Anura

Hochstetter's frog Leiopelma hochstetteri, copyright David M. Green.

Belongs within: Lepospondyli.
Contains: Alytoidea, Pipoidea, Pelobatoidea, Neobatrachia.

The Anura, frogs and toads, are a diverse group of terrestrial to aquatic vertebrates that lack an external tail at maturity and are often adapted for a hopping mode of locomotion. Larvae, if present, are aquatic and legless with a strong, finned tail.

Synapomorphies (from Frost et al. 2006): Prefrontal bone absent; prearticular bone absent; palatine absent; nine or fewer vertebrae; atlas with a single centrum; first spinal nerve exiting from spinal nerve canal via intervertebral foramen; caudal vertebral segments fused into urostyle; hindlimbs significantly longer than forelimbs, including elongation of ankle bones; radius and ulna, tibia and fibula fused; hyobranchial elements fused into hyoid plate; keratinous jaw sheaths and keratodonts present on larval mouthparts; larva with single median spiracle; skin with large subcutaneous lymph spaces; two m. protractor lentis attached to lens.

<==Anura (see below for synonymy)
    |--Bombinanura [Archaeobatrachia, Discoglossanura, Lalagobatrachia, Mesobatrachia, Ranoidei, Sokolanura] FG06
    |    |--Alytoidea FB17
    |    `--Pipanura FB17
    |         |--Pipoidea PW11
    |         `--Acosmanura FG06
    |              |  i. s.: Eurycephalella alcinae FB17
    |              |--Pelobatoidea EJK08
    |              `--Neobatrachia PW11
    `--Leiopelmatoidea FB17
         |--Ascaphus Stejneger 1899 FG06 [Ascaphidae PW11]
         |    |--A. montanus FG06
         |    `--A. truei FG06
         `--Leiopelmatidae [Amphicoela, Liopelmatina] FG06
             |--Vieraella herbstii Reig 1961 G88, M93
             |--Notobatrachus degiustoi Reig 1955 G88, M93
             `--Leiopelma Fitzinger 1861 [=Liopelma Günther 1869] FG06
                  |  i. s.: L. waitomoensis Worthy 1987 WH02
                  |--+--L. auroraensis Worthy 1987 WH02
                  |  |--L. hochstetteri Fitzinger 1861 WH02
                  |  `--L. markhami Worthy 1987 WH02
                  `--+--L. pakeka Bell, Daugherty & Hay 1998 BDH98
                     `--+--L. archeyi Turbott 1942 BDH98, WH02
                        `--L. hamiltoni McCulloch 1919 BDH98

Anura incertae sedis:
  Phractops F13
    |--P. brevipalmatus F13
    `--P. brevipes F13
  Callula baleata M89
  Notodelphys ovifera H04
  Liaobatrachus FB17
    |--L. grabaui ZBH03
    `--L. zhaoi FB17
  Enneabatrachus hechti R00
  Eobatrachus agilis Marsh 1887 (n. d.) R00
  Comobatrachus aenigmatis Hecht & Estes 1960 (n. d.) R00
  Paradiscoglossus americanus Estes & Sanchíz 1982 R00
  Theatonius lancensis Fox 1976 R00
  Gobiatidae R00
    |--Gobiatoides parvus Roček & Nessov 1993 R00
    `--Gobiates Špinar & Tatarinov 1986 R00
         |--G. asiaticus Roček & Nessov 1993 R00
         |--G. bogatchovi Roček & Nessov 1993 R00
         |--G. dzhyrakudukensis Roček & Nessov 1993 R00
         |--G. fritschi Roček & Nessov 1993 R00
         |--G. furcatus Roček & Nessov 1993 R00
         |--G. kermeentsavi Špinar & Tatarinov 1986 R00
         |--G. kizylkumensis Roček & Nessov 1993 R00
         |--G. leptocolaptus (Borsuk-Białynicka 1978) R00
         |--G. sosedkoi (Nessov 1981) [=Eopelobates sosedkoi] R00
         |--G. spinari Roček & Nessov 1993 R00
         `--G. tatarinovi Roček & Nessov 1993 R00
  Altanulia alifanovi Gubin 1993 R00
  Palaeophrynos gessneri Tschudi 1839 RR00
  Batrachus RR00
    |--B. lacustris Pomel 1853 RR00
    |--B. lemanensis Pomel 1853 RR00
    `--B. naiadum Pomel 1853 RR00
  Lutetiobatrachus gracilis RR00
  Liventsovkia jucunda RR00
  Ranavus scarabellii RR00

Anura [Acaudata, Anoura, Anuri, Anuria, Discoglossoidei, Ecaudata, Ecaudati, Heteromorpha, Leiopelmatanura, Miura, Pygomolgaei, Raniformia]

*Type species of generic name indicated

REFERENCES

[BDH98] Bell, B. D., C. H. Daugherty & J. M. Hay. 1998. Leiopelma pakeka, n. sp. (Anura: Leiopelmatidae), a cryptic species of frog from Maud Island, New Zealand, and a reassessment of the conservation status of L. hamiltoni from Stephens Island. Journal of the Royal Society of New Zealand 28 (1): 39–54.

[EJK08] Evans, S. E., M. E. H. Jones & D. W. Krause. 2008. A giant frog with South American affinities from the Late Cretaceous of Madagascar. Proceedings of the National Academy of Sciences of the USA 105 (8): 2951–2956.

[FB17] Feng, Y.-J., D. C. Blackburn, D. Liang, D. M. Hillis, D. B. Wake, D. C. Cannatella & P. Zhang. 2017. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 114 (29): E5864–E5870.

[FG06] Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green & W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1–370.

[F13] Fry, D. B. 1913. On a Varanus and a frog from Burnett River, Queensland, and a revision of the variations in Limnodynastes dorsalis, Gray. Records of the Australian Museum 10 (2): 17–34, pls 1–3.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[M93] Milner, A. R. 1993. Amphibian-grade Tetrapoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 665–679. Chapman & Hall: London.

[M89] Modigliani, E. 1889. Materiali per la fauna erpetologica dell’isola Nias. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 113–124.

[PW11] Pyron, R. A., & J. J. Wiens. 2011. A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution 61: 543–583.

[R00] Roček, Z. 2000. Mesozoic anurans. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1295–1331. Surrey Beatty & Sons.

[RR00] Roček, Z., & J.-C. Rage 2000. Tertiary Anura of Europe, Africa, Asia, North America, and Australia. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1332–1387. Surrey Beatty & Sons.

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

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807–814.

Sphenacodontidae

Pair of mounted skeletons of Dimetrodon, copyright D'Arcy Norman.

Belongs within: Synapsida.
Contains: Therapsida.

The Sphenacodontidae are a group of carnivorous 'pelycosaurs' known from the later Carboniferous to the Middle Permian, suggested by Kemp (1988) to be paraphyletic to the therapsids.

<==Sphenacodontidae K88
    |  i. s.: Secodontosaurus BH04
    |         Sphenacodon Marsh 1878 D07
    |           |--S. ferocior D07
    |           `--S. ferox Marsh 1878 C07
    |--Haptodus K88
    `--+--Therapsida K88
       `--Dimetrodon Cope 1878 K88, BH04
            |--*D. incisivus Cope 1878 (see below for synonymy) C07
            |--D. angelensis D07
            |--D. biradicatus C07
            |--D. dollovianus (Cope 1884) [=Embolophorus dollovianus] C07
            |--D. giganhomogenes Case 1907 C07
            |--D. gigas (Cope 1878) [=Clepsydrops gigas] C07
            |--D. grandis N85
            |--D. limbatus BH04
            |--D. longiramus Case 1907 C07
            |--D. macrospondylus (Cope 1884) [=Clepsydrops macrospondylus] C07
            |--D. milleri BH04
            |--D. natalis BH04
            |--D. navajovicus Case 1907 C07
            |--D. obtusidens (Cope 1880) [=Theropleura obtusidens] C07
            |--D. occidentalis BH04
            |--D. platycentrus Case 1907 C07
            `--D. teutonis Berman et al. 2001 BH04

*Dimetrodon incisivus Cope 1878 [incl. Clepsydrops limbatus Cope 1877, C. (Embolophorus) limbatus; incl. Dimetrodon rectiformis Cope 1878, D. semiradicatus Cope 1881] C07

*Type species of generic name indicated

REFERENCES

[BH04] Berman, D. S., A. C. Henrici, S. S. Sumida & T. Martens. 2004. New remains of Dimetrodon teutonis (Synapsida: Sphenacodontidae) from the Lower Permian of Germany. Annals of Carnegie Museum 73 (2): 48–56.

[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.

[K88] Kemp, T. S. 1988. Interrelationships of the Synapsida. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 2. Mammals pp. 1–22. Clarendon Press: Oxford.

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

Hueneosauria

Reconstruction of Cymbospondylus natans, copyright Teratophoneus.

Belongs within: Ichthyosauria.
Contains: Thunnosauria.

The Hueneosauria are a clade of ichthyosaurs defined by Maisch & Matzke (2000) as the most exclusive clade including Mixosaurus cornalianus and Ophthalmosaurus icenicus.

Synapomorphies (from Maisch & Matzke 2000): External naris placed largely laterally; anterior margin of orbit regularly rounded; ramus occipitalis of supratemporal well developed; processus transversus pterygoidei absent; posterior dorsal centra discoidal; sacral ribs indistinguishable; caudal vertebral centra short; interclavicle of triradiate shape; radius and ulna distinctly shortened; posterior margin of ulna convex or straight; distal carpals II–IV of similar size; metacarpal V with straight or concave posterior margin or absent; pubis and ischium with strongly convex medial margins, not forming symphysis; less than five toes present.

<==Hueneosauria
    |  i. s.: Guizhouichthyosaurus tangae Cao & Lu in Yin et al. 2000 (see below for synonymy) JH05
    |--+--Wimanius Maisch & Matzke 1998 MM03, MM00
    |  |    `--*W. odontopalatus Maisch & Matzke 1998 MM00
    |  `--Mixosauridae JH05
    |       |--Barracudasaurus Jiang, Hao et al. 2005 JH05
    |       |    `--*B. maotaiensis (Young 1965) [=Mixosaurus maotaiensis] JH05
    |       `--+--Mixosaurus Baur 1887 JH05, MM00 [incl. Sangiorgiosaurus Brinkmann 1998 MM00]
    |          |    |--*M. cornalianus (Bassani 1886) MM00
    |          |    `--M. kuhnschnyderi (Brinkmann 1998) MM00 [=Sangiorgiosaurus kuhnschnyderi H02]
    |          `--Phalarodontinae JH05
    |               |--Contectopalatus Maisch & Matzke 1998 JH05, MM00
    |               |    `--*C. atavus (Quenstedt 1852) MM00, H02 [=Mixosaurus atavus H02]
    |               `--Phalarodon Merriam 1910 JH05, MM00
    |                    |--*P. nordenskioeldii (Hulke 1873) MM00 [=Mixosaurus nordenskioldi H02]
    |                    |--P. atavus FF13
    |                    |--P. callawayi FF13
    |                    |--P. fraasi FF13
    |                    `--P. major (von Huene 1916) MM00
    `--Longipinnati [Euichthyosauria, Shastasauridae] MM00
         |--‘Cymbospondylus’ buchseri Sander 1989 FF13, MM00
         `--+--Cymbospondylidae MM00
            |    |--Phantomosaurus Maisch & Matzke 2000 MM03, MM00
            |    |    `--*P. neubigi (Sander 1997) [=Shastasaurus neubigi] MM00
            |    `--Cymbospondylus Leidy 1868 FF13, MM00
            |         |--*C. piscosus Leidy 1868 MM00
            |         |--C. germanicus D07
            |         |--C. grandis D07
            |         |--C. natans D07
            |         |--C. nevadanus D07
            |         |--C. parvus D07
            |         |--C. petrinus Leidy 1868 MM00
            |         `--C. piscosus D07
            `--+--+--Californosaurus Kuhn 1934 FF13, MM00 [incl. Delphinosaurus Merriam 1905 (preoc.) MM00]
               |  |    `--*C. perrini (Merriam 1902) MM00
               |  `--Toretocnemidae MM00
               |       |--Qianichthyosaurus Li 1999 MM03, MM00
               |       |    `--*Q. zhoui Li 1999 MM00
               |       `--Toretocnemus Merriam 1903 FF13, MM00 (see below for synonymy)
               |            |--*T. californicus Merriam 1903 MM00
               |            `--T. zitteli (Merriam 1903) MM00
               `--+--Merriamosaurus MM03 [incl. Rotundopteryx Maisch & Matzke 2000 MM03, MM00]
                  |    `--M. hulkei (Maisch & Matzke 2000) MM03 [=*Rotundopteryx hulkei MM00]
                  `--Merriamosauria MM00
                       |--Besanosaurus Dal Sasso & Pinna 1996 MM03, MM00
                       |    `--*B. leptorhynchus Dal Sasso & Pinna 1996 MM00
                       `--+--Thalattoarchon Fröbisch, Fröbisch et al. 2013 FF13
                          |    `--*T. saurophagis Fröbisch, Fröbisch et al. 2013 FF13
                          `--+--+--Shonisaurus Camp 1976 FF13, MM00
                             |  |    |--*S. popularis Camp 1976 MM00
                             |  |    `--S. sikanniensis D07
                             |  `--Shastasaurus Merriam 1895 FF13, MM00 [Shastasauria]
                             |       |  i. s.: S. alexandrae Merriam 1902 [incl. S. altispinus, S. osmonti] MM00
                             |       |--S. sikanniensis FF13
                             |       `--+--*S. pacificus Merriam 1895 MM00, FF13, MM00
                             |          `--S. liangae FF13
                             `--+--Mikadocephalus Maisch & Matzke 1997 JH05, MM00
                                |    `--*M. gracilirostris Maisch & Matzke 1997 MM00
                                `--+--Callawayia Maisch & Matzke 2000 MM03, MM00 (see below for synonymy)
                                   |    `--*C. neoscapularis (McGowan 1994) MM00 (see below for synonymy)
                                   `--Parvipelvia MM00
                                        |--Hudsonelpidia McGowan 1995 MM03, MM00
                                        |    `--*H. brevirostris McGowan 1995 MM00
                                        `--Neoichthyosauria MM00
                                             |  i. s.: Nannopterygius von Huene 1922 MM00
                                             |           `--*N. enthekiodon (Hulke 1871) MM00
                                             |--Suevoleviathan Maisch 1998 FF13, MM00 [Suevoleviathanidae]
                                             |    |--*S. disinteger (von Huene 1926) MM00
                                             |    `--S. integer (Bronn 1844) MM00
                                             `--+--+--Temnodontosaurus Lydekker 1889 FF13, MM00 (see below for synonymy)
                                                |  |    |--*T. platyodon (Conybeare 1822) MM00
                                                |  |    |--T. acutirostris (Owen 1840) MM00
                                                |  |    |--T. eurycephalus McGowan 1974 MM00
                                                |  |    |--T. longirostris D07
                                                |  |    |--T. nuertingensis (von Huene 1931) MM00
                                                |  |    `--T. trigonodon (von Theodori 1843) MM00
                                                |  `--Leptonectidae [Eurhinosauria, Eurhinosauridae] MM00
                                                |       |--+--Eurhinosaurus Abel 1909 FF13, MM00
                                                |       |  |    `--*E. longirostris (von Jaeger 1856) MM00
                                                |       |  `--Excalibosaurus McGowan 1986 FF13, MM00
                                                |       |       `--*E. costini McGowan 1986 [=Eurinosaurus costini] MM00
                                                |       `--Leptonectes McGowan 1996 FF13, MM00 (see below for synonymy)
                                                |            |--*L. tenuirostris (Conybeare 1822) MM00
                                                |            |--L. moorei McGowan & Milner 1999 MM00
                                                |            `--L. solei (McGowan 1993) MM00
                                                `--+--Thunnosauria FF13
                                                   `--Macgowania Motani 1999 FF13, MM00
                                                        `--*M. janiceps (McGowan 1996) [=Ichthyosaurus janiceps] MM00

Callawayia Maisch & Matzke 2000 MM03, MM00 [=Metashastasaurus Nicholls & Manabe 2001 H02]

*Callawayia neoscapularis (McGowan 1994) MM00 [=Shastasaurus neoscapularis MM00, *Metashastasaurus neoscapularis H02]

Guizhouichthyosaurus tangae Cao & Lu in Yin et al. 2000 [incl. Cymbospondylus asiaticus Li & You 2002] JH05

Leptonectes McGowan 1996 FF13, MM00 [incl. Leptopterygius von Huene 1922 (preoc.) MM00]

Temnodontosaurus Lydekker 1889 FF13, MM00 [incl. Proteosaurus Home 1814 MM00; Temnodontosauridae]

Toretocnemus Merriam 1903 FF13, MM00 [incl. Leptocheirus Merriam 1903 (preoc.) MM00, Merriamia Boulenger 1904 MM00]

*Type species of generic name indicated

REFERENCES

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

[FF13] Fröbisch, N. B., J. Fröbisch, P. M. Sander, L. Schmitz & O. Rieppel. 2013. Macropredatory ichthyosaur from the Middle Triassic and the origin of modern trophic networks. Proceedings of the National Academy of Sciences of the USA 110 (4): 1393–1397.

[H02] Haubold, H. 2002. Literaturbericht: Ichthyosaurier und Sauropterygier. Zbl. Geol. Paläont. Teil II 5-6: 367–403.

[JH05] Jiang, D.-Y., W.-C. Hao, M. W. Maisch, A. T. Matzke & Y.-L. Sun. 2005. A basal mixosaurid ichthyosaur from the Middle Triassic of China. Palaeontology 48 (4): 869–882.

[MM00] Maisch, M. W., & A. T. Matzke. 2000. The Ichthyosauria. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 298: 1–159.

[MM03] Maisch, M. W., & A. T. Matzke. 2003. Observations on Triassic ichthyosaurs. Part XII. A new Early Triassic ichthyosaur genus from Spitzbergen. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 229 (3): 317–338.

Ichthyosauria

Reconstruction of Grippia longirostris, copyright Dmitry Bogdanov.

Belongs within: Reptilia.
Contains: Hueneosauria.

The Ichthyosauria are a major clade of marine reptiles known from the Lower Triassic to the lower Upper Cretaceous, members of which evolved a remarkably fish-like body form (Maisch & Matzke 2000).

Characters (from Maisch & Matzke 2000): Premaxillary rostrum elongate; naris displaced posteriorly, close to orbit; upper temporal fenestra with very considerable contribution of supratemporal; ectopterygoid absent; thoracal ribs articulating exclusively with centrum; manus and pes exhibiting hyperphalangy; fibula immobile relative to femur.

<==Ichthyosauria [Eoichthyosauria, Ichthyopterygia]
    |--Thaisaurus Mazin, Suteethorn et al. 1991 MM03
    |    `--*T. chonglakmanii Mazin, Suteethorn et al. 1991 MM00
    `--+--Parvinatator Nicholls & Brinkman 1995 FF13, MM03
       |    `--*P. wapitiensis Nicholls & Brickman 1995 MM00
       `--+--Utatsusaurus Shikama, Kamei & Murata 1978 FF13, MM03
          |    `--*U. hataii Shikama, Kamei & Murata 1978 MM00
          `--+--Grippiidae [Grippidia] MM00
             |    |--Chaohusaurus Young & Dong 1972 MM03 (see below for synonymy)
             |    |    `--*C. geishanensis Young & Dong 1972 MM00
             |    `--Grippia Wiman 1929 FF13, MM03
             |         `--*G. longirostris Wiman 1929 MM00
             `--+--Hueneosauria FF13
                `--Quasianosteosaurus Maisch & Matzke 2003 [Quasianosteosauridae] MM03
                     `--*Q. vikinghoegdai Maisch & Matzke 2003 MM03

Ichthyosauria incertae sedis:
  Isfjordosaurus Motani 1999 MM03
    `--*I. minor (Wiman 1910) [=Pessopteryx minor] MM00
  Svalbardosaurus Mazin 1981 (n. d.) MM03
    `--S. crassidens MM03
  Himalayasaurus Young & Dong 1972 (n. d.) MM00
    `--*H. tibetensis Young & Dong 1972 (n. d.) MM00
  Tibetosaurus Dong in Young et al. 1982 (n. d.) MM00
    `--*T. tingjiensis Dong in Young et al. 1982 (n. d.) MM00
  Pessosaurus Wiman 1910 (n. d.) MM00
    |--*P. polaris (Hulke 1873) (n. d.) MM00
    `--P. suevicus von Huene 1916 MM00
  Macropterygius von Huene 1922 (n. d.) MM00
    `--*M. trigonus (Owen 1840) MM00
  Pachygonosaurus von Huene 1916 (n. d.) MM00
    `--*P. robustus Maisch & Matzke 1997 (n. d.) MM00
  Omphalosauridae H02
    |--Tholodus schmidi H02
    `--Omphalosaurus Merriam 1906 D07 [incl. Pessopteryx Wiman 1910 MM00]
         |--O. nettarhynchus D07
         |--O. nevadanus Merriam 1906 MM00
         |--O. nisseri (Wiman 1910) (see below for synonymy) MM00
         `--O. wolfi D07

Chaohusaurus Young & Dong 1972 MM03 [incl. Anhuisaurus Chen 1985 (preoc.) MM00, Chensaurus Mazin, Suteethorn et al. 1991 MM00]

Omphalosaurus nisseri (Wiman 1910) [=*Pessopteryx nisseri; incl. P. arctica Wiman 1910, P. pinguis Wiman 1910] MM00

*Type species of generic name indicated

REFERENCES

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

[FF13] Fröbisch, N. B., J. Fröbisch, P. M. Sander, L. Schmitz & O. Rieppel. 2013. Macropredatory ichthyosaur from the Middle Triassic and the origin of modern trophic networks. Proceedings of the National Academy of Sciences of the USA 110 (4): 1393–1397.

[H02] Haubold, H. 2002. Literaturbericht: Ichthyosaurier und Sauropterygier. Zbl. Geol. Paläont. Teil II 5-6: 367–403.

[MM00] Maisch, M. W., & A. T. Matzke. 2000. The Ichthyosauria. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 298: 1–159.

[MM03] Maisch, M. W., & A. T. Matzke. 2003. Observations on Triassic ichthyosaurs. Part XII. A new Early Triassic ichthyosaur genus from Spitzbergen. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 229 (3): 317–338.

Procolophonia

Reconstruction of Pareiasaurus serridens, copyright Nobu Tamura.

Belongs within: Parareptilia.

The Procolophonia are a clade of reptiles known from the mid-Permian to the late Triassic, including the large armoured Pareiasauria and the lizard-like Procolophonoidea. Some authors have also included the modern turtles in this clade.

Synapomorphies (from Laurin & Reisz 1995): Pineal foramen close to fronto-parietal suture; postorbital far from occiput; tabular absent; anterior maxillary narial foramen present; caniniform region absent; cranio-quadrate space large; pterygoid palatal ramus not reaching level of choana; transverse flange of pterygoid directed anterolaterally; cultriform process short; supraocciptal plate reduced to sagittal pillar; paroccipital process antero-posteriorly expanded; quadrate condyle articular surfaces nearly flat and antero-posteriorly short; foramen intermandibularis posterior located below coronoid process or farther posteriorly; meckelian fossa facing dorsally; fossa meckelii short; surangular not reaching coronoid eminence; prearticular extending no farther than coronoid eminence; retroarticular process transversely broad and dorsally concave; trunk neural arches swollen with narrow zygapophyseal buttresses; transverse processes or caudal ribs present on at least thirteen caudals; interclavicle T-shaped; anteriorly directed groove on interclavicle for clavicle attachment; scapula narrow and flat; glenoid short, bipartite; acetabular buttress large, overhanging strongly above acetabulum; femoral greater trochanter present on posterior edge of femur; astragalus and calcaneum sutured or fused to each other; fifth distal tarsal absent; dorsal dermal ossifications present.

<==Procolophonia
    |  i. s.: Nycteroleter [Nycteroleterida] LR95
    |--Pareiasauria [Serratidentia] MD07
    |    |  i. s.: Pareiasuchus SS02
    |    |         Anthodon pricei [=Nanoparia pricei] SS02
    |    |         Provelosaurus SS02
    |    |         Elginia Newton 1892 D07
    |    |           `--E. mirabilis D07
    |    |         Deltavjatia vjatkensis LR95
    |    |         Parasaurus LR95
    |    |         Shihtienfenia permica LR95, WH78
    |    |         Pareiasauridae TT05
    |    |           |--Proelginia permiana Hartmann-Weinberg 1937 TT05
    |    |           |--Scutosaurus Amalitzky 1922 TT05, D07
    |    |           |    `--S. karpinskii D07
    |    |           `--Pareiasaurus Owen 1876 D07
    |    |                |--P. americanus D07
    |    |                |--P. baini D07
    |    |                |--P. nasicornis D07
    |    |                |--P. omoeratus D07
    |    |                |--P. peringuegi D07
    |    |                `--P. serridens D07
    |    |--Bradysaurus MDS02
    |    `--+--Nochelesaurus MDS02
    |       `--Embrithosaurus MDS02
    `--Procolophonoidea MD07
         |  i. s.: Thelephon contritus MD07
         |--Owenettidae MD07
         |    |  i. s.: Saurodektes rogersorum MD07
         |    |--+--Owenetta rubidgei Broom 1939 MD07
         |    |  `--Barasaurus besairei MD07
         |    `--+--‘Owenetta’ kitchingorum MD07
         |       `--Candelaria barbouri MD07
         `--Procolophonidae [Telerpetidae] MD07
              |  i. s.: Burtensia MDS02
              |         Timanophon raridentatus MDS02, MD07
              |         Acadiella MDS02
              |         Haligonia MDS02
              |         Koiloskiosaurus MDS02
              |         Neoprocolophon MDS02
              |         Eumetabolodon MDS02
              |         Microtheledon MDS02
              |         Myocephalus MDS02
              |         Myognathus LR95
              |         Paoteodon LR95
              |         Sclerosaurus [Sclerosauridae] MD07
              |         Teratophon D-SMS06
              |--Coletta seca Gow 2000 MD07, MDS02
              |--Pintosaurus Piñeiro et al. 2004 D-SMS06
              |    `--*P. magnidentis Piñeiro et al. 2004 D-SMS06
              `--+--Sauropareion anoplus Modesto et al. 2001 MD07
                 `--+--Spondylolestinae MDS02
                    |    |--Contritosaurus simus MD07
                    |    |--Spondylolestes rubidgei MDS02
                    |    `--Phaanthosaurus MDS02
                    `--+--Thelerpeton oppressus MD07
                       |--+--Thelegnathus spinigenis MD07, MDS02
                       |  `--Leptopleuroninae MD07
                       |       |  i. s.: Libognathus sheddi PSH17
                       |       |--Tichvinskia vjatkensis MD07
                       |       `--+--Scoloparia glyphanodon MD07
                       |          `--+--Leptopleuron lacertinum MD07
                       |             `--Hypsognathus Gilmore 1928 MD07, D07
                       |                  `--H. fenneri MD07
                       `--Procolophoninae SS02
                            |--Orenburgia SS02
                            |--Samaria SS02
                            |--Lestanshoria SS02
                            |--Anomoiodon von Huene 1939 SS02
                            |    `--A. liliensterni von Huene 1939 SS02
                            |--Procolophon Owen 1876 MD07, D-SMS06
                            |    |--P. brasiliensis Cisneros & Schultz 2002 D-SMS06
                            |    |--P. pricei Lavina 1983 D-SMS06
                            |    `--P. trigoniceps Owen 1876 MD07
                            `--Kapes Ivakhnenko 1975 SS02
                                 |--*K. amaenus Ivakhnenko 1975 SS02
                                 |--K. bentoni Spencer & Storrs 2002 SS02
                                 |--K. komiensis (Ivakhnenko 1975) [=Macrophon komiensis] SS02
                                 `--K. majmesculae (Ochev 1968) (see below for synonymy) SS02

Kapes majmesculae (Ochev 1968) [=Tichvinskia majmesculae; incl. K. serotinus Novikov 1991] SS02

*Type species of generic name indicated

REFERENCES

[D-SMS06] Dias-da-Silva, S., S. P. Modesto & C. L. Schultz. 2006. New material of Procolophon (Parareptilia: Procolophonoidea) from the Lower Triassic of Brazil, with remarks on the ages of the Sanga do Cabral and Buena Vista formations of South America. Canadian Journal of Earth Sciences 43: 1685–1693.

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

[LR95] Laurin, M., & R. R. Reisz. 1995. A reevaluation of early amniote phylogeny. Zoological Journal of the Linnean Society 113: 165–223.

[MD07] Modesto, S. P., & R. Damiani. 2007. The procolophonoid reptile Sauropareion anoplus from the lowermost Triassic of South Africa. Journal of Vertebrate Paleontology 27 (2): 337–349.

[MDS02] Modesto, S. P., R. J. Damiani & H.-D. Sues. 2002. A reappraisal of Coletta seca, a basal procolophonid reptile from the Lower Triassic of South Africa. Palaeontology 45 (5): 883–895.

[PSH17] Pardo, J. D., B. J. Small & A. K. Huttenlocker. 2017. Stem caecilian from the Triassic of Colorado sheds light on the origins of Lissamphibia. Proceedings of the National Academy of Sciences of the USA 114 (27): E5389–E5395.

[SS02] Spencer, P. S., & G. W. Storrs. 2002. A re-evaluation of small tetrapods from the Middle Triassic Otter Sandstone Formation of Devon, England. Palaeontology 45 (3): 447–467.

[TT05] Tverdokhlebov, V. P., G. I. Tverdokhlebova, A. V. Minikh, M. V. Surkov & M. J. Benton. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69: 27–77.

[WH78] Wang Z. & Huang R.-J. 1978. Triassic charophytes of Shaanxi. Acta Palaeontologica Sinica 17 (3): 267–276.

Trematosauria

Reconstruction of Metoposaurus krasiejowensis, copyright Dmitry Bogdanov.

Belongs within: Stereospondylomorpha.

The Trematosauria are a clade of temnospondyls defined by Yates & Warren (2000) as including all species closer to Trematosaurus than Parotosuchus.

Synapomorphies (from Yates & Warren 2000): Interorbital space greater than 50% of skull width at mid orbital level; stapedial foramen absent; sharp rimmed depressions absent at posterior end of ventral surface of parasphenoid plate; denticle patches absent from all three coronoids; chorda tympanic foramen located on preaticular; hamate process absent.

<==Trematosauria [Trematosauroidei] YW00
    |  i. s.: Lyrocephalus euri CZ79
    |--+--Luzocephalus YW00
    |  `--Trematosauroidea YW00
    |       |--Thoosuchus jakovlevi YW00
    |       `--Lyrocephaliscus [Trematosauridae] YW00
    |            `--L. euri YW00
    `--+--Metoposauroidea YW00
       |    |--Almasaurus habbazi Dutuit 1972 YW00, CZ79
       |    |--Buettneria perfecta YW00
       |    |--Apachesaurus PI05
       |    `--Metoposaurus Lydekker 1890 D07
       |         |--M. bakeri D07
       |         |--M. diagnosticus YW00
       |         |--M. heimi D07
       |         |--M. krasiejowensis D07
       |         |--M. maleriensis DDL04
       |         |--M. santaecrucis D07
       |         `--M. stuttgartiensis D07
       `--+--Plagiosauroidea YW00
          |    |--Laidleria [Laidleriidae] YW00
          |    |    `--L. gracilis Kitching 1958 CZ79
          |    `--Plagiosauridae YW00
          |         |--Plagiosternum danilovi YW00
          |         `--Plagiosaurus YW00
          `--+--Brachyopoidea YW00
             |    |--Keratobrachyops australis YW00
             |    `--+--Brachyopidae YW00
             |       |    |--Xenobrachyops allos YW00
             |       |    |--Brachyops laticeps S87
             |       |    `--Batrachosuchus YW00
             |       |         `--‘Blinasaurus’ henwoodi YW00
             |       `--Chigutisauridae YW00
             |            |--Siderops kehli YW00
             |            `--Pelorocephalus Cabera 1944 YW00, D07 [incl. Chigutisaurus YW00]
             |                 |--P. cacheutensis D07
             |                 |--P. ischigualastensis D07
             |                 |--P. mendozensis D07
             |                 `--P. tenax D07
             `--Rhytidosteoidea CZ79
                  |--Indobrachyopidae CZ79
                  |    |--Derwentia warreni Cosgriff 1974 CZ79
                  |    `--+--Indobrachyops panchetensis Van Huene & Sahni 1958 CZ79
                  |       `--+--Rewana quadricuneata Howie 1972 CZ79
                  |          `--Mahavisaurus CZ79
                  |               |--M. australis (Lehmann 1966) [=Lyrosaurus australis] CZ79
                  |               `--M. dentatus Lehmann 1966 CZ79
                  `--Rhytidosteidae YW00
                       |  i. s.: Arcadia myriadens YW00
                       |         Trucheosaurus YW00
                       |--Deltasaurus CZ79
                       |    |--D. kimberleyensis Cosgriff 1965 CZ79
                       |    `--D. pustulatus Cosgriff 1965 CZ79
                       `--+--Pneumatostega Cosgriff & Zawiskie 1979 CZ79
                          |    `--*P. potamia Cosgriff & Zawiskie 1979 CZ79
                          `--+--Rhytidosteus capensis Owen 1884 CZ79
                             `--Peltostega CZ79
                                  |--P. erici Wiman 1916 CZ79
                                  `--P. wimani Nilsson 1946 CZ79

*Type species of generic name indicated

REFERENCES

[CZ79] Cosgriff, J. W., & J. M. Zawiskie. 1979. A new species of the Rhytidosteidae from the Lystrosaurus Zone and a review of the Rhytidosteoidea. Palaeontologia Africana 22: 1–27.

[DDL04] Datta, P. M., D. P. Das & Z.-X. Luo. 2004. A Late Triassic dromatheriid (Synapsida: Cynodontia) from India. Annals of Carnegie Museum 73 (2): 12–24.

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

[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.

[S87] Stephens, W. J. 1887. On the Biloela labyrinthodont (second notice). Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1113–1121, pl. 14.

[YW00] Yates, A. M., & A. A. Warren. 2000. The phylogeny of the ‘higher’ temnospondyls (Vertebrata: Choanata) and its implications for the monophyly and origins of the Stereospondyli. Zoological Journal of the Linnean Society 128: 77–121.

Parareptilia

Reconstruction of Milleretta rubidgei, copyright Nobu Tamura.

Belongs within: Reptilia.
Contains: Procolophonia.

The Parareptilia are a clade corresponding to the greater part of the previously recognised 'Anapsida' group of reptiles lacking a temporal fenestra in the skull. They are known from the Upper Permian and Triassic; their survival after this time depends on whether the modern turtles belong to this lineage.

<==Parareptilia MA04
    |--+--+--Chalcosaurus TT05 [Lanthanosuchida, Lanthanosuchidae, Lanthanosuchoidea MD07]
    |  |  |    `--C. lukjanovae (Ivakhnenko 1980) TT05
    |  |  `--Acleistorhinus Daly 1969 SS15, D07
    |  |       `--A. pteroticus D07
    |  `--+--Macroleter poezicus MD07
    |     `--Procolophonia MD07
    `--Millerettidae [Millerosauroidea] MD07
         |--Milleropsis LR95
         |--Millerosaurus LR95
         |--Heleophilus LR95
         `--Milleretta Watson 1957 D07
              `--M. rubidgei M06

*Type species of generic name indicated

REFERENCES

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

[LR95] Laurin, M., & R. R. Reisz. 1995. A reevaluation of early amniote phylogeny. Zoological Journal of the Linnean Society 113: 165–223.

[M06] Modesto, S. P. 2006. The cranial skeleton of the Early Permian aquatic reptile Mesosaurus tenuidens: implications for relationships and palaeobiology. Zoological Journal of the Linnean Society 146: 345–368.

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

[MD07] Modesto, S. P., & R. Damiani. 2007. The procolophonoid reptile Sauropareion anoplus from the lowermost Triassic of South Africa. Journal of Vertebrate Paleontology 27 (2): 337–349.

[SS15] Schoch, R. R., & H.-D. Sues. 2015. A Middle Triassic stem-turtle and the evolution of the turtle body plan. Nature 523: 584–587.

[TT05] Tverdokhlebov, V. P., G. I. Tverdokhlebova, A. V. Minikh, M. V. Surkov & M. J. Benton. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69: 27–77.

Thunnosauria

Reconstruction of Platypterygius bannovkensis, by Olorotitan.

Belongs within: Hueneosauria.

The clade Thunnosauria, defined by Maisch & Matzke (2000) as the most exclusive clade including Ichthyosaurus communis and Stenopterygius quadriscissus, includes the latest-surviving ichthyosaurs. Thunnosaurs had a compact body form relative to other ichthyosaurs with a short, powerful tail that would have provided most of the propulsive force when swimming.

See also: The tuna-lizards.

Synapomorphies (from Maisch & Matzke 2000): Quadratojugal small; squamosal small, delicate plate or absent; rib articulation in thorax bicipital; tail short; hindlimbs much shorter than forelimbs.

<==Thunnosauria [Ichthyosauridae]
    |--Stenopterygiidae MM00
    |    |--Chacaicosaurus Fernández 1994 MM00
    |    |    `--*C. cayi Fernández 1994 MM00
    |    `--Stenopterygius Jaekel 1904 FF13, MM000
    |         |--*S. quadricissus (Quenstedt 1856) MM00
    |         |--S. hauffianus von Huene 1922 MM00
    |         |--S. longifrons (Owen 1881) MM00
    |         `--S. megalorhinus von Huene 1922 MM00
    `--+--Ichthyosaurus De La Beche & Conybeare 1821 FF13, MM00 (see below for synonymy)
       |    |--*I. communis Conybeare 1822 MM00
       |    |--I. acutirostris N85
       |    |--I. australis F71
       |    |--I. breviceps Owen 1881 MM00
       |    |--I. conybeari Lydekker 1888 MM00
       |    |--I. intermedius Conybeare 1822 MM00
       |    |--I. marathonensis F71
       |    |--‘Protoichthyosaurus’ porostealis Appleby 1979 MM00
       |    `--‘*Protoichthyosaurus’ postaxalis Appleby 1979 MM00
       `--+--Aegirosaurus Bardet & Fernández 2000 MM03, MM00
          |    |--*A. leptospondylus (Wagner 1873) [=Ichthyosaurus leptospondylus] MM00
          |    `--‘Ophthalmosaurus’ monocharactus Appleby 1956 MM00
          `--Ophthalmosauridae [Ophthalmosauria] MM00
               |--Brachypterygius von Huene 1922 FF13, MM00 (see below for synonymy)
               |    |--*B. extremus (Boulenger 1904) MM00
               |    |--‘Otschevia’ alekseevi Arkhangelsky 2001 H02
               |    |--B. mordax (McGowan 1976) [=Grendelius mordax] MM00
               |    `--B. pseudoscythius (Efimov 1998) MM00 (see below for synonymy)
               `--+--+--Platypterygius von Huene 1922 FF13, MM00 (see below for synonymy)
                  |  |    |--*P. platydactylus (Broili 1907) MM00
                  |  |    |--P. americanus (Nace 1939) MM00
                  |  |    |--P. bannovkensis Arkhangelsky 1998 [=P. (Pervushovisaurus) bannovkensis] MM00
                  |  |    |--P. bedengensis (Efimov 1998) [=*Plutoniosaurus bedengensis] MM00
                  |  |    |--P. birjukovi (Otschev & Efimov 1985) [=*Simbirskiasaurus birjukovi] MM00
                  |  |    |--P. hauthali (von Huene 1926) MM00
                  |  |    |--P. hercynicus Kuhn 1946 MM00
                  |  |    |--P. kiprijanoffi Romer 1968 MM00
                  |  |    `--P. longmani Wade 1990 MM00
                  |  `--Caypullisaurus Fernández 1997 FF13, MM00
                  |       `--*C. bonapartei Fernández 1997 MM00
                  `--Ophthalmosaurus Seeley 1874 FF13, MM00 (see below for synonymy)
                       |--*O. icenicus Seeley 1874 MM00
                       |--‘*Khudiakovia’ calloviensis Arkhangelsky 1999 (n. d.) MM00
                       |--O. discus D07
                       |--O. gorodischensis (Efimov 1999) (see below for synonymy) MM00
                       |--O. natans (Marsh 1879) MM00
                       |--O. periallus (Fernández 1999) [=Mollesaurus periallus] MM00
                       |--‘Paraophthalmosaurus’ saratoviensis MM00
                       |--O. saveljeviensis (Arkhangelsky 1997) [=*Paraophthalmosaurus saveljeviensis] MM00
                       `--O. yasykovi (Efimov 1999) [=*Yasykovia yasykovi; incl. Y. kabanovi, Y. mittai, Y. sumini] MM00

Brachypterygius von Huene 1922 FF13, MM00 [incl. Grendelius McGowan 1976 MM00, Otschevia Efimov 1998 MM00]

Brachypterygius pseudoscythius (Efimov 1998) MM00 [=*Otschevia pseudoscythia MM00; incl. B. zhuravlevi Arkhangelsky 1998 MM00, O. zhuralevi H02]

Ichthyosaurus De La Beche & Conybeare 1821 FF13, MM00 [incl. Eurypterygius Jaekel 1904 MM00, Protoichthyosaurus Appleby 1979 MM00]

Ophthalmosaurus Seeley 1874 FF13, MM00 [incl. Ancanamunia Rusconi 1942 MM00, Apatodontosaurus Mehl 1927 MM00, Baptanodon Marsh 1880 MM00, Khudiakovia Arkhangelsky 1999 MM00, Mollesaurus Fernández 1999 MM00, Paraophthalmosaurus Arkhangelsky 1997 MM00, Sauranodon Marsh 1879 (preoc.) MM00, Undorosaurus Efimov 1999 MM00, Yasykovia Efimov 1999 MM00]

Ophthalmosaurus gorodischensis (Efimov 1999) [=*Undorosaurus gorodischensis; incl. U. khorlovensis, U. nessovi] MM00

Platypterygius von Huene 1922 FF13, MM00 [incl. Longirostria Arkhangelsky 1998 MM00, Myobradypterygius von Huene 1926 MM00, Myopterygius von Huene 1922 MM00, Pervushovisaurus Arkhangelsky 1998 MM00, Plutoniosaurus Efimov 1997 MM00, Simbirskiasaurus Ochev & Efimov 1985 MM00, Tenuirostria Arkhangelsky 1998 MM00]

*Type species of generic name indicated

REFERENCES

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

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

[FF13] Fröbisch, N. B., J. Fröbisch, P. M. Sander, L. Schmitz & O. Rieppel. 2013. Macropredatory ichthyosaur from the Middle Triassic and the origin of modern trophic networks. Proceedings of the National Academy of Sciences of the USA 110 (4): 1393–1397.

[H02] Haubold, H. 2002. Literaturbericht: Ichthyosaurier und Sauropterygier. Zbl. Geol. Paläont. Teil II 5-6: 367–403.

[MM00] Maisch, M. W., & A. T. Matzke. 2000. The Ichthyosauria. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 298: 1–159.

[MM03] Maisch, M. W., & A. T. Matzke. 2003. Observations on Triassic ichthyosaurs. Part XII. A new Early Triassic ichthyosaur genus from Spitzbergen. Neues Jahrbuch für Geologie und Paläontologie—Abhandlungen 229 (3): 317–338.

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

Last updated: 14 July 2018.

Stereospondylomorpha

Restoration of Benthosuchus sushkini, from here.

Belongs within: Temnospondyli.
Contains: Trematosauria.

The Stereospondylomorpha are a group of mostly aquatic temnospondyls known from the early Permian to early Cretaceous, though their diversity was much reduced after the end of the Triassic. They include the Permian Archegosauroidea, species of which were superficially crocodile-like with long narrow snouts.

Synapomorphies (from Yates & Warren 2000): Jugal extending anterior to orbit; prefrontal-jugal suture present; maxilla-nasal suture present; arcadian groove present on post glenoid area of mandible.

<==Stereospondylomorpha YW00
    |--Archegosauroidea YW00
    |    |  i. s.: Bageherpeton Dias & Barberena 2001 D07
    |    |           `--B. longignathus D07
    |    |--Actinodontidae YW00
    |    |    |--Cheliderpeton latirostre YW00
    |    |    `--Sclerocephalus [incl. Actinodon] YW00
    |    |         `--S. haeuseri YW00
    |    `--Archegosauridae YW00
    |         |--Melosauridae TT05
    |         |    |--Konzhukovia vetusta (Konzhukova 1955) YW00, TT05
    |         |    |--Tryphosuchus kinelensis (V’yushkov 1955) TT05
    |         |    `--Uralosuchus tverdochlebovae Gubin 1993 TT05
    |         `--+--Archegosaurus decheni YW00
    |            `--Platyoposaurus YW00
    |                 |--P. rickardi (Twelvetrees 1880) TT05
    |                 |--P. stuckenbergi YW00
    |                 `--P. vjuschkovi Gubin 1989 TT05
    `--Stereospondyli YW00
         |  i. s.: Aphaneramma S02
         |         Dutuitosaurus ouazzoui S02
         |         Kupferzellia S02
         |--Peltobatrachus Panchen 1959 YW00, D07
         |    `--P. pustulatus YW00
         `--+--Lapillopsis nana YW00
            `--+--Rhinesuchidae YW00
               |    |--Rhinesuchus PS88
               |    |--Rhineceps YW00
               |    `--Uranocentrodon [Uranocentrodontidae] YW00
               |         `--U. senekalensis (Van Hoepen 1911) CZ79
               `--+--Trematosauria YW00
                  `--Capitosauria [Capitosauroidei] YW00
                       |--+--Chomatobatrachus YW00
                       |  `--Lydekkerinidae CZ79
                       |       |--Limnoiketes paludinatans Parrington 1948 CZ79
                       |       |--Cryobatrachus kitchingi CZ79
                       |       `--Lydekkerina YW00
                       |            |--L. huxleyi (Lydekker 1889) (see below for synonymy) CZ79
                       |            `--L. kitchingi Broom 1950 CZ79
                       `--Capitosauroidea YW00
                            |--Mastodonsauridae
                            |    |--Heptasaurus cappelensis (Wepfer 1923) S02
                            |    `--Mastodonsaurus Jaeger 1828 YW00, D07
                            |         |--M. cappelensis D07
                            |         |--M. giganteus S02
                            |         |--M. lavisi SS02
                            |         |--M. robustus F71 [=Capitosaurus robustus S87]
                            |         `--M. torvus D07
                            `--Capitosauridae YW00
                                 |--Paracyclotosaurus davidi Watson 1958 YW00, F71
                                 |--Kestrosaurus dreyeri Haughton 1925 CZ79
                                 |--Parotosaurus peabodyi CZ79
                                 |--Parotosuchus YW00
                                 |    |--P. aliciae YW00
                                 |    `--P. gunganj YW00
                                 `--Benthosuchus CZ79
                                      |--B. korobkovi S02
                                      |--B. sushkini YW00
                                      `--B. uralensis S02

Inorganic: Mastodonsaurus minirotundomaculosus Okamura 1987 O87
           Benthosuchus sushkini minilorientalus Okamura 1987 O87

Lydekkerina huxleyi (Lydekker 1889) [incl. L. dutoiti Broom 1930, Broomulus dutoiti, L. putterilli Broom 1930, Putterillia platyceps Broom 1930] CZ79

*Type species of generic name indicated

REFERENCES

[CZ79] Cosgriff, J. W., & J. M. Zawiskie. 1979. A new species of the Rhytidosteidae from the Lystrosaurus Zone and a review of the Rhytidosteoidea. Palaeontologia Africana 22: 1–27.

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

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

[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.

[PS88] Panchen, A. L., & T. R. Smithson. 1988. The relationships of the earliest tetrapods. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 1–32. Clarendon Press: Oxford.

[S02] Schoch, R. R. 2002. The neurocranium of the stereospondyl Mastodonsaurus giganteus. Palaeontology 45 (3): 627–645.

[SS02] Spencer, P. S., & G. W. Storrs. 2002. A re-evaluation of small tetrapods from the Middle Triassic Otter Sandstone Formation of Devon, England. Palaeontology 45 (3): 447–467.

[S87] Stephens, W. J. 1887. On the Biloela labyrinthodont (second notice). Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1113–1121, pl. 14.

[TT05] Tverdokhlebov, V. P., G. I. Tverdokhlebova, A. V. Minikh, M. V. Surkov & M. J. Benton. 2005. Upper Permian vertebrates and their sedimentological context in the South Urals, Russia. Earth-Science Reviews 69: 27–77.

[YW00] Yates, A. M., & A. A. Warren. 2000. The phylogeny of the ‘higher’ temnospondyls (Vertebrata: Choanata) and its implications for the monophyly and origins of the Stereospondyli. Zoological Journal of the Linnean Society 128: 77–121.

Last updated: 3 August 2017.