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.

[NW07] Nussbaum, R. A., & S.-H. Wu. 2007. Morphological assessments and phylogenetic relationships of the Seychellean frogs of the family Sooglossidae (Amphibia: Anura). Zoological Studies 46 (3): 322–335.

[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.

[WKW73] Wallace, D. G., M.-C. King & A. C. Wilson. 1973. Albumin differences among ranid frogs: taxonomic and phylogenetic implications. Systematic Zoology 22 (1): 1–13.

Limnodynastinae

Spawning striped marsh frogs Limnodynastes peronii, copyright Froggydarb.

Belongs within: Neobatrachia.

The Limnodynastinae are a group of small to medium-sized toad-like terrestrial frogs found in Australia and New Guinea. They have an inguinal amplexus and many are foam-nesters (Frost et al. 2006).

<==Limnodynastinae [Limnodynastidae, Limnodynastini]
    |  i. s.: Opisthodon Steindachner 1867 FG06
    |           |--O. ornatus (Gray 1842) (see below for synonymy) FG06
    |           `--O. spenceri (Parker 1940) [=Limnodynastes spenceri] FG06
    |--+--Notaden Günther 1873 PW11, FG06
    |  |    |--N. bennettii PW11
    |  |    |--N. melanoscaphus FG06
    |  |    |--N. nichollsi TW05
    |  |    `--N. weigeli Shea & Johnston 1988 D11
    |  `--Neobatrachus Peters 1863 PW11, FG06 [incl. Neoruinosus Wells & Wellington 1985 R10]
    |       |  i. s.: N. albipes Roberts, Mahony et al. 1991 R10
    |       |         N. aquilonius Tyler, Davies & Martin 1981 R10
    |       |         N. fulvus Mahony & Roberts 1986 R10
    |       |         N. kunapalari Mahony & Roberts 1986 R10
    |       |         N. sutor Main 1957 R10
    |       |         N. wilsmorei (Parker 1940) R10
    |       |--N. pelobatoides (Werner 1914) PW11, R10
    |       `--+--N. pictus Peters 1863 PW11, R10
    |          `--N. sudelli (Lamb 1911) PW11, R10 [=*Neoruinosus sudelli R10; incl. N. centralis (Parker 1940) R10]
    `--+--Platyplectrum Günther 1863 PW11, DAP09
       |    |--P. spenceri PW11
       |    `--+--P. ornatum PW11
       |       `--Lechriodus Boulenger 1882 PW11, FG06
       |            |--L. fletcheri FG06
       |            `--L. melanopyga FG06
       `--+--Heleioporus Gray 1841 PW11, FG06
          |    |--H. albopunctatus ERK07
          |    |--H. australiacus FG06
          |    |--H. barycragus KF69
          |    |--H. eyrei KF69
          |    |--H. inornatus KF69
          |    `--H. psammophilus ERK07
          `--+--Adelotus Ogilby 1907 PW11, FG06
             |    `--A. brevis FG06
             `--+--Philoria Spencer 1901 PW11, FG06 [incl. Kyarranus Moore 1958 FG06]
                |    `--P. sphagnicola FG06
                `--Limnodynastes Fitzinger 1843 PW11, FG06 [incl. Megistolotis Tyler, Martin & Davis 1979 FG06]
                     |--+--L. salmini PW11
                     |  `--+--L. convexiusculus PW11
                     |     `--L. lignarius PW11
                     `--+--+--+--L. depressus PW11
                        |  |  `--L. fletcheri PW11
                        |  `--+--L. peronii PW11
                        |     `--L. tasmaniensis PW11
                        `--+--L. dorsalis (Gray 1841) PW11, F13 (see below for synonymy)
                           |--L. terraereginae PW11
                           `--+--L. dumerilii Peters 1863 PW11, F13 (see below for synonymy)
                              `--L. interioris Fry 1913 PW11, F13 [=L. dorsalis var. interioris F13]

Limnodynastes dorsalis (Gray 1841) PW11, F13 [=Cystignathus dorsalis F13, Wagleria dorsalis F13]

Limnodynastes dumerilii Peters 1863 PW11, F13 [=L. (Platyplectron) dumerilii F13, L. dorsalis var. dumerilii F13; incl. L. bibronii Krefft 1867 (n. n.) F13, Heliorana grayi Steindachner 1867 F13, Platyplectrum superciliare Keferstein 1867 F13, H. superciliaris F13]

Opisthodon ornatus (Gray 1842) [=Discoglossus ornatus, Limnodynastes ornatus; incl. *Opisthodon frauenfeldi Steindachner 1867] FG06

*Type species of generic name indicated

REFERENCES

[D11] Doughty, P. 2011. An emerging frog diversity hotspot in the northwest Kimberley of Western Australia: another new frog species from the high rainfall zone. Records of the Western Australian Museum 26 (2): 209–216.

[DAP09] Doughty, P., M. Anstis & L. C. Price. 2009. A new species of Crinia (Anura: Myobatrachidae) from the high rainfall zone of the northwest Kimberley, Western Australia. Records of the Western Australian Museum 25 (2): 127–144.

[ERK07] Edwards, D. L., J. D. Roberts & J. S. Keogh. 2007. Impact of Plio-Pleistocene arid cycling on the population history of a southwestern Australian frog. Molecular Ecology 16: 2782–2796.

[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.

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

[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.

[R10] Roberts, J. D. 2010. Taxonomic status of the Australian burrowing frogs Neobatrachus sudelli, N. centralis and Neoruinosus and clarification of the type specimen of N. albipes. Records of the Western Australian Museum 25 (4): 455–458.

[TW05] Thompson, G. G., P. C. Withers, K. A. McMaster & V. A. Cartledge. 2005. Burrows of desert-adapted frogs, Neobatrachus aquilonius and Notaden nichollsi. Journal of the Royal Society of Western Australia 88 (1): 17–23.

Neobatrachia

Table Mountain ghost frog Heleophryne rosei, copyright Vincent Carruthers.

Belongs within: Anura.
Contains: Ranoidea, Hyloidea, Limnodynastinae, Myobatrachinae.

The Neobatrachia are a clade containing the majority of living frogs, united by features such as the presence of a neopalatine bone in the skull.

Synapomorphies (from Frost et al. 2006): Neopalatine bone present; third distal carpal fused to other carpals; m. sartorius completely separated from m. semitendinosus; accessory head present on m. adductor longus; parahyoid bone absent.

<==Neobatrachia [Hemiphractinae]
    |  i. s.: Indobatrachus pusillus (Owen 1847) BB03
    |--Heleophrynidae [Heleophryninae] PW11
    |    |--Hadromophryne natalensis PW11
    |    `--Heleophryne Sclater 1898 FG06
    |         |--H. natalensis FG06
    |         |--H. purcelli BB03
    |         |--H. regis FG06
    |         `--H. rosei FG06
    `--Phthanobatrachia (see below for synonymy) FG06
         |--+--Nasikabatrachus Biju & Bossuyt 2003 FG06 [Nasikabatrachidae PW11]
         |  |    `--*N. sahyadrensis Biju & Bossuyt 2003 BB03
         |  `--Sooglossidae [Sooglossinae] PW11
         |       |--Sooglossus Boylenger 1906 [incl. Nesomantis Boulenger 1909] NW07
         |       |    |--*S. sechellensis (Boettger 1896) [=Arthroleptis sechellensis] NW07
         |       |    `--S. thomasseti (Boulenger 1909) [=*Nesomantis thomasseti] NW07
         |       `--Sechellophryne Nussbaum & Wu 2007 NW07
         |            |--*S. gardineri (Boulenger 1911) [=Nectophryne gardineri, Sooglossus gardineri] NW07
         |            `--S. pipilodryas (Gerlach & Willi 2002) [=Sooglossus pipilodryas] NW07
         `--+--Ranoidea EJK08
            `--+--Hyloidea PW11
               `--+--Calyptocephallelidae [Calyptocephalellinae, Calyptocephalellini] PW11
                  |    |--Calyptocephalella Strand 1928 FG06
                  |    |    `--C. gayi PW11
                  |    `--Telmatobufo Schmidt 1952 PW11, FG06
                  |         |--T. bullocki PW11
                  |         `--T. venustus FG06
                  `--Myobatrachidae [Australobatrachia, Myobatrachoidea] PW11
                       |  i. s.: Kankanophryne Heyer & Liem 1976 FKM82
                       |           `--*K. occidentalis FKM82
                       |         Glauertia FKM82
                       |--Limnodynastinae PW11
                       `--Myobatrachinae PW11

Phthanobatrachia [Athesphatanura, Ceratophrydes, Chthonobatrachia, Cladophrynia, Diphyabatrachia, Hyloides, Leptodactyliformes, Meridianura, Nobleobatrachia, Notogaeanura, Tinctanura] FG06

*Type species of generic name indicated

REFERENCES

[BB03] Biju, S. D., & F. Bossuyt. 2003. New frog family from India reveals an ancient biogeographical link with the Seychelles. Nature 425: 711–714.

[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.

[FKM82] Farris, J. S., A. G. Kluge & M. F. Mickevich. 1982. Phylogenetic analysis, the monothetic group method, and myobatrachid frogs. Systematic Zoology 31 (3): 317–327.

[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.

[NW07] Nussbaum, R. A., & S.-H. Wu. 2007. Morphological assessments and phylogenetic relationships of the Seychellean frogs of the family Sooglossidae (Amphibia: Anura). Zoological Studies 46 (3): 322–335.

[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.

Eleutherodactylus

Eleutherodactylus guantanamera, copyright Ariel Rodriguez.

Belongs within: Hyloidea.

Eleutherodactylus is a very diverse genus of directly developing, mostly small frogs found in Central and South America.

Eleutherodactylus Duméril & Bibron 1841 [incl. Pelorius Hedges 1989; Eleutherodactylini] FG06
    |--+--+--E. counouspeus (Schwartz 1964) PW11, FG06 [=Euhyas counouspea FG06]
    |  |  `--+--+--E. inoptatus Barbour 1914 PW11, FG06 [=Pelorius inoptatus FG06]
    |  |     |  `--+--E. chlorophenax Schwartz 1976 PW11, FG06 [=Pelorius chlorophenax FG06]
    |  |     |     `--E. nortoni Schwartz 1976 PW11, FG06 [=Pelorius nortoni FG06]
    |  |     `--+--+--E. bothroboans PW11
    |  |        |  `--E. ruthae Noble 1923 PW11, FG06 [=Pelorius ruthae FG06]
    |  |        `--+--E. hypostenor Schwartz 1965 PW11, FG06 [=Pelorius hypostenor FG06]
    |  |           `--E. parapelates Hedges & Thomas 1987 PW11, FG06 [=Pelorius parapelates FG06]
    |  `--+--+--E. richmondi Stejneger 1904 PW11, FG06 [=Euhyas richmondi FG06]
    |     |  `--E. unicolor PW11
    |     `--+--+--+--E. auriculatoides PW11
    |        |  |  `--E. patriciae PW11
    |        |  `--+--+--+--E. fowleri PW11
    |        |     |  |  `--E. lamprotes PW11
    |        |     |  `--+--E. sommeri PW11
    |        |     |     `--E. wetmorei PW11
    |        |     `--+--+--E. leberi PW11
    |        |        |  `--E. melacara PW11
    |        |        `--+--E. varians PW11
    |        |           `--+--E. guantanamera PW11
    |        |              `--E. ionthus PW11
    |        `--+--+--+--E. minutus Noble 1923 PW11, FG06 [=Euhyas minuta FG06]
    |           |  |  `--E. poolei PW11
    |           |  `--+--+--E. pituinus PW11
    |           |     |  `--+--E. parabates Schwartz 1964 PW11, FG06 [=Euhyas parabates FG06]
    |           |     |     `--+--E. abbotti PW11
    |           |     |        `--+--E. audanti PW11
    |           |     |           `--E. haitianus PW11
    |           |     `--+--E. eileenae PW11
    |           |        `--+--E. glamyrus PW11
    |           |           `--+--+--E. mariposa PW11
    |           |              |  `--E. ronaldi PW11
    |           |              `--+--E. bartonsmithi PW11
    |           |                 `--+--E. auriculatus PW11
    |           |                    `--E. principalis PW11
    |           `--+--+--+--E. barlagnei PW11
    |              |  |  `--E. pinchoni PW11
    |              |  `--+--E. johnstonei Barbour 1914 PW11, B-AC-O04
    |              |     `--+--E. amplinympha PW11
    |              |        `--E. locustus PW11
    |              `--+--+--E. antillensis PW11
    |                 |  `--+--E. brittoni PW11
    |                 |     `--+--E. cochranae PW11
    |                 |        `--E. hedricki PW11
    |                 `--+--E. gryllus PW11
    |                    `--+--+--E. coqui PW11
    |                       |  `--E. schwartzi PW11
    |                       `--+--E. portoricensis PW11
    |                          `--E. wightmanae PW11
    `--+--+--+--E. symingtoni Schwartz 1957 PW11, FG06 [=Euhyas symingtoni FG06]
       |  |  `--E. zeus Schwartz 1958 PW11, FG06 [=Euhyas zeus FG06]
       |  `--+--E. marnockii (Cope 1878) PW11, FG06 [=E. (Syrrhophus) marnockii FG06]
       |     `--+--E. nitidus Peters 1870 PW11, FG06 [=E. (Syrrhophus) nitidus FG06]
       |        `--E. pipilans (Taylor 1940) PW11, FG06 [=Syrrhophus pipilans FG06]
       `--+--+--+--E. klinikowskii Schwartz 1959 PW11, FG06 [=Euhyas klinikowskii FG06]
          |  |  `--E. zugi Schwartz 1958 PW11, FG06 [=Euhyas zugi FG06]
          |  `--+--+--+--E. brevirostris Shreve 1936 PW11, FG06 [=Craugastor brevirostris FG06]
          |     |  |  `--E. ventrilineatus Shreve 1936 PW11, FG06 [=Euhyas ventrilineata FG06]
          |     |  `--+--E. glandulifer Cochran 1935 PW11, FG06 [=Euhyas glandulifer FG06]
          |     |     `--E. sciagraphus Schwartz 1973 PW11, FG06 [=Euhyas sciagraphus FG06]
          |     `--+--E. paulsoni Schwartz 1964 PW11, FG06 [=Euhyas paulsoni FG06]
          |        `--+--+--E. furcyensis Shreve & Williams 1963 PW11, FG06 [=Euhyas furcyensis FG06]
          |           |  `--E. rufifemoralis PW11
          |           `--+--+--E. apostates Schwartz 1973 PW11, FG06 [=Euhyas apostates FG06]
          |              |  `--E. oxyrhyncus Duméril & Bibron 1841 PW11, FG06 [=Euhyas oxyrhyncha FG06]
          |              `--+--E. jugans Cochran 1937 PW11, FG06 [=Euhyas jugans FG06]
          |                 `--+--E. glanduliferoides Shreve 1936 PW11, FG06 [=Euhyas glanduliferoides FG06]
          |                    `--+--E. thorectes Hedges 1988 PW11, FG06 [=Euhyas thorectes FG06]
          |                       `--+--+--E. bakeri PW11
          |                          |  `--+--E. dolomedes Hedges & Thomas 1992 PW11, FG06 [=Euhyas dolomedes FG06]
          |                          |     `--E. glaphycompus Schwartz 1973 PW11, FG06 [=Euhyas glaphycompus FG06]
          |                          `--+--E. heminota PW11
          |                             `--+--E. corona Hedges & Thomas 1992 PW11, FG06 [=Euhuyas corona FG06]
          |                                `--+--E. amadeus Hedges, Thomas & Franz 1987 PW11, FG06 (see below for synonymy)
          |                                   `--+--E. caribe Hedges & Thomas 1992 PW11, FG06 [=Euhyas caribe FG06]
          |                                      `--E. eunaster PW11
          `--+--+--E. schmidti Noble 1923 PW11, FG06 [=Euhyas schmidti FG06]
             |  `--+--E. albipes Barbour & Shreve 1937 PW11, FG06 [=Euhyas albipes FG06]
             |     `--+--E. maestrensis Diaz, Cádiz & Navarro 2005 PW11, FG06 [=Euhyas maestrensis FG06]
             |        `--+--E. dimidiatus Cope 1862 PW11, FG06 [=Euhyas dimidiatus FG06]
             |           `--E. emiliae Dunn 1926 PW11, FG06 [=Euhyas emiliae FG06]
             `--+--E. greyi Dunn 1926 PW11, FG06 [=Euhyas greyi FG06]
                `--+--+--E. atkinsi Dunn 1925 PW11, FG06 [=Euhyas atkinsi FG06]
                   |  |--+--+--E. guanahacabibes Estrada & Rodriguez 1985 PW11, FG06 [=Euhyas guanahacabibes FG06]
                   |  |  |  `--+--E. casparii Dunn 1926 PW11, FG06 [=Euhyas casparii FG06]
                   |  |  |     `--E. planirostris Cope 1862 PW11, FG06 [=E. (Euhyas) planirostris FG06]
                   |  |  `--+--+--E. goini Schwartz 1960 PW11, FG06 [=Euhyas goini FG06]
                   |  |     |  `--+--E. rogersi PW11
                   |  |     |     `--E. tonyi Estrada & Hedges 1997 PW11, FG06 [=Euhyas tonyi FG06]
                   |  |     `--+--E. pezopetrus PW11
                   |  |        `--+--E. pinarensis PW11
                   |  |           `--+--E. blairhedgesi PW11
                   |  |              `--E. thomasi Schwartz 1959 PW11, FG06 [=Euhyas thomasi FG06]
                   |  `--+--+--E. gundlachi Schmidt 1920 PW11, FG06 [=Euhyas gundlachi FG06]
                   |     |  `--+--E. intermedius Barbour & Shreve 1937 PW11, FG06 [=Euhyas intermedia FG06]
                   |     |     `--E. varleyi Dunn 1925 PW11, FG06 [=Euhyas varleyi FG06]
                   |     `--+--+--E. cubanus Barbour 1942 PW11, FG06 [=Euhyas cubana FG06]
                   |        |  `--E. orientalis Barbour & Shreve 1937 PW11, FG06 [=Euhyas orientalis FG06]
                   |        `--+--E. etheridgei Schwartz 1958 PW11, FG06 [=Euhyas etheridgei FG06]
                   |           `--+--E. iberia Estrada & Hedges 1996 PW11, FG06 [=Euhyas iberia FG06]
                   |              `--+--E. jaumei Estrada & Alonso 1997 PW11, FG06 [=Euhyas jaumei FG06]
                   |                 `--E. limbatus Cope 1862 PW11, FG06 [=Euhyas limbata FG06]
                   `--+--+--+--E. alcoae Schwartz 1971 PW11, FG06 [=Euhyas alcoae FG06]
                      |  |  `--E. armstrongi Noble & Hassler 1933 PW11, FG06 [=Euhyas armstrongi FG06]
                      |  `--+--E. darlingtoni PW11
                      |     `--E. leoncei Shreve & Williams 1963 PW11, FG06 [=Euhyas leoncei FG06]
                      `--+--+--+--E. bresslerae PW11
                         |  |  `--+--E. acmonis Schwartz 1960 PW11, FG06 [=Euhyas acmonis FG06]
                         |  |     `--E. ricordii Duméril & Bibron 1841 PW11, FG06 [=E. (Euhyas) ricordii FG06]
                         |  `--+--E. probolaeus PW11
                         |     `--+--E. monensis Meerwarth 1901 PW11, FG06 [=Euhyas monensis FG06]
                         |        `--+--E. grahami Dunn 1926 PW11, FG06 [=Euhyas grahami FG06]
                         |           `--+--E. lentus PW11
                         |              `--+--E. rhodesi PW11
                         |                 `--E. weinlandi Barbour 1914 PW11, FG06 [=Euhyas weinlandi FG06]
                         `--+--+--E. cuneatus Cope 1862 PW11, FG06 (see below for synonymy)
                            |  `--E. turquinensis Barbour & Shreve 1937 PW11, FG06 [=Euhyas turquinensis FG06]
                            `--+--+--E. riparius PW11
                               |  `--E. rivularis Díaz, Estrada & Hedges 2001 PW11, DEH01 [=E. (Euhyas) rivularis DEH01]
                               `--+--E. toa Estrada & Hedges 1991 PW11, FG06 [=Euhyas toa FG06]
                                  `--+--E. luteolus Gosse 1851 PW11, FG06 [=Euhyas luteola FG06]
                                     `--+--+--+--E. sisyphodemus Crombie 1977 PW11, FG06 [=Euhyas sisyphodemus FG06]
                                        |  |  `--+--E. cavernicola Lynn 1954 PW11, FG06 [=Euhyas cavernicola FG06]
                                        |  |     `--E. grabhami Dunn 1926 PW11, FG06 [=Euhyas grabhami FG06]
                                        |  `--+--E. jamaicensis Barbour 1910 PW11, FG06 [=Euhyas jamaicensis FG06]
                                        |     `--+--E. nubicola Dunn 1926 PW11, FG06 [=E. (Euhyas) nubicola FG06]
                                        |        `--+--E. andrewsi Lynn 1937 PW11, FG06 [=Euhyas andrewsi FG06]
                                        |           `--E. orcutti Dunn 1928 PW11, FG06 [=Euhyas orcutti FG06]
                                        `--+--E. alticola Lynn 1937 PW11, FG06 [=Euhyas alticola FG06]
                                           `--+--E. fuscus Lynn & Dent 1943 PW11, FG06 [=Euhyas fusca FG06]
                                              `--+--E. junori Dunn 1926 PW11, FG06 [=Euhyas junori FG06]
                                                 `--+--E. gossei Dunn 1926 PW11, FG06 [=Euhyas gossei FG06]
                                                    `--+--E. griphus Crombie 1986 PW11, FG06 [=Euhyas griphus FG06]
                                                       `--+--+--E. cundalli PW11
                                                          |  `--E. glaucoreius Schwartz & Fowler 1973 PW11, FG06 (see below for synonymy)
                                                          `--+--E. pantoni Dunn 1926 PW11, FG06 [=Euhyas pantoni FG06]
                                                             `--E. pentasyringos Schwartz & Fowler 1973 PW11, FG06 (see below for synonymy)

Eleutherodactylus incertae sedis:
  E. angustidigitorum Taylor 1940 NW07, FG06 [=Syrrhophus angustidigitorum FG06]
  E. anolirex Lynch 1983 B-AC-O04
  E. (Eleutherodactylus) binotatus FG06
  E. boconoensis Rivero & Mayorga 1973 B-AC-O04
  E. chloronotus FG06
  E. chlorosoma Rivero 1984 B-AC-O04
  E. colostichos La Marca & Smith 1982 B-AC-O04
  E. conspicillatus B-AC-O04
  E. costaricensis M83
  E. diastema FG06
  E. discoidalis FG06
  E. duellmani FG06
  E. erythropleurus NW07
  E. ginesi (Rivero 1964) B-AC-O04
  E. guentheri NW07
  E. hedriki SWK87
  E. jasperi FG06
  E. karlschmidti Grant 1931 NW07, FG06 [=Euhyas karlschmidti FG06]
  E. lancinii Donoso-Barros 1968 B-AC-O04
  E. lentiginosus Rivero 1984 B-AC-O04
  E. malkini B-AC-O04
  E. megalops B-AC-O04
  E. melanoproctus Rivero 1984 B-AC-O04
  E. mercedesae FG06
  E. mondolfii Rivero 1984 B-AC-O04
  E. nicefori Cochran & Goin 1971 B-AC-O04
  E. nigrovittatus FG06
  E. paramerus Rivero 1984 B-AC-O04
  E. parvus NW07
  E. pedimontanus La Marca 2004 B-AC-O04
  E. pleurostriatus Rivero 1984 B-AC-O04
  E. prolixodiscus Lynch 1978 B-AC-O04
  E. pulidoi Rivero 1984 B-AC-O04
  E. rearki M83
  E. ridens M81
  E. supernatis FG06
  E. thymelensis FG06
  E. tubernasus Rivero 1984 B-AC-O04
  E. tubolus KF69
  E. underwoodi M83
  E. unistrigatus B-AC-O04
  E. urichi (Boettger 1894) B-AC-O04
  E. vanadisae La Marca 1984 B-AC-O04
  E. w-nigrum B-AC-O04
  E. yustizi Barrio-Amorós & Chacón-Ortiz 2004 B-AC-O04

Eleutherodactylus amadeus Hedges, Thomas & Franz 1987 PW11, FG06 [=Euhyas amadeus FG06]

Eleutherodactylus cuneatus Cope 1862 PW11, FG06 [=Euhyas cuneata FG06; incl. El. sierramaestrae DEH01]

Eleutherodactylus glaucoreius Schwartz & Fowler 1973 PW11, FG06 [=Euhyas glaucoreia FG06]

Eleutherodactylus pentasyringos Schwartz & Fowler 1973 PW11, FG06 [=Euhyas pentasyringos FG06]

*Type species of generic name indicated

REFERENCES

[B-AC-O04] Barrio-Amorós, C. L., & A. Chacón-Ortiz. 2004. Un nuevo Eleutherodactylus (Anura, Leptodactylidae) de la Cordillera de Mérida, Andes de Venezuela. Graellsia 60 (1): 3–11.

[DEH01] Díaz, L. M., A. R. Estrada & S. B. Hedges. 2001. A new riparial frog of the genus Eleutherodactylus (Anura: Leptodactylidae) from eastern Cuba. Caribbean Journal of Science 37 (1–2): 63–71.

[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.

[M81] Miyamoto, M. M. 1981. Congruence among character sets in phylogenetic studies of the frog genus Leptodactylus. Systematic Zoology 30 (3): 281–290.

[M83] Miyamoto, M. M. 1983. Biochemical variation in the frog Eleutherodactylus bransfordii: geographic patterns and cryptic species. Systematic Zoology 32 (1): 43–51.

[NW07] Nussbaum, R. A., & S.-H. Wu. 2007. Morphological assessments and phylogenetic relationships of the Seychellean frogs of the family Sooglossidae (Amphibia: Anura). Zoological Studies 46 (3): 322–335.

[SWK87] Snyder, N. F. R., J. W. Wiley & C. B. Kepler. 1987. The Parrots of Luquillo: Natural history and conservation of the Puerto Rican parrot. Western Foundation of Vertebrate Zoology: Los Angeles.

Brachycephalidae

Steindachner's robber frog Ischnocnema guentheri, copyright Mario Sacramento.

Belongs within: Hyloidea.

The Brachycephalidae are a group of small, leaf-litter dwelling frogs found in southern Brazil (Frost et al. 2006).

Characters (from Frost et al. 2006): Distinguishable sternum absent through fusion; epicoracoid cartilages completely ossified with coracoids and clavicles; digits reduced; axillary amplexus; direct development with very large terrestrial eggs.

Brachycephalidae [Brachycephalina] FG06
    |  i. s.: ‘Eleutherodactylus’ juipoca FG06
    |--Phyllonastes Heyer 1977 FG06
    |--Cornufer Tschudi 1838 [Cornuferinae] FG06
    |--Brachycephalus Fitzinger 1826 FG06
    |    `--B. ephippium FG06
    `--Ischnocnema Reinhardt & Lütken 1862 PW11, FG06
         |  i. s.: I. sanderi Padial, Reichle & De la Riva 2005 PRDR05
         |         I. quixensis FG06
         |--+--I. holti PW11
         |  `--I. juipoca PW11
         `--+--I. parva PW11
            `--+--I. guentheri PW11
               `--I. hoehnei 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.

[PRDR05] Padial, J. M., S. Reichle & I. De la Riva. 2005. Ischnocnema sanderi Padial, Reichle & De la Riva, 2005 (Amphibia, Anura, Leptodactylidae): a nomenclatural note. Graellsia 61 (1): 143.

[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.

Neospathodus

Platform element of Neospathodus, from Sweet (1988).

Belongs within: Gondolellidae.

Neospathodus is a genus of conodonts known from the early Triassic represented only by segminate platform (Pa) elements, the remaining positions of the standard apparatus perhaps lacking mineralised elements (Sweet 1988).

Neospathodus Mosher 1968 S88
    |--N. arcucristatus RR79
    |--N. bransoni CB86
    |--N. collinsoni S88
    |--N. conservativus K79
    |--N. constricta TE01
    |--N. cristagalli K79
    |--N. dieneri TE01
    |--N. divergens RR79
    |--N. homeri TE01
    |--N. hungaricus TE01
    |--N. kummeli K79
    |--N. newpassensis K79
    |--N. pakistanensis TE01
    |--N. timorensis K79
    |--N. triangularis TE01
    `--N. waageni S88

*Type species of generic name indicated

REFERENCES

[CB86] Chen D.-Q. & Bao H. 1986. Lower Permian ostracodes from the Chihsia Formation of Jurong and Longtan, Jiangsu Province. Acta Micropalaeontologica Sinica 3 (2): 107–132.

[K79] Kummel, B. 1979. Triassic. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A351–A389. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[RR79] Ross, C. A., & J. R. P. Ross. 1979. Permian. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A291–A350. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[S88] Sweet, W. C. 1988. The Conodonta: Morphology, taxonomy, paleoecology, and evolutionary history of a long extinct animal phylum. Clarendon Press: Oxford.

[TE01] Tong, J., & D. H. Erwin. 2001. Triassic gastropods of the southern Qinling Mountains, China. Smithsonian Contributions to Paleobiology 92: 1–47.

Neogondolella

Pa element of Neogondolella, from Sweet (1988).

Belongs within: Gondolellidae.

Neogondolella is a genus of conodonts known from the Late Permian and Triassic, at least some species of which possessed an apparatus with ramiform elements in the S and M positions and segminiplanate elements in the Pa position (Sweet 1988).

Neogondolella Bender & Stoppel 1965 S88
    |--*N. mombergensis S88
    |--N. bifurcata S88
    |--N. bisselli RR79
    |--N. bitteri S88
    |--N. bulgarica S88
    |--N. carinata RR79
    |--N. carpathica S88
    |--N. clarki S88
    |--N. constricta K79
    |--N. excelsa TE01
    |--N. gracilis S88
    |--N. idahoensis RR79
    |--N. jubata K79
    |--N. leveni S88
    |--N. longa S88
    |--N. milleri K79
    |--N. navicula TE01
    |--N. noah S88
    |--N. orientalis S88
    |--N. phosphoriensis S88
    |--N. praebisselli S88
    |--N. regale S88
    |--N. rosenkrantzi RR79
    |--N. serrata DX84
    |    |--N. s. serrata RR79
    |    `--N. s. postserrata RR79
    |--N. shoshonensis S88
    |--N. subcarinata S88
    |--N. timorensis S88
    |--N. transita S88
    `--N. triangularis S88

*Type species of generic name indicated

REFERENCES

[DX84] Ding Y., Xia G., Duan C., Li W., Liu X. & Liang Z. 1984. Study on the early Permian stratigraphy and fauna in Zhesi district, Nei Mongol Zizhiqu (Inner Mongolia). Bulletin Tianjin Institure Geol. Min. Res. 10.

[K79] Kummel, B. 1979. Triassic. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A351–A389. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[RR79] Ross, C. A., & J. R. P. Ross. 1979. Permian. In: Robison, R. A., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt A. Introduction. Fossilisation (Taphonomy), Biogeography and Biostratigraphy pp. A291–A350. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[S88] Sweet, W. C. 1988. The Conodonta: Morphology, taxonomy, paleoecology, and evolutionary history of a long extinct animal phylum. Clarendon Press: Oxford.

[TE01] Tong, J., & D. H. Erwin. 2001. Triassic gastropods of the southern Qinling Mountains, China. Smithsonian Contributions to Paleobiology 92: 1–47.

Drepanoistodontidae

Element of Drepanoistodus suberectus, copyright Olle Hints.

Belongs within: Conodonti.

The Drepanoistodontidae are a group of conodonts known from the Ordovician possessing apparatus composed of recurved and/or geniculate coniform elements (Sweet 1988). Members of this family comprise the core of what has been recognised as the form-family Distacodontidae; however, the use of this latter name was discouraged by Sweet (1988) due to the type genus Distacodus being a nomen dubium of uncertain affinities.

<==Drepanoistodontidae [Distacodontacea, Distacodontidae] S88
    |--Acodina Stauffer 1940 H62
    |    `--A. cuspidata (Stauffer 1940) WSQ86
    |--Proacodus Müller 1959 M62
    |    `--*P. obliquus Müller 1959 M62
    |--Acontiodus Pander 1856 H62
    |    |--*A. latus Pander 1856 H62
    |    `--A. franconicus LM05
    |--Distacodus Pander 1856 (n. d.) S88 [=Machairodus Pander 1856 non Kaup 1833 H62; incl. Machairodia Smith 1907 H62]
    |    |--*D. incurvus (Pander 1856) [=*Machairodus incurvus] H62
    |    `--D. ensiformis H62
    |--Furnishina Müller 1959 M62
    |    |--*F. furnishi Müller 1959 M62, DJ71 [=Sagittodontus furnishi DJ71]
    |    |--F. alata Szaniawski 1971 RNP03
    |    `--F. polonica Szaniawski 1971 RNP03
    |--Paltodus Pander 1856 S88, H62
    |    |--*P. subequalis Pander 1856 H62
    |    |--P. deltifer J79
    |    |--P. jemtlandicus PBJ03
    |    |--P. spurius LM05
    |    `--P. truncatus H62
    |--Paroistodus Lindström 1971 S88
    |    |--P. horridus (Barnes & Poplawski 1973) S88
    |    |--P. numarcuatus S88
    |    |--P. originalis PBJ03
    |    |--P. parallelus PBJ03
    |    `--P. proteus PBJ03
    `--Drepanoistodus Lindström 1971 SS05
         |--*D. forceps (Lindström 1955) [=Oistodus forceps] SS05
         |--D. angulensis PBJ03
         |--D. basiovalis PBJ03
         |--D. concavus PBJ03
         |--D. latus PBJ03
         |--D. lucidus PBJ03
         `--D. suberectus (Branson & Mehl 1933) SS05 (see below for synonymy)

Drepanoistodus suberectus (Branson & Mehl 1933) SS05 [=Oistodus suberectus SS05, Drepanodus suberectus DJ71; incl. D. homocurvatus DJ71, Oistodus inclinatus DJ71]

*Type species of generic name indicated

REFERENCES

[DJ71] Druce, E. C., & P. J. Jones. 1971. Cambro-Ordovician conodonts from the Burke River Structural Belt, Queensland. Commonwealth of Australia, Bureau of National Development, Bureau of Mineral Resources, Geology and Geophysics, Bulletin 110: 1–159.

[H62] Hass, W. H. 1962. Conodonts. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W3–W69. Geological Society of America, and University of Kansas Press.

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