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Crocodylomorpha
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| Crocodylomorphs Temporal range: Late Triassic–Present,
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| Skeleton of Terrestrisuchus, an early crocodylomorph | |||||
| Scientific classification | |||||
| Kingdom: | Animalia | ||||
| Phylum: | Chordata | ||||
| Class: | Reptilia | ||||
| Clade: | Archosauria | ||||
| Clade: | Pseudosuchia | ||||
| Clade: | Paracrocodylomorpha | ||||
| Clade: | Loricata | ||||
| Clade: | Crocodylomorpha Hay, 1930 | ||||
| Type species | |||||
| Crocodylus niloticus Laurenti, 1768
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| Subgroups | |||||
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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.
Evolutionary history
[edit]
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
[edit]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:
- Eusuchia: true crocodiles (which includes crown-group Crocodylia)
- Mesosuchia: 'middle' crocodiles
- Thalattosuchia: sea crocodiles
- Protosuchia: first crocodiles
- Sphenosuchia: wedge crocodiles
- Sebecosuchia: Sebecus crocodiles
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
[edit]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]
Biology
[edit]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.[13] 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,[14] but in Crocodyliformes all metatarsal V phalanges have been lost.[10]
References
[edit]- ^ a b 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.
- ^ Irmis, Randall B.; Nesbitt, Sterling J.; Sues, Hans-Dieter (January 2013). "Early Crocodylomorpha". Geological Society, London, Special Publications. 379 (1): 275–302. Bibcode:2013GSLSP.379..275I. doi:10.1144/SP379.24. ISSN 0305-8719.
- ^ Melstrom, Keegan M.; Irmis, Randall B. (July 2019). "Repeated Evolution of Herbivorous Crocodyliforms during the Age of Dinosaurs". Current Biology. 29 (14): 2389–2395.e3. Bibcode:2019CBio...29E2389M. doi:10.1016/j.cub.2019.05.076. PMID 31257139.
- ^ 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): 514. Bibcode:2019NatSR...9..514W. doi:10.1038/s41598-018-36795-1. ISSN 2045-2322. PMC 6346023. PMID 30679529.
- ^ 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.
- ^ 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). Bibcode:2022JVPal..42E3828M. doi:10.1080/02724634.2023.2193828. ISSN 0272-4634.
- ^ 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.
- ^ 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.
- ^ 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.
- ^ a b 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.
- ^ 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.
- ^ 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.
- ^ 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.
- ^ 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
[edit]- Benton, M. J. (2004), Vertebrate Palaeontology, 3rd ed. Blackwell Science Ltd
- Hay, O. P. 1930 (1929–1930). Second Bibliography and Catalogue of the Fossil Vertebrata of North America. Carnegie Institution Publications, Washington, 1, 990 pp.
External links
[edit]- Crocodylomorpha - webpages by Ross Elgin on the University of Bristol server
- Major subgroups classification (used here)
- Crocodylomorpha from Palaeos
Crocodylomorpha
View on GrokipediaEvolutionary history
Origins in the Triassic
Crocodylomorpha emerged during the Late Triassic, with the earliest known fossils dating to the Carnian stage approximately 235–227 million years ago.[1] These initial records include basal sphenosuchians such as Carnufex carolinensis from the Pekin Formation in North Carolina, representing one of the oldest documented members of the clade.[7] Similarly, Saltoposuchus connectens from the Tübingen Formation in Germany exemplifies early crocodylomorph morphology, though its fossils are slightly younger within the Norian stage.[8] These small-bodied forms, typically under 2 meters in length, indicate a rapid initial appearance following the diversification of pseudosuchian archosaurs. Key adaptations distinguishing crocodylomorphs from their archosaur ancestors included a fully erect limb posture, which enhanced terrestrial mobility and stamina compared to the more sprawling gait of earlier pseudosuchians.[9] Elongated hindlimbs further supported agile, cursorial locomotion, allowing these early crocodylomorphs to inhabit diverse terrestrial environments.[1] Dentition showed early signs of anisodonty, with differentiated teeth suited for grasping prey, marking a shift toward specialized carnivory inherited and refined from archosaur forebears.[10] By the Norian and Rhaetian stages (approximately 227–201 million years ago), crocodylomorphs underwent an initial radiation, evidenced by fossils from the Chinle Formation in North America, where taxa like Hesperosuchus agilis and cf. Sphenosuchus sp. document increased diversity in fluvial and floodplain settings.[1] In South America, the Los Colorados Formation yields remains of non-crocodyliforms such as Pseudhesperosuchus jachaleri and protosuchids, highlighting a broad Pangaean distribution during this interval.[11] This expansion coincided with arid to semi-arid habitats across the supercontinent Pangaea, where seasonal river systems and ephemeral lakes supported small, agile predators amid the early stages of continental rifting.[12] These conditions, characterized by hot, dry climates with intermittent wet phases, facilitated the ecological niche occupancy by basal crocodylomorphs before their further diversification in the Mesozoic.[13]Mesozoic diversification and key radiations
The Triassic-Jurassic mass extinction event approximately 201 million years ago triggered a rapid radiation of crocodylomorphs, the sole surviving pseudosuchian lineage, leading to significant morphological disparity by the Early Jurassic.[14] This diversification was marked by the emergence of thalattosuchians, a clade that adapted to fully marine environments during the Jurassic. Teleosaurids, such as Teleosaurus, inhabited coastal and nearshore habitats with robust, armored bodies suited for ambush predation in shallow waters, while metriorhynchids like Metriorhynchus and Geosaurus evolved streamlined forms, reduced armor, and paddle-like limbs for open-ocean pelagic lifestyles, representing one of the most extreme aquatic specializations among archosaurs.[15] Early neosuchians, basal relatives of modern crocodylians, also arose, exemplified by semi-aquatic forms such as Goniopholis from the Late Jurassic Morrison Formation in North America, which featured generalized skulls for versatile feeding in riverine and floodplain ecosystems.[16] In the Cretaceous, crocodylomorph diversity peaked with expansions into terrestrial and semi-aquatic niches, particularly in Gondwana, where notosuchians exhibited remarkable morphological innovation. Protosuchians, early crocodylomorphs with terrestrial traits, persisted in some regions, but notosuchians dominated with adaptations for upright posture, varied locomotion, and specialized dentitions reflecting omnivorous, herbivorous, or insectivorous diets. For instance, Anatosuchus minor from the Early Cretaceous of Niger possessed a broad, overhanging snout with low-pressure sensory pits, enabling it to forage for small fish, amphibians, and insects in shallow waters or mudflats.[17] Similarly, Pakasuchus kapilimai from the mid-Cretaceous of Tanzania displayed mammal-like heterodont teeth, including shearing carnassials and grinding molars for processing diverse terrestrial prey such as insects and small vertebrates, highlighting convergent evolution with early mammals.[18] Pholidosaurs, long-snouted neosuchians like Pholidosaurus, occupied semi-aquatic habitats in Laurasia with elongated rostra for piscivory, bridging basal and advanced crocodyliform forms.[19] Key radiations underscored crocodylomorphs' ecological versatility, including the rise of giant apex predators in Late Cretaceous coastal environments. Deinosuchus, an enormous alligatoroid reaching over 10 meters in length, preyed on large dinosaurs and marine reptiles along the Western Interior Seaway in North America, evidenced by bite marks on hadrosaur and tyrannosaurid bones, and tolerated brackish to saltwater conditions.[20] Fossil assemblages from iconic sites illustrate this diversification: the Morrison Formation yielded neosuchians like Goniopholis amid fluvial deposits; the Solnhofen Limestone preserved exceptionally detailed thalattosuchians such as Cricosaurus in lagoonal settings; and the Bahariya Formation in Egypt contained diverse crocodyliforms, including the aberrant Stomatosuchus with its filter-feeding pouch-like snout, in deltaic and coastal facies.[16][21][22] These adaptations allowed crocodylomorphs to exploit a broad array of niches, from fully aquatic to predominantly terrestrial, before the end-Cretaceous extinction.[23]Cenozoic survival and modern forms
The Cretaceous–Paleogene (K–Pg) extinction event at approximately 66 million years ago resulted in the extinction of most non-crocodylian crocodylomorph lineages, though recent evidence indicates brief survival of at least some notosuchians into the early Paleocene.[24] Only members of the crown-group Crocodylia, particularly within Eusuchia, survived long-term into the Cenozoic.[3] This survival is attributed to the ecological versatility of eusuchians, which allowed them to persist through the mass extinction that eliminated most other pseudosuchians; a 2025 study further highlights the clade's resilience across multiple extinctions due to ecomorphological adaptability.[25][26] Post-K–Pg, eusuchian lineages underwent diversification, marking a recovery phase amid the recovery of continental ecosystems.[3] In the Paleogene, early alligatoroids and crocodyloids emerged and diversified primarily in North America and Europe. For instance, the alligatoroid Alligator prenasalis represents one of the earliest definitive records of the genus Alligator in North America, known from Late Eocene deposits in Nebraska, indicating an expansion of alligatoroid forms during this period.[27] In Europe, Paleogene assemblages include basal alligatoroids such as Diplocynodon and Hassiacosuchus, alongside stem crocodyloids and planocraniids, reflecting a radiation of these groups in temperate to subtropical environments following the extinction.[28] These fossils document the initial Cenozoic establishment of modern crocodylian body plans and ecological roles.[28] During the Neogene and into the Quaternary, crocodylomorphs—now predominantly crocodylians—migrated toward tropical regions as global climates cooled, adapting to warmer, wetland-dominated ecosystems in the tropics.[29] Gigantism became prominent among some lineages, exemplified by Purussaurus, a Miocene caimanine from South American deposits that reached lengths of approximately 12 meters, serving as a top predator in fluvial systems.[30] This period saw crocodylians integrate into contemporary ecosystems, with their diversity stabilizing around 25 extant species today, shaped by biogeographic shifts and climatic constraints.[29] Recent molecular clock analyses estimate the divergence of the crown-group Crocodylia between 100 and 80 million years ago, aligning with Late Cretaceous origins and supporting the survival of these lineages through the K–Pg boundary.[31] These estimates, derived from integrated morphological and genetic data, highlight the deep evolutionary history leading to modern forms.[31]Taxonomy and systematics
Classification and major groups
Crocodylomorpha is defined as the most inclusive clade containing Crocodylus niloticus but excluding Poposaurus gracilis, encompassing the crown-group Crocodylia (extant crocodilians) and various stem-groups such as basal forms and intermediate taxa.[1] This clade originated in the Late Triassic and includes approximately 26 extant species alongside a rich diversity of extinct relatives that exhibited terrestrial, semiaquatic, and fully aquatic lifestyles.[4][32] The classification of crocodylomorphs traces back to Carl Linnaeus, who in the 10th edition of Systema Naturae (1758) placed the Nile crocodile within the genus Lacerta as Lacerta crocodilus, grouping it among lizards based on superficial morphological similarities rather than phylogenetic relationships.[33] Over the 19th and early 20th centuries, Linnaean taxonomy evolved through hierarchical groupings like orders and families, often emphasizing anatomical features such as skull structure, but these systems lacked explicit consideration of evolutionary ancestry. The shift to cladistic approaches in the mid-20th century, influenced by Willi Hennig's principles, emphasized monophyletic groups defined by shared derived characters; A. D. Walker's 1968 erection of Crocodylomorpha as a clade uniting Crocodylia with sphenosuchians marked a key transition, later refined in cladistic analyses by Jacques Gauthier in 1986.[34][35] Major subgroups of Crocodylomorpha include basal sphenosuchians, which represent early terrestrial forms from the Late Triassic to Middle Jurassic, and crocodyliforms such as Protosuchia, early Mesozoic basal crocodyliforms with primitive features like upright limb posture.[4] Mesoeucrocodylia, defined as all crocodyliforms more closely related to Crocodylus niloticus than to Protosuchus richardsoni, encompasses more derived forms, uniting Notosuchia (primarily terrestrial, diverse in the Cretaceous with adaptations like herbivory) sister to Neosuchia (semiaquatic, leading to the extant branch Eusuchia).[4] Other notable subgroups are Thalattosuchia, a Jurassic marine radiation with paddle-like limbs, and Neosuchia, which includes Eusuchia as the crown group containing all living crocodylians (Alligatoridae, Crocodylidae, and Gavialidae).[4] Among extinct families, Goniopholididae, basal neosuchians characterized by robust skulls and semiaquatic habits, ranged from the Middle Jurassic to the Early Cretaceous across Laurasia.[36] Dyrosauridae, a mesoeucrocodylian group with elongated snouts and marine affinities, persisted from the Late Cretaceous (Campanian) through the Middle Eocene, surviving the Cretaceous-Paleogene extinction and achieving a global distribution in the Paleogene.[37]Phylogenetic relationships
The phylogenetic relationships within Crocodylomorpha have been elucidated through numerous cladistic analyses, revealing a basal dichotomy where non-crocodyliform crocodylomorphs, often referred to as sphenosuchians, represent the earliest-diverging lineages and serve as the sister group to all more derived crocodylomorphs. These basal forms, such as Hesperosuchus and Litargosuchus, exhibit primitive archosaurian features including slender builds and terrestrial adaptations, with no evidence of monophyly for Sphenosuchia itself in recent parsimony-based trees.[5] Following this split, Protosuchia emerges as an early offshoot within Crocodyliformes, comprising basal crocodyliforms like Protosuchus that retain small body sizes and generalized morphologies, positioned as successive outgroups to more advanced clades in comprehensive morphological datasets.[5] Key debates persist regarding the placement of certain groups, notably Thalattosuchia, whose monophyly is generally supported by shared cranial and postcranial traits such as elongated snouts and paddle-like limbs adapted for marine life, but whose exact position remains contentious. Early analyses recovered Thalattosuchia as the sister group to Crocodyliformes, while others nest it within Neosuchia; recent expanded datasets from the 2020s, incorporating broader taxon sampling, support positions either as basal crocodylomorphs, within Neosuchia, or sister to Mesoeucrocodylia, depending on dataset and outgroup choice, highlighting the influence of outgroup choice on tree topology.[38] Similarly, Notosuchia exhibits debated affinities, with some lineages showing convergent evolutionary trends toward terrestrial mammal-like adaptations, including heterodont dentition, ziphodont teeth for shearing, and upright posture, suggesting niche overlap with mammalian herbivores and carnivores in Gondwanan ecosystems during the Cretaceous.[23] Central to crocodylomorph interrelationships is the node Mesoeucrocodylia, defined as all crocodyliforms more closely related to Crocodylus niloticus than to Protosuchus richardsoni, which unites Notosuchia (encompassing sebecids in consensus topologies) with the crown-group radiation of Alligatoroidea and Crocodyloidea via Neosuchia. This clade is characterized by derived features like procoelous vertebrae and enhanced cranial kinesis, marking a major diversification in the Early Jurassic. In simplified cladogram form:Crocodylomorpha
├── Basal crocodylomorphs (incl. sphenosuchians, paraphyletic)
└── [Crocodyliformes](/page/Crocodyliformes)
├── Protosuchia (basal crocodyliforms)
└── Mesoeucrocodylia
├── Notosuchia (incl. [Sebecidae](/page/Sebecidae))
└── Neosuchia
├── Thalattosuchia (debated position)
└── [Eusuchia](/page/Eusuchia)
└── Crocodylia
├── [Alligatoroidea](/page/Alligatoroidea)
├── [Gavialoidea](/page/Gavialoidea)
└── [Crocodyloidea](/page/Crocodyloidea)
Crocodylomorpha
├── Basal crocodylomorphs (incl. sphenosuchians, paraphyletic)
└── [Crocodyliformes](/page/Crocodyliformes)
├── Protosuchia (basal crocodyliforms)
└── Mesoeucrocodylia
├── Notosuchia (incl. [Sebecidae](/page/Sebecidae))
└── Neosuchia
├── Thalattosuchia (debated position)
└── [Eusuchia](/page/Eusuchia)
└── Crocodylia
├── [Alligatoroidea](/page/Alligatoroidea)
├── [Gavialoidea](/page/Gavialoidea)
└── [Crocodyloidea](/page/Crocodyloidea)