Pseudosuchia

Mounted skeleton of Postosuchus at Museum of Texas Tech University, copyrigh Dallas Krentzel.

Belongs within: Archosauriformes.
Contains: Stagonolepidae, Crocodylomorpha.

The Pseudosuchia are a clade of reptiles including the crocodilians and a number of related fossil taxa once classified as 'thecodontians' such as the aetosaurs, ornithosuchids and 'rauisuchians'. Higher relationships within the clade have differed between analyses though most recent authors have agreed that the 'rauisuchians' (Rauisuchidae, Prestosuchidae and Poposauridae) do not form a single clade. Indeed, an analysis of the archosauriforms by Nesbitt et al. (2017) did not even recover monophyly for the individual rauisuchian families.

Synapomorphies (from Benton & Clark 1988): Septomaxilla absent; lower temporal fenestra reduced in size, triangular with a dorsal point, quadratojugal sloping forward towards postorbital; axial diapophysis reduced or absent; pubo-ischiadic plate absent, pubis and ischium long and narrow, pubis expanding distally into broad forward-facing plate; long distal process of pubis bent down into subvertical orientation; pubis longer than ischium; proximal head of femur turned inwards at about 45°; fully developed crocodile-normal tarsus, capable of rotation of 60–70°; digit V in foot reduced in length, shorter than digit I.

<==Pseudosuchia (see below for synonymy) NS09
    |--Nundasuchus songeaensis NB17
    `--+--Ornithosuchidae B04
       |    |--Venaticosuchus B04
       |    |--Parringtonia gracilis R72, NB17
       |    |--Dyoplax R72
       |    |--Riojasuchus Bonaparte 1969 IN07, R72
       |    |    `--R. tenuisceps IN07
       |    `--Ornithosuchus Newton 1894 NB17, GQ01 [incl. Dasygnathoides R72]
       |         |--*O. longidens (Huxley 1877) [=Dasygnathus longidens; incl. O. woodwardi Newton 1894] GQ01
       |         `--O. rusconii P88
       `--+--+--Stagonolepidae NB17
          |  `--Revueltosaurus Hunt 1989 NS09, PI05
          |       |--R. callenderi Hunt 1989 PI05
          |       `--R. hunti Heckert 2002 PI05
          `--+--Kollamasuchus gonzalezdiazi NB17
             `--+--+--Turfanosuchus dabanensis NB17
                |  `--+--Yonghesuchus sangbiensis NB17
                |     `--Gracilisuchus Romer 1972 NB17, D07
                |          `--G. stipanicicorum GPS06
                `--+--Ticinosuchus Krebs 1965 NB17, D07 (see below for synonymy)
                   |    `--T. ferox Krebs 1965 G00
                   `--+--+--Qianosuchus mixtus NB17
                      |  `--+--+--Arizonasaurus Welles 1947 NB17, G00
                      |     |  |    `--A. babbitti NS09
                      |     |  `--Xilousuchus sapingensis NB17
                      |     `--+--Poposaurus gracilis Mehl 1915 NB17, G00
                      |        `--+--Lotosaurus Zhang 1975 NB17, D07
                      |           |    `--L. adentus Zhang 1975 G00
                      |           `--+--Sillosuchus longicervix Alcober & Parrish 1997 NB17, G00
                      |              `--+--Shuvosaurus Chatterjee 1993 NB17, D07
                      |                 |    `--S. inexpectatus G00
                      |                 `--Effigia okeeffeae NB17
                      `--+--Prestosuchus chiniquensis Huene 1942 NB17, G00 [incl. P. loricatus Huene 1942 G00]
                         `--+--Saurosuchus Reig 1959 NB17, D07
                            |    `--S. galilei Reig 1959 G00
                            `--+--Batrachotomus kupferzellensis Gower 1999 NB17, G00
                               `--+--Fasolasuchus tenax Bonaparte 1978 NB17, G00
                                  `--+--+--Rauisuchus tiradentes Huene 1942 NB17, G00
                                     |  `--+--Postosuchus NB17
                                     |     |    |--P. alisonae NB17
                                     |     |    `--P. kirkpatricki Chatterjee 1985 G00
                                     |     `--Polonosuchus silesiacus NB17
                                     `--+--Trialestes BC88
                                        |--Erpetosuchus Newton 1894 CX04, D07 [=Herpetosuchus R72]
                                        |    `--E. granti D07
                                        `--Crocodylomorpha NB17

Pseudosuchia incertae sedis:
  Belodon C77
  Strigosuchus R72
  Wangisuchus Young 1974 G00
    `--W. tzeyii G00
  Vytshegdosuchus zheshartensis Sennikov 1988 G00
  Energosuchus garjainovi Ochev 1986 G00
  Dongusia Huene 1940 G00
  Pallisteria R72
  Tsylmosuchus G00
    |--T. donensis Sennikov 1990 G00
    |--T. jakovlevi Sennikov 1990 G00
    `--T. samariensis Sennikov 1990 G00
  Episcoposaurus G00
  Dolichobrachium G00
  Chirotherium G00
  Vjushkovisaurini G00
    |--Vjushkovisaurus berdjanensis Ochev 1982 G00
    |--Jaikosuchus magnus Sennikov 1990 G00
    |--Jushatyria vjushkovi Sennikov in Kalandadze & Sennikov 1985 G00
    `--Youngosuchus Sennikov in Kalandadze & Sennikov 1985 G00
         `--*Y. sinensis (Young 1973) [=Vjushkovia sinensis] G00
  Stagonosuchus nyassicus Huene 1938 G00
  Zanclodon Plieninger 1846 G00
  Tikisuchus romeri Chatterjee & Majumdar 1987 G00
  Luperosuchus fractus Romer 1971 G00
  Hoplitosuchus raui Huene 1942 G00
  Procerosuchus Huene 1944 G00, R72
    `--P. celer Huene 1942 G00
  Heptasuchus clarki Dawley, Zawiskie & Cosgriff 1979 G00
  Lythrosuchus langstoni Long & Murray 1995 G00
  Bromsgroveia walkeri Galton 1985 G00
  Teratosaurus suevicus v. Meyer 1861 G00
  Chatterjeea [Chatterjeeidae] G00
    `--C. elegans Long & Murray 1995 G00

Nomen nudum: Mandasuchus tanyauchen Charig in Appleby et al. 1967 G00

Pseudosuchia [Crocodylotarsi, Crurotarsi, Dromaeosuchia, Poposauridae, Prestosuchidae, Procrocodylomorpha, Rauisuchia, Rauisuchidae, Rauisuchiformes, Rauisuchinae, Rauisuchini, Suchia] NS09

Ticinosuchus Krebs 1965 NB17, D07 [incl. Mandasuchus Charig in Appleby et al. 1967 (n. n.) G00; Ticinosuchinae]

*Type species of generic name indicated

REFERENCES

[B04] Benton, M. J. 2004. Origin and relationships of Dinosauria. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2^nd ed. pp. 7–19. University of California Press: Berkeley.

[BC88] Benton, M. J., & J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 295–338. Clarendon Press: Oxford.

[CX04] Clark, J. M., X. Xu, C. A. Forster & Y. Wang. 2004. A Middle Jurassic ‘sphenosuchian’ from China and the origin of the crocodylian skull. Nature 430: 1021–1024.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[GQ01] Gauthier, J., & K. de Queiroz. 2001. Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name "Aves". In: Gauthier, J., & L. F. Gall (eds) New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom pp. 7–41. Peabody Museum of Natural History, Yale University: New Haven.

[G00] Gower, D. J. 2000. Rauisuchian archosaurs (Reptilia, Diapsida): an overview. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 218 (3): 447–488.

[IN07] Irmis, R. B., S. J. Nesbitt, K. Padian, N. D. Smith, A. H. Turner, D. Woody & A. Downs. 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317: 358–361.

[NB17] Nesbitt, S. J., R. J. Butler, M. D. Ezcurra, P. M. Barrett, M. R. Stocker, K. D. Angielczyk, R. M. H. Smith, C. A. Sidor, G. Niedźwiedzki, A. G. Sennikov & A. J. Charig. 2017. The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature 544: 484–487.

[NS09] Nesbitt, S. J., N. D. Smith, R. B. Irmis, A. H. Turner, A. Downs & M. A. Norell. 2009. A complete skeleton of a Late Triassic saurischian and the early evolution of dinosaurs. Science 326: 1530–1533.

[PI05] Parker, W. G., R. B. Irmis, S. J. Nesbitt, J. W. Martz & L. S. Browne. 2005. The Late Triassic pseudosuchian Revueltosaurus callenderi and its implications for the diversity of early ornithischian dinosaurs. Proceedings of the Royal Society of London Series B—Biological Sciences 272: 963–969.

[P88] Paul, G. S. 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon & Schuster: New York.

[R72] Romer, A. S. 1972. The Chañares (Argentina) Triassic reptile fauna. XVI. Thecodont classification. Breviora 395: 1–24.

Pholidosauridae

Reconstruction of Dyrosaurus phosphaticus, copyright Nobu Tamura.

Belongs within: Neosuchia.

The Pholidosauridae are a group of crocodyliforms known from the Cretaceous, most of which are slender-snouted, fish-eating specialists adapted to near-shore marine habitats (Sereno et al. 2001).

<==Pholidosauridae
    |--Pholidosaurus decipiens SL01, GPS06
    `--+--+--Terminonaris ropbusta SL01, GPS06
       |  `--Sarcosuchus Broin & Tarquet 1966 SL01, D07
       |       |--S. hartti SL01
       |       `--S. imperator Broin & Taquet 1966 SL01
       `--+--Sokotosuchus ianwilsoni GPS06
          `--Dyrosauridae GPS06
               |--Congosaurus LB02
               |--Rhabdognathus SF03
               |--Hyposaurus rogersii GPS06
               |--Chenanisuchus lateroculi GPS06
               `--Dyrosaurus Thomas 1893 SL01, D07
                    |--D. minor SF03
                    `--D. phosphaticus GPS06

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3–47.

[SL01] Sereno, P. C., H. C. E. Larsson, C. A. Sidor & B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.

Thalattosuchia

Reconstruction of Geosaurus giganteus scaled to a human diver, from Young et al. (2012).

Belongs within: Neosuchia.

The Thalattosuchia are a clade of marine crocodyliforms known from the Jurassic. Basal members of the clade had an extremely long and tubular hyperdentate snout though this became striking modified in some derived members of the clade (Gasparini et al. 2006).

<==Thalattosuchia
    |--Teleosauridae SF03
    |    |--Steneosaurus bollensis LB02, GPS06
    |    |--Machimosaurus LB02
    |    |--Pelagosaurus Brown 1841 LB02, SF03
    |    |    `--P. typus GPS06
    |    `--Teleosaurus Deslongchamps 1866 D07
    |         |--T. chapmanni D07
    |         |--T. hastifer D07
    |         `--T. mandelslohi D07
    `--Metriorhynchidae GPS06
         |--Metriorhynchus Meyer 1830 GPS06, D07
         |    |--M. casamiquelai GPS06
         |    |--M. moreli D07
         |    `--M. superciliosus GPS06
         `--+--Dakosaurus GPS06
            |    |--D. andiniensis GPS06
            |    `--D. maximus GPS06
            `--Geosaurus Cuvier 1842 GPS06, D07
                 |--G. araucanensis GPS06
                 |--G. cynodus Gervais 1867–1869 C07
                 |--G. giganteus D07
                 |--G. gracilis D07
                 |--G. suevicus GPS06
                 `--G. vignaudi D07

*Type species of generic name indicated

REFERENCES

[C07] Case, E. C. 1907. Revision of the Pelycosauria of North America. Carnegie Institute of Washington Publication 55: 1–176, 35 pls.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3–47.

Neosuchia

Spectacled caiman Caiman crocodilus, copyright Berrucomons.

Belongs within: Metasuchia.
Contains: Thalattosuchia, Pholidosauridae, Crocodylidae.

The Neosuchia are a clade of crocodylomorphs that has been defined as including all taxa closer to Crocodylus niloticus than to Notosuchus terrestris. This clade includes all living crocodilians.

<==Neosuchia [Metamesosuchia] SF03
    |  i. s.: Susisuchus Salisbury, Frey et al. 2003 [Susisuchidae] SF03
    |           `--*S. anatoceps Salisbury, Frey et al. 2003 SF03
    |         Brillanceausuchus Michard et al. 1990 SF03, LB02
    |           `--B. babouriensis SF03
    |         Gilchristosuchus palatinus SF03
    |--Atoposauridae GPS06
    |    |--Theriosuchus pusillus GPS06
    |    |--Montsechosuchus depereti [=Alligatorium depereti] BC88
    |    `--Alligatorium [incl. Alligatorellus, Atoposaurus] BC88
    |         `--A. meyeri BC88
    `--+--+--Thalattosuchia GPS06
       |  `--Pholidosauridae GPS06
       `--+--Goniopholidae GPS06
          |    |--Sunosuchus SL01
          |    |--Eutretauranosuchus delfsi GPS06
          |    `--Goniopholis CRA04
          |         |--G. gracilidens [=Nannosuchus gracilidens] BC88
          |         `--G. paulistanus Roxo 1936 CRA04
          `--+--Shamosuchus Mook 1924 BC88, SF03 [incl. Paralligator BC88]
             |--Bernissartia Dollo 1885 GPS06, N85
             |    `--B. fagesii Dollo 1883 SF03
             `--Eusuchia GPS06
                  |--Borealosuchus formidabilis GPS06
                  |--Leidyosuchus BC88
                  |--Hylaeochampsa Owen 1874 GPS06, BC88
                  |    `--H. vectiana GPS06
                  |--Stomatosuchus Stromer 1925 [Stomatosuchidae] BC88
                  |    `--S. inermis SF03
                  `--Crocodylia BC88
                       |  i. s.: Diplocynodon RB05
                       |           |--D. darwini RRO00
                       |           |--D. hantoniensis RRO00
                       |           `--D. styriacus RB05
                       |         Pallimnarchus pollens F71
                       |         Rimasuchus Storrs 2003 LA04
                       |           `--*R. lloydi (Fourtau 1918) [=Crocodylus lloydi] LA04
                       |--+--Gavialidae AS09
                       |  |    |--Gavialis gangeticus GPS06
                       |  |    `--Tomistoma Müller 1846 AS09, LA04 [Tomistominae]
                       |  |         |--T. coppensi Pickford 1994 LA04
                       |  |         |--T. dowsoni Fourtau 1918 LA04
                       |  |         |--T. lusitanica Antunes 1967 LA04
                       |  |         `--T. schlegelii USDI77
                       |  `--Eosuchus lerichei GPS06
                       `--+--+--Crocodylidae GPS06
                          |  `--Osteolaemus tetraspis BC88, USDI77
                          |       |--O. t. tetraspis USDI77
                          |       `--O. t. osborni USDI77
                          `--Alligatoroidea D07
                               |  i. s.: Deinosuchus Colbert & Bird 1958 D07 [incl. Phobosuchus N85]
                               |           |--D. hatchery D07
                               |           |--D. riograndensis D07
                               |           `--D. rugosus D07
                               `--Alligatorinae ML14
                                    |--Navajosuchus mooki ML14
                                    `--+--+--Stangerochampsa mccabei AS09
                                       |  `--Alligator GPS06
                                       |       |--A. mississippiensis Daudin 1801 NS10
                                       |       `--A. sinensis SF03
                                       `--+--Paleosuchus BC88
                                          |    |--P. palpebrosus DS86
                                          |    `--P. trigonatus DS86
                                          `--Caiman BC88
                                               |  i. s.: C. sclerops GWN71
                                               |         C. yacare USDI77
                                               |--C. latirostris BC88
                                               `--+--C. crocodilus BC88
                                                  |    |--C. c. crocodilus B81
                                                  |    |--C. c. apaporiensis USDI77
                                                  |    `--C. c. fuscus B81
                                                  `--Melanosuchus niger BC88, DS86

Inorganic: Caiman crocodilus protominilorientalus Okamura 1987 O87

*Type species of generic name indicated

REFERENCES

[AS09] Alfaro, M. E., F. Santini, C. Brock, H. Alamillo, A. Dornburg, D. L. Rabosky, G. Carnevale & L. J. Harmon. 2009. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proceedings of the National Academy of Sciences of the USA 106 (32): 13410–13414.

[BC88] Benton, M. J., & J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 295–338. Clarendon Press: Oxford.

[B81] Brooks, D. R. 1981. Hennig's parasitological method: a proposed solution. Systematic Zoology 30 (3): 229–249.

[CRA04] Carvalho, I. de S., L. C. B. Ribeiro & L. dos S. Avilla. 2004. Uberabasuchus terrificus sp. nov., a new Crocodylomorpha from the Bauru Basin (Upper Cretaceous), Brazil. Gondwana Research 7 (4): 975–1002.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[DS86] Dixon, J. R., & P. Soini. 1986. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Milwaukee Public Museum: Milwaukee.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[GWN71] Gorman, G. C., A. C. Wilson & M. Nakanishi. 1971. A biochemical approach towards the study of reptilian phylogeny: evolution of serum albumin and lactic dehydrogenase. Systematic Zoology 20 (2): 167–185.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[LA04] Llinás Agrasar, E. 2004. Crocodile remains from the Burdigalian (lower Miocene) of Gebel Zelten (Libya). Geodiversitas 26 (2): 309–321.

[ML14] Mitchell, K. J., B. Llamas, J. Soubrier, N. J. Rawlence, T. H. Worthy, J. Wood, M. S. Y. Lee & A. Cooper. 2014. Ancient DNA reveals elephant birds and kiwi are sister taxa and clarifies ratite bird evolution. Science 344 (6186): 898–900.

[NS10] Nesbitt, S. J., C. A. Sidor, R. B. Irmis, K. D. Angielczyk, R. M. H. Smith & L. A. Tsuji. 2010. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464: 95–98.

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[RB05] Rage, J.-C., & S. Bailon. 2005. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France). Geodiversitas 27 (3): 413–441.

[RRO00] Rossmann, T., M. Rauhe & F. Ortega. 2000. Studies on Cenozoic crocodiles: 8. Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions. Paläontologische Zeitschrift 74: 379–392.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3-47.

[SL01] Sereno, P. C., H. C. E. Larsson, C. A. Sidor & B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

Crocodylidae

American crocodile Crocodylus acutus, photographed by Tomascastelazo.

Belongs within: Neosuchia.

The family Crocodylidae includes the crocodiles Crocodylus and their closest fossil relatives. The false gharial Tomistoma schlegelii may also be a member of this group, though its phylogenetic position has been open to debate. Members of this family have relatively narrow, elongate heads. The largest living species, the saltwater crocodile Crocodylus porosus, may reach lengths of well over six metres. Other species may be considerably smaller; the Australian freshwater crocodile C. johnstoni, for instance, is usually less than two metres in length.

<==Crocodylidae
    |--Rhamphosuchus SL01
    |--Euthecodon PS01
    |    |--E. arambourgi Ginsburg & Buffetaut 1978 LA04
    |    |--E. brumpti Joleaud 1920 LA04
    |    `--E. nitriae Fourtau 1920 LA04
    `--Crocodylus MA04
         |  i. s.: C. affinis C77
         |         C. chamensis Cope 1874 C77
         |         C. elliottii C77
         |         C. grypus Cope 1875 C77
         |         C. heterodon C77
         |         C. liodon Marsh 1871 C77
         |         C. megarhinus SF03
         |         C. (Botosaurus) selaslophensis Etheridge 1917 F71
         |         C. wheelerii Cope 1875 C77
         |--+--C. johnstoni B81, SWS14
         |  `--+--C. porosus B81
         |     `--+--C. palustris B81
         |        |    |--C. p. palustris USDI77
         |        |    `--C. p. kimbula USDI77
         |        `--+--C. novaeguineae B81
         |           |    |--C. n. novaeguineae USDI77
         |           |    `--C. n. mindorensis USDI77
         |           `--C. siamensis B81
         `--+--+--C. cataphractus B81
            |  `--C. niloticus Laurenti 1768 B81, MA04
            `--+--+--C. acutus B81
               |  `--C. rhombifer B81
               `--+--C. intermedius B81
                  `--C. moreletii B81

*Type species of generic name indicated

REFERENCES

[B81] Brooks, D. R. 1981. Hennig's parasitological method: a proposed solution. Systematic Zoology 30 (3): 229–249.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[LA04] Llinás Agrasar, E. 2004. Crocodile remains from the Burdigalian (lower Miocene) of Gebel Zelten (Libya). Geodiversitas 26 (2): 309–321.

[MA04] Modesto, S. P., & J. S. Anderson. 2004. The phylogenetic definition of Reptilia. Systematic Biology 53 (5): 815–821.

[PS01] Pickford, M., & B. Senut. 2001. The geological and faunal context of Late Miocene hominid remains from Lukeino, Kenya. Comptes Rendus de L’Academie des Sciences—Series IIA—Earth and Planetary Sciences 332: 145–152.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3–47.

[SL01] Sereno, P. C., H. C. E. Larsson, C. A. Sidor & B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.

[SWS14] Somaweera, R., D. Woods & T. Sonneman. 2014. A note on the Australian freshwater crocodiles inhabiting Tunnel Creek cave, West Kimberley. Records of the Western Australian Museum 29 (1): 82–87.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants—republication of list of species. Federal Register 42: 36420–36431.

Last updated: 6 April 2017.

Metasuchia

Reconstruction of Araripesuchus wegeneri, by Todd Marshall.

Belongs within: Crocodyliformes.
Contains: Neosuchia.

The Metasuchia were defined by Sereno et al. (2001) as the most exclusive clade containing both recent crocodilians and Notosuchus. This differs somewhat from its original usage by Benton & Clark (1988) for all members of the clade Mesoeucrocodylia excluding Thalattosuchia, which have subsequently been classified as derived metasuchians.

The Notosuchia are a group of often small, terrestrial crocodyliforms known from the Cretaceous of Africa and South America. Many notosuchians were markedly heterodont, with the rear teeth become multicusped and/or molariform (O'Connor et al. 2010). The Peirosauridae have been recognised as a group of moderately high-snouted crocodyliforms from the same time and area; it is possible that this group is paraphyletic to the later neosuchians (O'Connor et al. 2010).

<==Metasuchia (see below for synonymy)
    |  i. s.: Brasileosaurus pachecoi Huene 1931 CRA04
    |         Elosuchidae LB02
    |           |--Stolokrosuchus Larsson & Gado 2000 LB02
    |           |    `--S. lapparenti Larsson & Gado 2000 LB02
    |           `--Elosuchus de Lapparent de Broin 2002 LB02
    |                |--*E. cherifiensis (Lavocat 1955) (see below for synonymy) LB02
    |                `--E. felixi de Lapparent de Broin 2002 LB02
    |--Peirosauridae [Lomasuchinae] CRA04
    |    |--+--Peirosaurus tormini Price 1955 OS10, CRA04
    |    |  `--Lomasuchus Gasparini, Chiappe & Fernandez 1991 OS10, CRA04
    |    |       `--L. palpebrosus Gasparini, Chiappe & Fernandez 1991 CRA04
    |    `--+--+--Anatosuchus minor OS10
    |       |  `--Neosuchia OS10
    |       `--Mahajangasuchini CRA04
    |            |--Mahajangasuchus insignis OS10
    |            `--Uberabasuchus Carvalho, Ribeiro & Avilla 2004 CRA04
    |                 `--*U. terrificus Carvalho, Ribeiro & Avilla 2004 CRA04
    `--+--Araripesuchus Price 1959 OS10, SF03
       |    |--A. wegeneri OS10
       |    `--+--A. buitreraensis OS10
       |       `--+--A. gomesii Price 1959 OS10, CRA04
       |          `--A. patagonicus Ortega et al. 2001 GPS06, LB02
       `--Notosuchia OS10
            |--Uruguaysuchus [Uruguaysuchidae] OS10
            |    `--U. aznarezi GPS06
            `--Ziphosuchia [Baurusuchoidea, Sebecosuchia] CRA04
                 |  i. s.: Bergisuchus Kuhn 1968 [Bergisuchidae] RRO00
                 |           `--*B. dietrichbergi Kuhn 1968 RRO00
                 |         Ilchunaia parca Rusconi 1946 RRO00
                 |         Cynodontosuchus rothi Woodward 1896 RRO00
                 |         Ayllusuchus fernandezi Gasparini 1984 RRO00
                 |--+--Sebecidae CRA04
                 |  |    |--Sebecus LB02
                 |  |    |    |--S. huilensis RRO00
                 |  |    |    |--S. icaeorhinus RRO00
                 |  |    |    `--S. querejazus Buffetaut & Marshall 1991 LB02
                 |  |    `--Libycosuchus Stromer 1914 OS10, SF03 [Libycosuchidae]
                 |  |         `--L. brevirostris GPS06
                 |  `--Candidodontidae CRA04
                 |       |--+--Mariliasuchus amarali Carvalho & Bertini 1999 OS10, CRA04
                 |       |  `--+--Adamantinasuchus navae OS10
                 |       |     `--Pakasuchus O’Connor, Sertich et al. 2010 OS10
                 |       |          `--*P. kapilimai O’Connor, Sertich et al. 2010 OS10
                 |       `--+--Candidodon itapecuruense OS10
                 |          `--Itasuchidae CRA04
                 |               |--Malawisuchus mwakasyungutiensis OS10
                 |               `--Itasuchus CRA04
                 |                    |--I. camposi SF03 [=Cairisuchus camposi RRO00]
                 |                    `--I. jesuinoi Price 1955 CRA04
                 `--Chimaerasuchidae CRA04
                      |--Simosuchus Buckley 2000 OS10, D07
                      |    `--S. clarki OS10
                      `--+--Notosuchus Woodward 1896 OS10, SF03 [Notosuchidae]
                         |    `--N. terrestris OS10
                         `--+--+--Chimaeresuchus Wu & Suess 1995 OS10, D07
                            |  |    `--C. paradoxus GPS06
                            |  `--Sphagesaurus Price 1950 OS10, D07
                            |       `--S. huenei Price 1950 CRA04
                            `--+--Comahuesuchus brachybuccalis OS10, GPS06
                               `--+--+--Bretesuchus [Bretesuchidae] GPS06
                                  |  |    `--B. bonapartei Gasparini et al. 1993 RRO00
                                  |  `--Iberosuchus macrodon GPS06
                                  `--Baurusuchidae CRA04
                                       |--Stratiosuchus maxhechti (Campos et al. 2001) CRA04
                                       `--Baurusuchus Prince 1945 OS10, D07
                                            `--B. pachecoi Price 1945 CRA04

Elosuchus cherifiensis (Lavocat 1955) [=Thoracosaurus cherifiensis, Thoracosuchus (l. c.) cherifiensis, Eleiosuchus africanus ms] LB02

Metasuchia [Gondwanasuchia, Notosuchiformes, Notosuchimorpha, Peirosauriformes, Peirosaurimorpha, Peirosauroidea, Terriasuchia]

*Type species of generic name indicated

REFERENCES

Benton, M. J., & J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 295–338. Clarendon Press: Oxford.

[CRA04] Carvalho, I. de S., L. C. B. Ribeiro & L. dos S. Avilla. 2004. Uberabasuchus terrificus sp. nov., a new Crocodylomorpha from the Bauru Basin (Upper Cretaceous), Brazil. Gondwana Research 7 (4): 975–1002.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[OS10] O'Connor, P. M., J. J. W. Sertich, N. J. Stevens, E. M. Roberts, M. D. Gottfried, T. L. Hieronymus, Z. A. Jinnah, R. Ridgely, S. E. Ngasala & J. Temba. 2010. The evolution of mammal-like crocodyliforms in the Cretaceous period of Gondwana. Nature 466: 748–751.

[RRO00] Rossmann, T., M. Rauhe & F. Ortega. 2000. Studies on Cenozoic crocodiles: 8. Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions. Paläontologische Zeitschrift 74: 379–392.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formation of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3–47.

Sereno, P. C., H. C. E. Larsson, C. A. Sidor & B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.

Last updated: 3 August 2017.

Crocodyliformes

Fossilised skeleton and reconstruction of Protosuchus richardsoni, photographed by Ryan Somma.

Belongs within: Crocodylomorpha.
Contains: Metasuchia.

The Crocodyliformes correspond to the 'Crocodylia' of many earlier authors (with the latter generally now restricted to the crown group of crocodilians). The basalmost forms, the protosuchians, were still relatively lightly built taxa, though undeniably crocodile-like. Members of the basal lineage including modern species, the Mesoeucrocodylia, are united by the fusion of the frontal bones of the skull, together with the development of a secondary palate from the posterior extension of the palatine bones.

Synapomorphies (from Benton & Clark 1988): Otoccipital contacting ventromedial part of quadrate, enclosing carotid artery and forming passage for cranial nerves IX-XI; basisphenoid rostrum dorsoventrally expanded; basipterygoid process reduced, basipterygoid joint closed suturally; pterygoid extending dorsally, forming ventrolateral edge of trigeminal foramen; otoccipitals broadly meeting dorsal to foramen magnum; eustachian tubes enclosed between basioccipital and basisphenoid; antorbital fenestra much smaller than orbit; premaxilla and maxilla sutured together along butt joint; parietal lacking broad occipital portion; ventromedial end of coracoid anteroposteriorly expanded, anterior edge concave; quadrate hollow with several fenestrae on dorsal surface; mastoid antrum extending through supraoccipital; two large palpebrals present; dorsal head of quadrate contacting laterosphenoid; skull table in temporal region with nearly flat dorsal surface; postorbital medial to jugal on postorbital bar; scapula dramatically broadening dorsally.

Crocodyliformes [Crocodilia]
    |--Protosuchia SL01
    |    |--Orthosuchus stormbergi GPS06
    |    `--Protosuchidae [Stegomosuchidae] GPS06
    |         |  i. s.: Notochampsa R72
    |         |         Erythrochampsa R72
    |         |         Stegomosuchus R72
    |         |         Platyognathus R72
    |         |         Microchampsa R72
    |         |--Edentosuchus tienshanensis GPS06
    |         `--+--Hemiprotosuchus leali GPS06
    |            `--Protosuchus Brown 1934 GPS06, D07 (see below for synonymy)
    |                 |--P. haughtoni D07
    |                 |--P. micmac D07
    |                 `--P. richardsoni GPS06
    `--Mesoeucrocodylia [Mesosuchia] SL01
         |  i. s.: Brachychampsa Gilmore 1911 D07
         |           |--B. montana SF03
         |           `--B. sealeyi D07
         |--+--Zaraasuchus shepardi GPS06
         |  `--Gobiosuchus [Gobiosuchidae] GPS06
         |       `--G. kielanae GPS06
         `--+--+--Hsisosuchus chungkingensis GPS06
            |  `--Metasuchia GPS06
            `--+--Zosuchus davidsoni GPS06
               `--+--Shantungosuchus hangjinensis GPS06
                  `--Sichuanosuchus shuhanensis GPS06

Crocodyliformes incertae sedis:
  Mekosuchinae D07
    |--Mekosuchus inexpectatus WH02
    `--Quinkana Molnar 1961 D07
         |--Q. babarra D07
         |--Q. fortirostrum D07
         |--Q. meboldi D07
         `--Q. timera D07
  Trematochampsidae RRO00
    |--Trematochampsa taqueti Buffetaut 1974 LB02
    |--Eremosuchus elkoholicus RRO00
    |--Hamadasuchus rebouli RRO00
    `--Amargasuchus minor RRO00
  Crocidileimus robustus Gervais 1873 LB02
  Diplocynodus C77
    |--D. sphenops Cope 1875 C77
    `--D. subulatus C77
  Allodaposuchus precedens Nopcsa 1928 PST99
  Dolichochampsa minima SF03
  Aigialosuchus Persson 1959 LS04
  Pristichampsus rollinatii RRO00
  Asiatosuchus depressifrons RRO00

Protosuchus GPS06 [incl. Archaeosuchus R72, Baroqueosuchus Busbey & Gow 1984 BC88, Lesothosuchus Whetstone & Whybrow 1983 BC88]

*Type species of generic name indicated

REFERENCES

[BC88] Benton, M. J., & J. M. Clark. 1988. Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 295–338. Clarendon Press: Oxford.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GPS06] Gasparini, Z., D. Pol & L. A. Spalletti. 2006. An unusual marine crocodyliform from the Jurassic–Cretaceous boundary of Patagonia. Science 311: 70–73.

[LB02] Lapparent de Broin, F. de. 2002. Elosuchus, a new genus of crocodile from the Lower Cretaceous of the north of Africa. Comptes Rendus Palevol 1: 275–285.

[LS04] Lindgren, J., & M. Siverson. 2004. The first record of the mosasaur Clidastes from Europe and its palaeogeographical implications. Acta Palaeontologica Polonica 49 (2): 219–234.

[PST99] Pereda Suberbiola, X., & P. Taquet. 1999. Restes de Rhabdodon (dinosaure ornithopode) de Transylvanie donnés par Nopcsa au Muséum national d'Histoire naturelle de Paris. Geodiversitas 21 (2): 157–166.

[R72] Romer, A. S. 1972. The Chañares (Argentina) Triassic reptile fauna. XVI. Thecodont classification. Breviora 395: 1–24.

[RRO00] Rossmann, T., M. Rauhe & F. Ortega. 2000. Studies on Cenozoic crocodiles: 8. Bergisuchus dietrichbergi Kuhn (Sebecosuchia: Bergisuchidae n. fam.) from the Middle Eocene of Germany, some new systematic and biological conclusions. Paläontologische Zeitschrift 74: 379–392.

[SF03] Salisbury, S. W., E. Frey, D. M. Martill & M. C. Buchy. 2003. A new crocodilian from the Lower Cretaceous Crato Formatioin of north-eastern Brazil. Palaeontographica Abteilung A: Paläozoologie—Stratigraphie 270: 3–47.

[SL01] Sereno, P. C., H. C. E. Larsson, C. A. Sidor & B. Gado. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516–1519.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

Last updated: 3 August 2017.