Telmatobufo

Frost, Darrel R., Grant, Taran, Faivovich, Julián, Bain, Raoul H., Haas, Alexander, De Sá, Célio F. B. Haddad Rafael O., Channing, Alan, Wilkinson, Mark, Donnellan, Stephen C., Raxworthy, Christopher J., Campbell, Jonathan A., Blotto, Boris L., Moler, Paul, Drewes, Robert C., Nussbaum, Ronald A., Lynch, John D., Green, David M. & Wheeler, And Ward C., 2006, The Amphibian Tree Of Life, Bulletin of the American Museum of Natural History 2006 (297), pp. 1-291 : 1-291

erbera and Telmatobufo (presumably

Batrachophrynidae Cope, 1875: 9 . Type genus: apomorphic condition at this level of univer­ Batrachophrynus Peters, 1873 . sality) and horizontal in Batrachophrynus 194 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Telmatobufo and Caudiverbera exhibit the lecular synapomorphies are summarized in condition of the trigeminal nerve passing appendix 5. Further study is needed. medial to the m. adductor mandibulae (the ‘‘E’’ condition); the condition is unknown in [322] FAMILY: LIMNODYNASTIDAE LYNCH, 1971 Batrachophrynus . This characteristic is otherwise known sporadically in Ceratophrys Limnodynastini J.D. Lynch, 1971: 83 . Type ge­ and Lepidobatrachus , some bufonids, Crau­ nus: Limnodynastes Fitzinger, 1843 . gastor , Mixophyes , some hyperoliids, most microhylids, a few ranids, rhacophorids, IMMEDIATELY MORE INCLUSIVE TAXON: Rhinophrynus , and in Sooglossus thomasseti [321] Myobatrachoidea Schlegel, 1850. (J.D. Lynch , 1986). We regard this condition SISTER TAXON: [334] Myobatrachidae as a likely synapomorphy of Batrachophryn­ Schlegel, 1850. idae (or minimally Caudiverbera 1 Telma­ RANGE: Australia and New Guinea, includtobufo), although the distribution of this fea­ ing the Aru Islands. ture across the anuran tree requires further CONTENT: Adelotus Ogilby, 1907 ; Heleiostudy. porus Gray, 1841; Lechriodus Boulenger ,

1882; Limnodynastes Fitzinger, 1843 ; Neo­ [321] SUPERFAMILY: MYOBATRACHOIDEA batrachus Peters, 1863; Notaden Günther ,

SCHLEGEL, 1850 1873; Opisthodon Steindachner, 1867 (see IMMEDIATELY MORE INCLUSIVE TAXON: Systematic Comments and appendix 7); Phi­ [319] Australobatrachia new taxon. loria Spencer, 1901 (including Kyarranus SISTER TAXON: [320] Batrachophrynidae Moore, 1958 ). Cope, 1875. CHARACTERIZATION AND DIAGNOSIS: Lim­ RANGE: Australia and New Guinea. nodynastids are predominantly small to mod­ CONTENT: [322] Limnodynastidae Lynch , erately­sized toad­like terrestrial frogs. Am­ 1971, and [334] Myobatrachidae Schlegel , plexus is inguinal, and with the exception of 1850. Neobatrachus and Notaden , all species are CHARACTERIZATION AND DIAGNOSIS: See the foam­nesters (Martin, 1967). characterization and diagnosis of Australo­ See ‘‘Characterization and diagnosis’’ of batrachia for morphological characters that Australobatrachia for morphological characmay optimize on this branch with further ters that may optimize on this branch. Ford study. At present, there are no morphological and Cannatella (1993) suggested the followcharacters that can be documented to opti­ ing to be a morphological synapomorphy of mize on this branch so justification for rec­ Limnodynastidae : connection between the m. ognizing this taxon is based entirely on mo­ submentalis and m. intermandibularis. But, lecular evidence (listed in appendix 5). with the transfer of Mixophyes to Myoba­ SYSTEMATIC COMMENTS: We recognize two trachidae on the basis of molecular evidence, families within Myobatrachoidea, corre­ this morphological character requires verifisponding substantially to Limnodynastidae cation. Davies (2003a) noted that Limnodyn­ and Myobatrachidae of previous usage (Zug astidae are united by the character of fusion et al., 2001; Davies, 2003a, 2003b), differing of the first two vertebrae. Molecular evidence mildly only in the transfer of Mixophyes is decisive in support of this taxon; see apfrom Limnodynastidae to Myobatrachidae pendix 5 for diagnostic transformations. and the firm attachment of Rheobatrachus SYSTEMATIC COMMENTS : Our results sug­ (formerly Rheobatrachidae ) to Myobatrachi­ gest strongly that Limnodynastes as currently dae. See those accounts for further discus­ formulated is polyphyletic. Schäuble et al. sion. Haas (2003) suggested a number of (2000) provided a tree of species of Limnomorphological characters that optimize on dynastes which corresponds in some ways his terminal taxon, Limnodynastes peronii . with our results, but which differs in others. Because this was the only myobatrachoid in Their maximum­likelihood results, based on

that study, all of these characters might be 450 bp of 16S mtDNA and 370 bp of ND4, synapomorphies of various monophyletic suggest that Adelotus sits within the Limnogroups within this taxon. Hypothesized mo­ dynastes ornatus group ( L. ornatus and L.

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 195

spenceri ), and that this overall group forms Uperoliidae Günther, 1858b: 346. Type genus: the sister taxon of the remaing Limnodynas­ Uperoleia Gray, 1841 .

tes in the arrangement Limnodynastes dor­ Criniae Cope, 1866: 89. Type genus: Crinia salis group 1 ( L. peronii group 1 L. salmini Tschudi, 1838.

Rheobatrachinae Heyer and Liem, 1976: 11. Type group). Our results, based on denser taxon

genus: Rheobatrachus Liem, 1973 .

sampling and substantially more data, place

Adelotus outside of Limnodynastes (sensu IMMEDIATELY MORE INCLUSIVE TAXON: lato), but place Neobatrachus , Notaden , [321] Myobatrachoidea Schlegel, 1850.

Lechriodus , and Heleioporus within a para­ SISTER TAXON: [322] Limnodynastidae phyletic Limnodynastes , or, alternatively, Lynch , 1971.

place Limnodynastes ornatus as the sister RANGE: Australia and New Guinea.

taxon of Lechriodus fletcheri , and far away CONTENT: Arenophryne Tyler, 1976 ; Assa from Limnodynastes (including Megistolotis Tyler, 1972 ; Crinia Tschudi, 1838 ; Geocrias a synonym as suggested by Schäuble et nia Blake, 1973; Metacrinia Parker, 1940 ; al., 2000), which is the sister taxon of Hel­ Mixophyes Günther, 1864 ; Myobatrachus eioporus . In order to alleviate the polyphyly Schlegel, 1850; Paracrinia Heyer and Liem , of Limnodynastes , we resurrect the name Op­ 1976; Pseudophryne Fitzinger, 1843 ; Rheoisthodon Steindachner, 1867 (type species: batrachus Liem, 1973; Spicospina Roberts , Opisthodon frauenfeldi Steindachner, 1867 , Horwitz, Wardell­Johnson, Maxson, and Maby monotypy [5 Discoglossus ornatus Gray , hony, 1997; Taudactylus Straughan and Lee, 1842 ]) for the former Limnodynastes ornatus 1966; Uperoleia Gray, 1841 .

group (i.e., Opisthodon ornatus [Gray, 1842] CHARACTERIZATION AND DIAGNOSIS: Myob­ and O. spenceri [Parker, 1940]). This renders atrachids are predominantly small frogs of Opisthodon as the sister taxon of Lechriodus , heterogeneous appearance. All are assumed and Limnodynastes as the sister taxon of Hel­ to have inguinal amplexus, except Mixophyes eioporus , assuming that both Opisthodon and (which has axillary amplexus). Davies Lechriodus are monophyletic. We suggest (2003b) noted that although Mixophyes had that the molecular characters that optimize on traditionally been associated with Limnodynthe branch labeled Limnodynastes ornatus astidae, its placement there was always probare synapomorphies of Opisthodon (appen­ lematic due to its lack of most limnodynasdix 5). See appendix 7 for new combinations tinae characteristics. All myobatrachids are produced by this generic change. assumed to have a typical biphasic life his­ J.D. Lynch (1971: 76) distinguished two tory (Martin, 1967). None of our morphologtribes within his Cycloraninae (equivalent to

ical characters optimize on this branch be­

cause no member of this taxon was studied our Limnodynastinae with the removal of

to Pelodryadinae ): Cycloranini

by Haas (2003), although Ford and Canna­ Cyclorana tella (1993) suggested the following to be a ( Cyclorana , Heleioporus , Mixophyes , Neo­

likely synapomorphy: (1) broad alary process batrachus, and Notaden ), characterized by

of premaxilla (absent in Mixophyes , but also laying eggs in dry burrows in a foam nest,

present in the leptodactylids Adenomera , and Limnodynastini ( Adelotus , Lechriodus ,

Pseudopaludicola , and Physalaemus [in the Limnodynastes , and Philoria ), which lay

sense of including Engystomops and Eupemtheir eggs in water or in moist terrestrial

phix]). In our topology, this character could sites. When these characteristics are opti­

be reversed in or convergent in

Mixophyes

mized on our cladogram, they provide a rath­

Rheobatrachus and the clade bracketed by er confusing picture of life history evolution

Taudactylus and Arenophryne . Regardless, in limnodynastine frogs, and our data do not the molecular evidence appears to be decisupport recognition of these taxa. sive (see appendix 5 for molecular synapo­

morphies for branch 334).

[334] FAMILY: MYOBATRACHIDAE SCHLEGEL, SYSTEMATIC COMMENT : We expected

1850

as