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.

Staphylinini

Acylophorus pratensis, copyright Richard Migneault.

Belongs within: Staphylininae.
Contains: Amblyopinina, Cyrtoquediina, Indoquedius, Quedius, Philonthina, Staphylinina.

The Staphylinini are a cosmopolitan group of relatively large staphylinid beetles, including the rove beetles that are the most familiar members of the family. Representatives of the Staphylinini have antennae inserted closer to the eyes than to each other, a well developed apical maxillary palpomere, a relatively wide neck (if any), and no separate sclerotised plate anterior to the prosternum (Newton et al. 2001). A major clade within the group has been referred to as the 'Staphylinini propria', members of which are often distinguished by a dorsal basal ridge on the head capsule, a complete fusion of the pronotum and prosternum inside the procoxal cavity, and a transverse ridge on the hind coxa (Brunke et al. 2016).

Staphylinini [Quediini]
    |  i. s.: Valdiviodes BC16
    |         Alesiella BC16
    |         Quediomacrus BC16
    |         Lonia BC16
    |         Quelaestrygon BC16
    |         Strouhalium gracilicorne BC16
    |--Cretoquedius BC16
    `--+--Atanygnathus Jacobson 1909 NT01 (see below for synonymy)
       |    `--A. acuminatus BC16
       `--+--Antimerus punctipennis BC16
          |--+--Afroquedius sexpunctatus BC16
          |  `--Amblyopinina BC16
          `--+--Cyrtoquediina BC16
             `--+--+--Erichsonius Fauvel 1874 NT01 (see below for synonymy)
                |  |    |--E. nanus BC16
                |  |    `--E. patella N90
                |  `--Acylophorina BC16
                |       |  i. s.: Australotarsius grandis BC16
                |       |         Acylohsellus BC16
                |       |         Stevensia longipennis BC16
                |       |--Hemiquedius Casey 1915 BC16, NT01
                |       |    `--H. ferox (LeConte 1878) NT01
                |       `--+--Anaquedius Casey 1915 BC16, NT01
                |          |    `--A. vernix (LeConte 1878) NT01
                |          `--+--+--Anchocerus BC16
                |             |  `--Paratolmerus siamensis BC16
                |             `--Acylophorus Nordmann 1837 BC16, NT01 (see below for synonymy)
                |                  |--A. capensis BC16
                |                  |--A. pratensis BC16
                |                  `--A. wagenschieberi BC16
                `--+--Indoquedius BC16
                   `--+--Quediina BC16
                      |    |--Quedius BC16
                      |    |--Pseudorientis BC16
                      |    |--Velleiopsis BC16
                      |    `--Beeria Hatch 1957 NT01
                      |         `--B. nematocera (Casey 1915) [incl. B. punctata Hatch 1957] NT01
                      `--Staphylinini propria BC16
                           |  i. s.: Eucibdelus BC16
                           |         Ioma BC16
                           |         Weiserianum BC16
                           |         Haematodes BC16
                           |         Euristus BC16
                           |--+--Philothalpus BC16
                           |  |    |--P. bilobus BC16
                           |  |    `--P. falini BC16
                           |  `--Xanthopygina BC16
                           |       |  i. s.: Gastrisus Sharp 1876 NT01
                           |       |         Trigonopselaphus Gemminger & Harold 1868 (see below for synonymy) NT01
                           |       |--+--Isanopus sallaei BC16
                           |       |  `--+--Nordus fungicola BC16
                           |       |     `--Plociopterus BC16
                           |       `--+--Glenus flohri BC16
                           |          `--Xanthopygus Kraatz 1857 BC16, GN09 (see below for synonymy)
                           |               |--X. chapareanus BC16
                           |               `--X. xanthopygus Nordmann 1837 [incl. X. cacti Horn 1868] NT01
                           `--+--+--Philonthina BC16
                              |  `--Algon BC16
                              |       |--A. hollowayae BC16
                              |       `--A. oculatus BC16
                              `--+--Staphylinina BC16
                                 `--Anisolinina BC16
                                      |--+--Misantlius gebieni BC16
                                      |  `--Hesperosoma pederseni BC16
                                      `--+--Tolmerinus BC16
                                         `--+--Tympanophorus Nordmann 1837 BC16, NT01
                                            |    |--T. concolor Sharp 1884 NT01
                                            |    `--T. puncticollis Erichson 1840 (see below for synonymy) NT01
                                            `--+--Anisolinus BC16
                                               `--Pammegus ruficollis BC16

Acylophorus Nordmann 1837 BC16, NT01 [incl. Amacylophorus Smetana 1971 NT01, Palpacylophorus Smetana 1971 NT01]

Atanygnathus Jacobson 1909 NT01 [incl. Tanygnathinus Reitter 1909 NT01, Tanygnathus Erichson 1839 non Wagler 1832 NT01; Tanygnathinina BC16]

Erichsonius Fauvel 1874 NT01 [incl. Actobius Fauvel 1876 NT01, Sectophilonthus Tottenham 1949 NT01; Erichsoniina BC16]

Trigonopselaphus Gemminger & Harold 1868 [incl. Trigonophorus Nordmann 1837 nec Stephens 1829 nec Solier 1851] NT01

Tympanophorus puncticollis Erichson 1840 [incl. Xanthopygus borealis Hatch 1957, T. borealis] NT01

Xanthopygus Kraatz 1857 BC16, GN09 [incl. Heteropygus Bernhauer 1906 NT01, Lampropygus Sharp 1884 NT01]

*Type species of generic name indicated

REFERENCES

[BC16] Brunke, A. J., S. Chatzimanolis, H. Schillhammer & A. Solodovnikov. 2016. Early evolution of the hyperdiverse rove beetle tribe Staphylinini (Coleoptera: Staphylinidae: Staphylininae) and a revision of its higher classification. Cladistics 32 (4): 427–451.

[GN09] Grebennikov, V. V., & A. F. Newton. 2009. Good-bye Scydmaenidae, or why the ant-like stone beetles should become megadiverse Staphylinidae sensu latissimo (Coleoptera). European Journal of Entomology 106: 275–301.

[N90] Newton, A. F., Jr. 1990. Insecta: Coleoptera Staphylinidae adults and larvae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1137–1174. John Wiley & Sones: New York.

[NT01] Newton, A. F., M. K. Thayer, J. S. Ashe & D. S. Chandler. 2001. Staphylinidae Latreille, 1802. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 272–418. CRC Press: Boca Raton.

Last updated: 16 December 2020.

Aleochara

Aleochara curtula, copyright U. Schmidt.

Belongs within: Aleocharini.

Aleochara is a genus of aleocharine staphylinids characterised by a partially concealed head and five-segmented tarsi with the first segment much longer than the second. They are commonly associated with decaying matter (Olliff 1886).

Characters (from Olliff 1886): Mentum very short, narrowed and slightly emarginate in front. Ligula very short, broad, bifid. Maxillary palpi 5-segmented, the second and third segment subequal, the fifth very small. Labial palpi 4-segmented, the first segment longer than the others, the fourth very small. Mandibles unarmed. Labrum transverse, truncate in front. Head ovate, retracted, not narrowed behind. Antennae short, thickened towards the extremity, 1st, 2nd and 3rd segments elongate, the following, with the exception of the 11th, transverse. Prothorax broadly transverse, narrowed in front, rounded at the sides. Elytra slightly rounded behind. Abdomen either parallel-sided or narrowed behind. Legs rather short; intermediate coxae somewhat remote; tarsi 5:5:5, first segment of posterior pair distinctly longer than 2nd.

Aleochara Gravenhorst 1802 O86
    |--A. actae Olliff 1886 O86
    |--A. analis (Macleay 1871) [=Oxypoda analis] O86
    |--A. asiatica O86
    |--A. baliola Olliff 1886 O86
    |--A. bilineata P91
    |--A. bipunctata C01
    |--A. bisulcata (Redt. 1867) [=Oxypoda bisulcata] O86
    |--A. brachialis Jekel 1873 O86
    |--A. crassiuscula G89
    |--A. croceipennis Motschulsky 1858 [incl. A. maculipennis Kraatz 1859, A. sanguinipennis Kraatz 1859] O86
    |--A. curtula (Goeze 1777) M04
    |--A. fuscipes O86
    |--A. haemorrhoidalis O86
    |--A. insuavis Olliff 1886 O86
    |--A. lata Gravenhorst 1802 M04
    |--A. marginata Fauvel 1877 O86
    |--A. mastersi Macleay 1871 O86
    |--A. moesta C01
    |--A. nitida C01
    |--A. (Isochara) perturbans Fenyes 1921 F21
    |--A. puberula Klug 1883 [incl. A. armitagei Wollaston 1854, A. decorata Aubé 1850, A. dubia Fauvel 1863] O86
    |--A. punctum Fauvel 1878 O86
    |--A. scutellaris Lucas 1846 E12
    |--A. semirubra Fauvel 1877 O86
    |--A. speculifera Erichson 1842 O86
    |--A. subaenea ED04
    |--A. tristis C01
    `--A. vicina Olliff 1886 O86

*Type species of generic name indicated

REFERENCES

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] Band II. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[ED04] Ewers, R. M., & R. K. Didham. 2004. A first assessment of New Zealand beetles restricted to large forest areas. Weta 27: 33–40.

[F21] Fenyes, A. 1921. New genera and species of Aleocharinae with a polytomic synopsis of the tribes. Bulletin of the Museum of Comparative Zoology 65 (2): 15–36.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 5–72.

[M04] Maruyama, M. 2004. Redescription of the genus Creochara (Coleoptera: Staphylinidae: Aleocharinae: Aleocharini) and its systematic position. Canadian Entomologist 136: 621–637.

[O86] Olliff, A. S. 1886. A revision of the Staphylinidae of Australia. Part I. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 403–473, pl. 7.

[P91] Petrushov, A. Z. 1991. Selection and mechanisms of ambush resistance in Metaseiulus occidentalis N. (Acarina: Phytoseiidae). In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 741–748. SPB Academic Publishing: The Hague.

Psocodea

Mating pair of Rhyopsocus eclipticus (male on left), copyright Diane Young.

Belongs within: Paraneoptera.
Contains: Psocathropetae, Trogioidea, Lepidopsocidae, Psocomorpha, Nanopsocetae, Amphientometae.

The Psocodea include the barklice, small insects that mostly feed on fungi and other micro-organisms, as well as the true parasitic lice. It is now generally accepted that the latter are a derived subgroup of the former. The most basal division in the Psocodea is between the clades Trogiomorpha and Psocina; Trogiomorpha have antennae with more than twenty segments and a strong posterior spine on the paraproct (the lateral sclerites alongside the anus) whereas Psocina have fewer antennal segments and lack the paraproct spine. Psocina also have the hypopharyngeal filaments fused for at least part of their length whereas they remain separate in Trogiomorpha. Trogiomorpha are divided between the Psocathropetae and Atropetae, with the Atropetae having a short, broad head, a conical sensillum on the inner side of the second segment of the maxillary palp, and no nodulus on the fore wing (if present) (New & Lienhard 2007). Within the Psocina, members of the Troctomorpha lineage have antennae with at least some segments secondarily annulated, wings lacking a pterostigma and often a triangular or T-shaped sclerite on the large subgenital plate (Mockford 1993).

Characters (from Rasnitsyn 2002, for Psocidea): Size small, rarely medium. General appearance highly variable depending on free living or parasitic habit and mode of parasitism. Head with clypeus strongly convex to house cibarial muscles (secondarily less developed in some parasitic forms). Mandible chewing (reverted and thus tearing instead of biting in elephant louse, reduced in sucking lice). Lacinia free of remaining maxilla, rod-like. Hypopharynx with two ovoid sclerites. Pronotum small. Wings often lost, when present roof-like at rest, rather poor in venation, coupled in flight except in some Palaeozoic species with homonomous wings. Legs cursorial or, in parasites, clinging, with tarsus l-3-segmented (4-segmented in some extinct forms). 1st abdominal sternum lost. Ovipositor present or lost. Aedeagus lost in living species. Cercus lost. Extant forms with abdominal ganglia fused in single ganglionic mass and with 4 or less Malpighian tubes. Ovaries polytrophic. Chromosomes holokinetic.

<==Psocodea [Copeognatha, Corrodentia, Psocida, Psocidea, Psocoptera]
    |--Trogiomorpha [Trogiina] Y02
    |    |--Psocathropetae NL07
    |    `--Atropetae [Trogiformia] NL07
    |         |--Trogioidea NL07
    |         |--Lepidopsocidae NL07
    |         `--Psoquillidae [Psoquilloidea] L02
    |              |--Balliella Badonnel 1949 M93
    |              |    `--*B. ealensis Badonnel 1949 M93
    |              |--Psoquilla Hagen 1865 [incl. Heteropsocus Verrill 1902] L02
    |              |    |--*P. marginepunctata Hagen 1865 NL07 (see below for synonymy)
    |              |    `--P. infuscata Badonnel 1949 B49
    |              |--Eosilla NL07
    |              |    |--*E. (Eosilla) jacobsoni Ribaga 1908 NL07
    |              |    `--E. (Empheriella) denervosa NL07
    |              `--Rhyopsocus Hagen 1876 [incl. Deipnopsocus Enderlein 1903, Rhyopsocopsis Pearman 1929] M93
    |                   |--*R. eclipticus Hagen 1876 [incl. R. phillipsae Sommerman 1956] M93
    |                   |--R. afer (Badonnel 1948) [=Deipnopsocus afer] B49
    |                   |--R. bentonae Sommerman 1956 M93
    |                   |--R. bicornis Badonnel 1986 B86
    |                   |--R. disparilis (Pearman 1931) M93 (see below for synonymy)
    |                   |--R. maculosus M93
    |                   |--R. micropterus Mockford 1971 M93
    |                   |--R. orthatus B86
    |                   |--R. pandanicola Thornton, Lee & Chui 1972 NL07
    |                   |--R. peregrinus (Pearman 1929) L98 [=*Rhyopsocopsis peregrina NL07]
    |                   |--R. phillipsae Sommerman 1956 S56
    |                   |--R. spheciophilus (Enderlein 1903) L98 [=*Deipnopsocus speciophilus NL07]
    |                   `--R. texanus (Banks 1930) (see below for synonymy) M93
    `--Psocina R02
         |--Psocomorpha Y02
         `--Troctomorpha [Amphientomiformia, Amphientomomorpha, Troctopsocoidea] Y02
              |--Nanopsocetae YL10
              `--Amphientometae YL10

Psocodea incertae sedis:
  Tshekarcephalus [Tshekarcephalidae] R02
    `--T. bigladipotens R02
  Surijokopsocus [Surijokopsocidae] R02
  Sinopsocus oligovenus Lin 1976 L02
  Diploperipsocus phagococcus Li & Mockford 1993 L02
  Mascaropsocus spinosus Badonnel & Pearman 1971 SN08
  Miotroctes Pierce 1960 YL10
    `--M. rousei Pierce 1960 YL10
  Westphalopsocus Azar, Nel et al. in Nel, Roques et al. 2013 [Westphalopsocidae] NR13
    `--*W. pumilio Azar, Nel et al. in Nel, Roques et al. 2013 NR13
  Zygopsocus [Zygopsocidae] NR13

*Psoquilla marginepunctata Hagen 1865 NL07 [incl. *Heteropsocus dispar Verrill 1902 NL07, M93, P. dispar B49]

Rhyopsocus disparilis (Pearman 1931) M93 [=Deipnopsocus spheciophilus var. disparilis M93, R. spheciophilus disparilis MG56]

Rhyopsocus texanus (Banks 1930) [=Deipnopsocus texanus; incl. R. pescadori García Aldrete 1984, R. squamosus Mockford & Gurney 1956] M93

*Type species of generic name indicated

REFERENCES

[B49] Badonnel, A. 1949. Psocoptères de la Cote d'Ivoire. Mission Paulian-Delamare (1945). Revue Française d'Entomologie 16: 20–46.

[B86] Badonnel, A. 1986. Psocoptères de Colombie (Insecta, Psocoptera). Spixiana 9 (2): 179–223.

[L02] Li F. 2002. Psocoptera of China vol. 1. Science Press: Beijing.

[L98] Lienhard, C. 1998. Faune de France. France et Régions Limitrophes. 83. Psocoptères Euro-Méditerranéens. Fédération Française des Sociétés de Sciences Naturelles: Paris.

[M93] Mockford, E. L. 1993. North American Psocoptera (Insecta). Sandhill Crane Press, Inc.

[NR13] Nel, A., P. Roques, P. Nel, A. A. Prokin, T. Bourgoin, J. Prokop, J. Szwedo, D. Azar, L. Desutter-Grandcolas, T. Wappler, R. Garrouste, D. Coty, D. Huang, M. S. Engel & A. G. Kirejtshuk. 2013. The earliest known holometabolous insects. Nature 503: 257–261.

[NL07] New, T. R., & C. Lienhard. 2007. The Psocoptera of Tropical South-east Asia. Brill: Leiden.

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Last updated: 27 August 2020.