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.

Arthroleptinae

Female Arthroleptis reichei, copyright Vincenzo Mercurio.

Belongs within: Laurentobatrachia.

The Arthroleptinae are a group of frogs found in sub-Saharan Africa, males of which often have hyperelongate third fingers.

<==Arthroleptinae [Arthroleptina, Arthroleptini] FG06
    |--Coracodichus FG06
    |--Cardioglossa Boulenger 1900 FG06
    |    |--+--C. gracilis PW11
    |    |  `--C. schioetzi PW11
    |    `--+--C. elegans PW11
    |       `--+--C. gratiosa PW11
    |          `--+--+--C. leucomystax PW11
    |             |  `--C. occidentalis PW11
    |             `--+--C. pulchra PW11
    |                `--+--C. manengouba PW11
    |                   `--C. oreas PW11
    `--Arthroleptis Smith 1849 [incl. Schoutedenella De Witte 1921] FG06
         |  i. s.: A. batesi FG06
         |         A. crusculum Angel 1950 [=Schoutedenella crusculum] FG06
         |         A. discodactyla (Laurent 1954) [=Schoutedenella discodactyla] FG06
         |         A. hematogaster (Laurent 1954) [=Schoutedenella hematogaster] FG06
         |         A. lameerei de Witte 1921 [=Schoutedenella lameerei] FG06
         |         A. loveridgei de Witte 1933 [=Schoutedenella loveridgei] FG06
         |         A. milletihorsini Angel 1922 [=Schoutedenella milletihorsini] FG06
         |         A. mossoensis (Laurent 1954) [=Schoutedenella mossoensis] FG06
         |         A. nimbaensis Angel 1950 [=Schoutedenella nimbaensis] FG06
         |         A. phrynoides (Laurent 1976) [=Schoutedenella phrynoides] FG06
         |         A. pyrrhoscelis Laurent 1952 [=Schoutedenella pyrrhoscelis] FG06
         |         A. spinalis Boulenger 1919 FG06
         |         A. troglodytes Poynton 1963 [=Schoutedenella troglodytes] FG06
         |         A. vercammeni (Laurent 1954) [=Schoutedella vercammeni] FG06
         |         A. xenochirus Boulenger 1905 [=Schoutedenella xenochirus] FG06
         |         A. zimmeri Ahl 1925 FG06
         |--+--A. xenodactylus Boulenger 1909 PW11, FG06 [=Schoutedenella senodactyla FG06]
         |  `--+--A. schubotzi Nieden 1911 PW11, FG06 [=Schoutedenella schubotzi FG06]
         |     `--A. xenodactyloides Hewitt 1933 PW11, FG06 [=Schoutedenella xenodactyloides FG06]
         `--+--+--A. taeniatus PW11 [=Schoutedenella taeniata FG06]
            |  `--+--A. aureoli PW11
            |     `--A. sylvaticus (Laurent 1954) PW11, FG06 [=Schoutedenella sylvatica FG06]
            `--+--+--A. francei PW11
               |  `--A. wahlbergii PW11
               `--+--+--A. tanneri PW11
                  |  `--+--A. reichei PW11
                  |     `--+--A. affinis PW11
                  |        `--A. nikeae PW11
                  `--+--A. stenodactylus PW11
                     `--+--+--A. adelphus PW11
                        |  `--A. poecilonotus PW11
                        `--+--A. krokosua PW11
                           `--A. variabilis PW11

*Type species of generic name indicated

REFERENCES

[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.

[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.

Laurentobatrachia

Peacock tree frog Leptopelis vermiculatus, copyright Dawson.

Belongs within: Ranoidea.
Contains: Arthroleptinae, Hyperolius, Heterixalus.

The Laurentobatrachia are a clade of frogs found in sub-Saharan Africa and Madagascar.

Characters (from Frost et al. 2006): Larval m. levator manidbulae externus in two portion; commissura proximalis III absent; intercalary elements absent; external metatarsal tubercle poorly developed or absent; terminal phalanges claw-shaped.

Laurentobatrachia [Arthroleptoidae]
    |--Arthroleptidae FG06
    |    |--Arthroleptinae FG06
    |    `--+--Leptopelis Günther 1859 FG06 [Leptopelinae PW11, Leptopelini]
    |       |    |  i. s.: L. mossambicus FG06
    |       |    |--+--L. argenteus FG06
    |       |    |  `--L. brevirostris PW11
    |       |    `--+--L. modestus PW11
    |       |       `--+--L. vermiculatus PW11
    |       |          `--+--L. natalensis PW11
    |       |             `--+--+--L. bocagii PW11
    |       |                |  `--L. concolor PW11
    |       |                `--+--L. kivuensis PW11
    |       |                   `--L. palmatus PW11
    |       `--Astylosterninae [Astylosternidae, Astylosternini] PW11
    |            |--Leptodactylodon Andersson 1903 FG06
    |            |    `--L. bicolor FG06
    |            `--+--+--Nyctibates Boulenger 1904 FG06
    |               |  |    `--N. corrugatus FG06
    |               |  `--Scotobleps Boulenger 1900 PW11, FG06
    |               |       `--S. gabonicus FG06
    |               `--+--Trichobatrachus Boulenger 1900 FG06
    |                  |    `--T. robustus FG06
    |                  `--Astylosternus Werner 1898 FG06
    |                       |--A. diadematus PW11
    |                       `--+--A. batesi PW11
    |                          `--A. schioetzi PW11
    `--Hyperoliidae [Hyperoliinae, Kassininae, Kassinini]
         |  i. s.: Chlorolius Perret 1988 FG06
         |         Arlequinus Perret 1988 FG06
         |         Chrysobatrachus Laurent 1951 FG06
         |--Cryptothylax Laurent & Combaz 1950 PW11, FG06
         |    `--C. greshoffii PW11
         `--+--+--Phlyctimantis Laurent & Combaz 1950 PW11, FG06
            |  |    |--P. leonardi PW11
            |  |    `--+--‘Kassina’ maculata PW11
            |  |       `--P. verrucosus PW11
            |  `--+--Kassina Girard 1853 PW11, FG06
            |     |    |--K. senegalensis FG06
            |     |    `--K. wealii WKW73
            |     `--+--Paracassina Peracca 1907 FG06
            |        `--Semnodactylus Hoffman 1939 FG06
            |             `--S. wealii PW11
            `--+--+--Callixalus Laurent 1950 FG06
               |  `--Acanthixalus Laurent 1944 PW11, FG06
               |       |--A. sonjae PW11
               |       `--A. spinosus FG06
               `--+--Opisthothylax Perret 1966 PW11, FG06
                  |    `--O. immaculatus FG06
                  `--+--Hyperolius PW11
                     `--+--Morerella cyanophthalma PW11
                        `--+--+--Heterixalus FG06
                           |  `--Tachycnemis Fitzinger 1843 PW11, FG06 [Tachycneminae]
                           |       `--T. seychellensis NW07
                           `--+--Kassinula Laurent 1940 FG06
                              `--Afrixalus Laurent 1944 PW11, FG06
                                   |  i. s.: A. pygmaeus FG06
                                   |         A. weidholzi KF69
                                   |--+--A. knysnae PW11
                                   |  `--+--A. delicatus PW11
                                   |     `--A. stuhlmanni PW11
                                   `--+--A. fornasini PW11
                                      `--+--A. laevis PW11
                                         `--+--A. dorsalis PW11
                                            `--A. paradorsalis PW11

*Type species of generic name indicated

REFERENCES

[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.

[KF69] Kluge, A. G., & J. S. Farris. 1969. Quantitative phyletics and the evolution of anurans. Systematic Zoology 18 (1): 1–32.

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