revision and cranial description of Terrestrisuchus gracilis (Archosauria, Crocodylomorpha) from the Upper Triassic of Pant-y-Ffynnon Quarry (Southern Wales)". Papers in Palaeontology. 9 (6). Bibcode:2023PPal....9E1534S. doi:10.1002/spp2.1534.

Sources

وصلات خارجية

قالب:Pseudosuchia