Crocodyliformes

Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudosuchians to survive the K-Pg extinction event.

Crocodyliformes
Temporal range: Late Triassic–Recent,
Protosuchus, an early crocodyliform
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Pseudosuchia
Clade: Loricata
Superorder: Crocodylomorpha
Clade: Crocodyliformes
Hay, 1930
Clades

In 1988, Michael J. Benton and James M. Clark argued that all traditional names for well-known groups of animals should be restricted to their crown clades, that is, used only for natural groups comprising all living members of any given lineage. This posed a problem for the crocodilians, because the name Crocodylia, while used in various ways by various scientists, had always included not only living crocodilians but many of their extinct ancestors known only from the fossil record.[1]

Benton and Clark's solution to this issue was to restrict the name Crocodylia to the group containing modern alligators, crocodiles, and gharials, plus any extinct members of those specific families. The traditional group "Crocodylia" was replaced by the name Crocodyliformes, which included many of the extinct families that the new definition left out. Clark and Benton did not initially provide an exact definition for Crocodyliformes; but, in 2001, Paul Sereno and colleagues defined it as the clade including Protosuchus richardsoni and the Nile crocodile, plus all descendants of their common ancestor.[2]

Chris Brochu agreed with the assessment that Crocodylia as a name has never had stable contents, and that a series of clades larger than the crown group Crocodylia (including Crocodyliformes and the slightly more inclusive clade Crocodylomorpha) was a good solution.[3] However, in a 2008 paper, Jeremy Martin and Benton reversed the previous opinion (co-authored by Benton) that Crocodylia should be restricted to the crown group, suggesting that Crocodyliformes should be considered a synonym of a more inclusive Crocodylia, and thus replaced.[4] Brochu and colleagues rejected this proposal, arguing that the crown definition of Crocodylia is the standard meaning both within and beyond the crocodyliform systematics community.[5]

Ecology

While all extant crocodilians are carnivorous, the ecological roles of Mesozoic crocodyliforms were more diverse, and included omnivory and herbivory. Herbivorous forms were present from the early Jurassic to the late Cretaceous and are thought to have evolved at least three times.[6]

Fossil tracks

Tracks of a crocodyliform are known since Cretaceous, at least. Tracks representing the ichnofamily Batrachopodidae are described from the Early Cretaceous (late Aptian) Calonda Formation (Angola) by Mateus et al. (2017), who name a new ichnotaxon Angolaichnus adamanticus.[7]

Phylogeny

Below is a simplified cladogram based on Fiorelli and Calvo (2007).[8]

Crocodylomorpha 
 
 

Terrestrisuchus

 
 

Dibothrosuchus

 
 
 Crocodyliformes 
 
 Gobiosuchidae 
 

Gobiosuchus

 
 

Zaraasuchus

 
 
 
 

Orthosuchus

 
 
 Protosuchidae 
 

Protosuchus

 
 

Hemiprotosuchus

 
 
 
 

"Kayenta Form"

 
 

Edentosuchus

 
 
 
 
 
 

Mesoeucrocodylia

 
 
 

In 2012, paleontologists Mario Bronzati, Felipe Chinaglia Montefeltro, and Max C. Langer conducted a broad phylogenetic analysis to produce supertrees of Crocodyliformes, including 184 species. The most parsimonious trees were highly resolved, meaning the phylogenetic relationships found in the analysis were highly likely. Below is a consensus tree from the study:[9]

 
 

Kayentasuchus walkeri

 
Crocodyliformes
Protosuchia
 

Orthosuchus stormbergi

 
 
 
 

Protosuchus richardsoni

 
 

Hemiprotosuchus leali

 
 
 
 

Kayenta Form

 
 

Edentosuchus tienshanensis

 
 
 
 
 
 
 

Zaraasuchus shepardi

 
 

Las Hoyas Croc

 
 

Gobiosuchus kielanae

 
 
 
 

Eopneumatosuchus colberti

 
 
 

Zosuchus davidsoni

 
 
 
 

Sichuanosuchus shuhanensis

 
 

Sichuanosuchus huidongensis

 
 
 
 

Shantungosuchus hangjinensis

 
 

Shantungosuchus chuhsiensis

 
 

Shantungosuchus brachycephalus

 
 

Neuquensuchus universitas

 
 
 
 
 
 
 

Shartegosuchus asperopalatum

 
 

Fruita Form (Fruitachampsa callisoni)

 
 
 
 
 

Hsisosuchus dashanpuensis

 
 

Hsisosuchus chungkingensis

 
 
 
Notosuchia
 
 

Araripesuchus wegeneri

 
 
 

Araripesuchus tsangatsangana

 
 

Araripesuchus buitreraensis

 
 
 

Araripesuchus patagonicus

 
 

Araripesuchus gomesii

 
 
 
 
 
 
 

Uruguaysuchus terrai

 
 

Uruguaysuchus aznarezi

 
 
 
 

Libycosuchus brevirostris

 
 
 

Simosuchus clarki

 
 
 
 

Malawisuchus mwakasyungutiensis

 
 

Candidodon itapecuruense

 
 
 
 
 

Notosuchus terrestris

 
 
 

Comahuesuchus brachybuccalis

 
 
 

Mariliasuchus amarali

 
 
 

Yacarerani boliviensis

 
 

Pakasuchus kapilimai

 
 

Adamantinasuchus navae

 
 
 
 
 
 
 

Stratiotosuchus maxhechti

 
 

Pehuenchesuchus enderi

 
 

Pabwehshi pakistanensis

 
 

Iberosuchus macrodon

 
 

Eremosuchus elkoholicus

 
 

Doratodon

 
 

Bergisuchus dietrichbergi

 
 

Baurusuchus salgadoensis

 
 

Baurusuchus pachechoi

 
 
 

Chimaerasuchus paradoxus

 
 
 

Sphagesaurus huenei

 
 

Sphagesaurus montealtensis

 
 
 
 
 
 

Sebecus huilensis

 
 

Sebecus icaeorhinus

 
 
 
 

Itaborai Croc

 
 

Bretesuchus bonapartei

 
 
 
 
 
 
 
 
 
 
 
 

Anatosuchus minor

 
 
 
 

Barcinosuchus gradilis

 
 

Itasuchus jesuinoi

 
 
 

Miadanasuchus oblita

 
 

Trematochampsa taqueti

 
 

Caririsuchus camposi

 
 
 
 
 

Kaprosuchus saharicus

 
 

Mahajangasuchus insignis

 
 
Peirosauridae
 

Hamadasuchus rebouli

 
 
 

Montealtosuchus arrudacamposi

 
 

Uberabasuchus terrificus

 
 
 

Peirosaurus torminni

 
 

Lomasuchus palpebrosus

 
 
 
 
 
 
 

Neosuchia

 
 
 
 
 
 
 
 
 
 

References

  1. Benton, M.J. and Clark, J.M. (1988). "Archosaur phylogeny and the relationships of the Crocodylia." Pp. 295–338 in Benton, M.J. (ed.), The phylogeny and classification of the Tetrapods, volume 1. Oxford: Clarendon Press.
  2. Sereno, P.C.; Larson, H.C.E.; Sidor, C.A.; Gado, B. (2001). "The giant crocodyliform Sarcosuchus from the Cretaceous of Africa". Science. 294 (5546): 1516–1519. Bibcode:2001Sci...294.1516S. doi:10.1126/science.1066521. PMID 11679634. S2CID 22956704.
  3. Brochu, C.A. (2003). "Phylogenetic approaches toward crocodylian history". Annu. Rev. Earth Planet. Sci. 31 (31): 357–397. Bibcode:2003AREPS..31..357B. doi:10.1146/annurev.earth.31.100901.141308.
  4. 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.
  5. Brochu, A.C.; Wagner, J.R.; Jouve, S.; Sumrall, C.D.; Densmore, L.D. (2009). "A correction corrected: consensus over the meaning of Crocodylia and why it matters". Systematic Biology. 58 (5): 537–543. doi:10.1093/sysbio/syp053. PMID 20525607.
  6. Melstrom, K. M.; Irmis, R. B. (2019). "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. PMID 31257139.
  7. Octávio Mateus; Marco Marzola; Anne S. Schulp; Louis L. Jacobs; Michael J. Polcyn; Vladimir Pervov; António Olímpio Gonçalves; Maria Luisa Morais (2017). "Angolan ichnosite in a diamond mine shows the presence of a large terrestrial mammaliamorph, a crocodylomorph, and sauropod dinosaurs in the Early Cretaceous of Africa". Palaeogeography, Palaeoclimatology, Palaeoecology. 471: 220–232. Bibcode:2017PPP...471..220M. doi:10.1016/j.palaeo.2016.12.049.
  8. Fiorelli, LE; Calvo, JO (2007). "The first "protosuchian" (Archosauria: Crocodyliformes) from the Cretaceous (Santonian) of Gondwana". Arquivos do Museu Nacional, Rio de Janeiro. 65 (4): 417–459.
  9. 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.
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