Poyntonophrynus, Frost & Grant & Faivovich & Bain & Haas & De Sá & Channing & Wilkinson & Donnellan & Raxworthy & Campbell & Blotto & Moler & Drewes & Nussbaum & Lynch & Green & Wheeler, 2006

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

Poyntonophrynus for the Bufo vertebralis ly of this taxon is a testable proposition.

group of Tandy and Keith (1972; cf. Poyn­ Other than the Bufo pardalis group (see ton, 1964) and Cunningham and Cherry above), we have no unambiguous evidence

222 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

tying the African 22­chromosome toad taxon was considered to comprise a number groups ( B. gracilipes and B. mauritanicus of casually­defined species groups, most of groups) or such African species of unknown which require reevaluation. Although Tschukaryotype such as the B. pentoni group and di (1845) provided an erroneous South B. arabicus group to any of the African (or American type locality for the type species, other) bufonid groups. Additional evidence it was recognized as early as 1882 (Boulen­ and study will be needed to resolve their ger, 1882) that Anaxyrus melancholicus placement, which very clearly is not within Tschudi, 1845, is a junior synonym of the Bufo (sensu stricto). For the moment, we Mexican Bufo compactilis Wiegmann, 1834 . merely place the generic name ‘‘ Bufo ’’ in This was most recently detailed by Pramuk quotation marks in combination with these and Mendelson (2003). (See appendix 7 for species to denote their formal exclusion from content and new and revived combinations.) Bufo (sensu stricto). A partial junior synonym of Anaxyrus is

(14) Nannophryne Günther, 1870 (type Incilius Cope (1863: 50) . Under the provispecies: Nannophryne variegata Günther , sions of the ‘‘Principle of First Revisor’’ 1870, by monotypy). We resurrect the name (Art. 24; ICZN, 1999) we designate Bufo Nannophryne for Bufo variegatus (Günther, cognatus Say, 1823 , as the type species of 1870). Although we did not include this tax­ Incilius to solidify this synonymy, which othon in our analysis, the molecular evidence erwise could have been assigned through one provided by Pauly et al. (2004) suggests of the originally included species to threaten strongly that this taxon, like Rhinella (the the priority of Cranopsis . Bufo margaritifer group), is only distantly re­ (16) [519] Cranopsis Cope, 1875 ‘‘1876’’ lated to other New World ‘‘ Bufo ’’. Martin (type species: Bufo fastiodosus Cope, 1875 (1972) provided osteological differentia that ‘‘1876’’). We apply the name Cranopsis to serve to diagnose the taxon among ‘‘ Bufo ’’, the predominantly Middle American taxon but its exact phylogenetic position among subtended by branch 519. Although we know bufonids remains to be determined. Prior to of no morphological synapomorphy for this Pauly et al. (2004), some authors placed B. taxon, species within it generally exhibit a variegatus near the B. spinulosus group (e.g., distinctive appearance. Nevertheless, see ap­ Blair, 1972c), whereas others (e.g., Cei, pendix 5 for molecular synapomorphies. This 1980) have declined to place it in any species group is composed of the former Bufo valgroup . Pauly et al. (2004) placed it far from liceps group and allies. See appendix 7 for the B. spinulosus group, and attaching near content and new and revived combinations. the base of the bufonid exemplars that they (17) [522] Chaunus Wagler, 1828 (type studied. It remains possible that Nannophry­ species: Chaunus marmoratus Wagler, 1828 ne will be found to be most closely related [5 Bufo granulosus Spix, 1824 ]). We recto Rhaebo , in which case Rhaebo will take ognize the predominantly South American nomenclatural precedence for the larger taxon subtended by branch 522 as Chaunus . group. No morphological characters are known to

(15) [513] Anaxyrus Tschudi, 1845 (type diagnose this group, which is diagnosed species: Anaxyrus melancholicus Tschudi , completely on the basis of molecular data 1845 [5 Bufo compactilis Wiegmann , (see appendix 5, branch 522). Rhamphophry­ 1833]). We recognize the North American ne and Rhinella may well be found to be clade of ‘‘ Bufo ’’ subtended by branch 513 nested within Chaunus (see Graybeal, 1997: (see appendix 5) as the genus Anaxyrus her fig. 13; Pauly et al., 2004), in which case, Tschudi, 1845. We are unaware of any mor­ Rhinella Fitzinger, 1826 , will take precephological synapomorphy for this group, al­ dence, but evidence has yet to be produced though, with exceptions, they do have a dif­ to support this synonymy without recourse to ferent look and feel than the predominantly accepting a specific model of molecular evo­ Middle­American ( Cranopsis ) and South­ lution (Pauly et al., 2004).

American ( Chaunus ) taxa. Recognition of Pauly et al. (2004) suggested on the basis this taxon is consistent with our results and of fewer data, more analytical assumptions, those of Pauly et al. (2004). Formerly, this but denser sampling that the Bufo margari­

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 223

tifer group (see below) is imbedded within tus. But, because we did not study that spethis group. This remains an open question, cies, and because its sole reason for being but we suggest that decisive resolution will placed outside of Nectophrynoides is its loss require denser taxon sampling and more data, of columella, a character strongly contingent not additional analytical assumptions. on immediate outgroups, we refrain from this

There are several other groups of ‘‘ Bufo ’’ action until the appropriate phylogenetic and various individual species we have not comparisons can be made. addressed because we did not include any of In addition, the following monotypic genthem in our analysis and because there is no era have been named since the publication of substantial published evidence on their phy­ Graybeal and Cannatella (1995) and Graylogenetic placement. All of these we simply beal (1997): Churamiti Channing and Stantreat as incertae sedis within Bufonidae , ley, 2002, and Parapelophryne Fei, Ye , and tacked to the generic label ‘‘ Bufo ’’ (see ap­ Jiang, 2003. Neither obviously renders any pendix 7 for a list). The reader will note that other taxon paraphyletic. Clearly, a detailed the bulk are Asian taxa, residing in geo­ revision of Bufonidae without reference to graphic areas suggesting that they will be geographic boundaries is badly needed. found to be related to a number of non­ Bufo genera. Only additional work will elucidate [108] RANOIDES NEW TAXON this.

We think that our proposed breakup of ETYMOLOGY: Rana (Latin: frog) 1 oides ‘‘ Bufo ’’ will promote more rapid progress in (Greek: having the form of). The taxon is the field, because the sociological principle identical in content to the regulated superthat drives much of systematics is to show family name Ranoidea , but with an ending that other workers are wrong (Hull, 1988), change made to remove the implication that and many graduate students will certainly it is regulated by the International Code of take aim at our hypotheses. Most systema­ Zoological Nomenclature (ICZN, 1999). tists recognize that, traditionally, the first (See nomenclatural comment under Ranoides species to receive novel generic names have in appendix 6.) been those that are highly autapomorphic, IMMEDIATELY MORE INCLUSIVE TAXON: and subsequent authors are usually hesitant [107] Phthanobatrachia new taxon. to apply these names to more generalized SISTER TAXON: [314] Hyloides new taxon. forms. Having taken the controversial first RANGE: Worldwide temperate and tropical step, we hope that other workers will step in regions, except New Zealand, most of Aus­ and address the rather large number of prob­ tralia, and southern South America. lems that we have identified. There is much CONCEPT AND CONTENT : Ranoides new taxwork to be done in bufonids, and we intend on is a monophyletic group composed of our taxonomic proposal to serve as a frame­ [109] Allodapanura new taxon and [180] work that will guide additional studies. Natatanura new taxon.

We do not find any compelling reason to CHARACTERIZATION AND DIAGNOSIS: Haas maintain the sister monotypic genera Alti­ (2003) suggested the following characters phrynoides Dubois, 1987 ‘‘1986’’ and Spi­ that we regard as synapomorphies of our nophrynoides Dubois, 1987 ‘‘1986’’. Gran­ Ranoides: (1) insertion of m. rectus cervicis dison (1981) and Graybeal and Cannatella on proximal ceratobranchialia III and IV (1995) showed these African toads to be each (Haas 39.2); (2) ramus mandibularis (cranial other’s closest relatives. Acting as First Re­ nerve V 3) is either posterior (ventral) to m. visor, we consider Altiphrynoides Dubois , levator mandibulae externus group or runs 1987 ‘‘1986’’, to be a senior synonym of Spi­ through it—a change from being anterior nophrynoides Dubois, 1987 ‘‘1986’’. (See (dorsal) to the externus group (Haas 65.0/1); appendix 7 for the single new combination.) and (3) firmisternal shoulder girdle (epicor­ ‘‘ Nectophrynoides ’’ cryptus in their tree (fig. acoids are fully fused along their length;). osof

26) is not part of a monophyletic group with Haas 144.2; convergent in Dendrobatidae other Nectophrynoides . We were tempted to J.D. Lynch (1973: 146) suggested that an name a new genus for Nectophrynoides cryp­ sified omosternum is a synapomorphy 224 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

‘‘Ranoidea’’ (our Ranoides, excluding Mi­ Liem, 1970). In addition, Tyler (1971a) sugcrohylidae and Brevicipitidae ). This may be, gested that the presence of the m. cutaneous but there are alternative optimizations. pectoris could be a synapomorphy of Rano­ Among others, the ossified omosternum may ides, although with several reversals. have been gained at the level of Ranoides and lost independently in Microhylidae and [109] ALLODAPANURA NEW TAXON Brevicipitidae ; gained at the level of Rano­ ETYMOLOGY: Allodapos­ (Greek: strange, ides, lost at Allodapanura, and regained at foreign, or belonging to another kind) 1 an­ Laurentobatrachia; or gained independently oura (Greek: without a tail, i.e., frog), referin Laurentobatrachia, Natatanura, and Hem­ encing the exotic diversity of morphotypes isotidae. (See also appendix 5, branch 108, in this taxon. for molecular synapomorphies.) IMMEDIATELY MORE INCLUSIVE TAXON: SYSTEMATIC COMMENTS: Ranoides in our [108] Ranoides new taxon. sense is coextensive with the Recent content SISTER TAXON: [180] Natatanura new taxof the superfamily Ranoidea Rafinesque , on. 1814, of Dubois (2005). RANGE: North and South America; sub­Sa­ A preliminary survey of literature (Liem, haran Africa; India and Korea to northern 1970; Tyler, 1972, 1982; Burton, 1986, Australia. 1998b) as well as examination of a few ex­ CONCEPT AND CONTENT : Allodapanura new emplars of selected genera of several families taxon is a monophyletic group composed of suggests another likely synapomorphy of [110] Microhylidae Günther, 1858 (1843), Ranoides, worthy of additional investigation. and [143] Afrobatrachia new taxon. Anteromedially differentiated elements of CHARACTERIZATION AND DIAGNOSIS: Morthe m. intermandibularis are present in Ar­ phological characters in our analysis (from throleptidae, Brevicipitidae , Cacosterninae Haas, 2003 ) that are synapomorphies are (1) ( Pyxicephalidae ), Ceratobatrachidae , Hemi­ m. tympanopharyngeus present (Haas 20.1); sotidae, Hyperoliidae , Microhylidae , Pty­ and (2) arcus subocularis round in cross secchadenidae (however, absent in Hildebrand­ tion (Haas 82.2). In addition, absence of the tia), Petropedetidae , Phrynobatrachidae , and palatine bone in adults (Haas 146.0; a reverare absent in Alytidae , Batrachophrynidae sal from the acosmanuran condition), may (although present in Batrachophrynus brach­ optimize on this branch (to reappear on the ydactylus), Bombinatoridae , Heleophrynidae , branch subtending Afrobatrachia), or, alter­ Limnodynastidae , Myobatrachidae , Peloba­ natively, the palatine may be lost in Microtidae , Sooglossidae , and Hemiphractidae hylidae and independently in Xenosyneuni­ (Beddard, 1908 ‘‘1907’’, 1911; Tyler, 1972; tanura. Similarly, the presence of palatal Tyler and Duellman, 1995; Burton, 1998b). folds may optimize on this branch and be This taxonomic distribution suggest that the reversed in Laurentobatrachia, or may appear presence of differentiated elements of the m. twice, once on the branch subtending Microintermandibularis is a synapomorphy of Ran­ hylidae as well as on the branch subtending oides. Many details about the morphological Xenosyneunitanura. Regardless, the primary diversity and taxonomic distribution of this evidence for the recognition of this taxon is character remain unknown and several in­ molecular (see appendix 5). stances of homoplasy are known within Hyloides (see Tyler, 1971b, 1971c, 1972; [110] FAMILY: MICROHYLIDAE GÜNTHER, 1858 Burton, 1998b, and Tyler and Duellman, (1843) 1995, for examples within Noblebatrachia), and there are possibly multiple subsequent Hylaedactyli Fitzinger, 1843: 33. Type genus: Hy­

laedactylus Duméril anbd Bibron, 1841. transformations within Natatanura. (This Gastrophrynae Fitzinger, 1843: 33. Type genus: character does not seem to occur in at least Gastrophryne Fitzinger, 1843 . some Dicroglossidae [exemplars of Occidoz­ Micrhylidae Günther, 1858b: 346. Type genus: yga, Euphlyctis , Nannophrys ] or Nyctiba­ Micrhyla Duméril and Bibron, 1841 (an incortrachidae [ Lankanectes , Nyctibatrachus ], but rect subsequent spelling of Microhyla Tschudi , is present in Mantellidae and Rhacophoridae ; 1838).

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 225

Asterophrydidae Günther, 1858b: 346 . Type ge­ 1986; Altigius Wild, 1995 ; Arcovomer Car­

nus: Asterophrys Tschudi, 1838 . valho, 1954; Chiasmocleis Méhely, 1904 ;

Kalophrynina Mivart, 1869: 289. Type genus: Gastrophrynoides Noble, 1926 ; Glyphoglos­

Kalophrynus Tschudi, 1838 . sus Günther, 1869 ‘‘1868’’; Hyophryne Car­

Xenorhinidae Mivart, 1869: 286. Type genus: Xe­ valho, 1954; Keferstein, 1867; Hypopachus

norhina Peters, 1863.

Kalophrynus Tschudi, 1838 ; Metaphrynella

Dyscophidae Boulenger, 1882: 179 . Type genus:

Dyscophus Grandidier, 1872 . Parker, 1934; Micryletta Dubois, 1987 ;

Cophylidae Cope, 1889: 248. Type genus: Cophy­ Myersiella Carvalho, 1954 ; Otophryne Bou­

la Boettger, 1880. lenger, 1900; Paradoxophyla Blommers­

Genyophrynidae Boulenger, 1890: 326. Type ge­ Schlösser and Blanc, 1991; Phrynella Bou­

nus: Genyophryne Boulenger, 1890 . lenger, 1887; Phrynomantis Peters , 186731;

Rhombophryninae Noble, 1931: 529. Type genus: Ramanella Rao and Ramanna, 1925 ; Relic­

Rhombophryne Boettger, 1880 . tivomer Carvalho, 1954; Stereocyclops Cope ,

Sphenophryninae Noble, 1931: 531. Type genus: 1870 ‘‘1869’’; Synapturanus Carvalho ,

Sphenophryne Peters and Doria, 1878 , by mon­ 1954; Syncope Walker, 1973 ; Uperodon Du­

otypy. méril and Bibron, 1841. (See Systematic

Melanobatrachinae Noble, 1931: 538 . Type ge­

Comments.)

nus: Melanobatrachus Beddome, 1878 .

Kaloulinae Noble, 1931: 538. Type genus: Kal­ CHARACTERIZATION AND DIAGNOSIS: A large

oula Gray, 1831. number of morphological characters in our

Hoplophryninae Noble, 1931: 538–539 . Type ge­ analysis (from Haas, 2003) are synapomor­

nus: Hoplophryne Barbour and Loveridge , phies of Microhylidae : (1) keratodonts absent

1928. in larvae (Haas 3.0); (2) keratinized jaw

Scaphiophryninae Laurent, 1946: 337 . Type ge­ sheaths absent in larvae (Haas 6.0); (3) vena

nus: Scaphiophryne Boulenger, 1882 . caudalis dorsalis present in larvae (Haas

Pseudohemisiinae Tamarunov, 1964a: 132. Type 14.1); (4) spiracle position median posterior

genus: Pseudohemisus Mocquard, 1895 . (Haas 18.2); (5) m. geniohyoideus origin in

Otophryninae Wassersug and Pyburn, 1987: 166 . larvae from connective tissue lateral to glan­

Type genus: Otophryne Boulenger, 1900 .

dula thyroidea (Haas 19.4); (6) m. interhyoi­

Phrynomantini Burton, 1986: 405–450. Type ge­

nus: ‘‘ Phrynomantis Peters, 1867 ’’.

deus posterior in larvae extensive and strong­

Barygenini Burton, 1986: 405–450. Type genus: ly developed (Haas 24.2); (7) m. diaphrag­

Barygenys Parker, 1936 . matopraecordialis absent in larvae (Haas

Callulopini Dubois, 1988a: 3. Type genus: Cal­ 25.0); (8) lateral fibers of m. subarcualis rec­

lulops Boulenger, 1888. tus II–IV invade interbranchial septum IV musculature in larvae (Haas 29.1); (9) m. su­

IMMEDIATELY MORE INCLUSIVE TAXON: barcualis rectus II–IV split into medial and

[109] Allodapanura new taxon. lateral separate muscles (Haas 30.1); (10) m.

SISTER TAXON: [143] Afrobatrachia new subarcualis rectus I portion with origin from

taxon. ceratobranchial III absent (Haas 35.0); (11)

RANGE: North and South America; East ventral portion of the m. subarcualis rectus I

and South Africa; India and Korea to north­ inserts laterally on ceratohyal (Haas 36.1);

ern Australia. (12) origin of m. suspensoriohyoideus from

CONTENT: [135] Asterophryninae Günther, posterior palatoquadrate (Haas 46.1); (13) m.

1858 (including Genyophryninae Boulenger , interhyoideus and m. intermandibularis in

1890), [118] Cophylinae Cope, 1889 , Dys­ close proximity (Haas 47.0); (14) m. man­

cophinae Boulenger, 1882, [121] Gastro­ dibulolabialis inserting exclusively on carti­

phryninae Fitzinger, 1843, [130] Microhyli­ lago labialis inferior (Haas 49.1); (15) m. le­

nae Günther, 1858 (1843), Scaphiophryninae vator mandibulae internus anterior (Haas

Laurent, 1946, as well as several nominal

genera unassigned to subfamily either be­ 31 We realize, of course, that Phrynomantis Peters ,

cause we did not study them and assignment 1867, is the sole member of Phrynomerinae Noble, 1931 . But, beyond the autapomorphic intercalary phaits to subfamily based on published evidence is langeal elements, we have only weak evidence for not possible, or because they fall outside of placement. In this case, recognition of a monotypic subexisting subfamilies: Adelastes Zweifel , family serves no purpose.

226 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

58.2); (16) m. levator mandibulae longus monophyletic taxonomy we propose the foloriginates exclusively from arcus subocularis lowing taxonomic changes: (1) place Aster­ (Haas 60.2); (17) profundus and superficialis ophryinae and Genyophryninae in one subportions of m. levator mandibulae longus not family, Asterophryinae (following Savage, overlapping and parallel (Haas 62.1); (18) ra­ 1973); (2) restrict Dyscophinae to Dyscophus mus mandibularis (cranial nerve V 3) between (also following Savage, 1973) and transfer portions of m. levator mandibulae longus Calluella from Dyscophinae to Microhylimuscle (Haas 64.1); (19) processus suboticus nae; (3) retain Cophylinae , but note that it quadrati present (Haas 76.1); (20) partes cor­ appears to be imbedded within a cluster of pores forming medial body (Haas 87.2); (21) ‘‘microhyline’’ genera that, once their phydistal end of cartilago meckeli expanded and logeny is better resolved, may require some flattened with no fossa (Haas 94.2); (22) hy­ reconstitution of Cophylinae ; and (4) partipobranchial plates fused (Haas 107.1); (23) tion Microhylinae into a New World group, commissura proximalis I present (Haas Gastrophryninae , and an Old World group, 109.1); (24) processus branchialis closed Microhylinae , with several genera left incer­ (Haas 114.1); (25) accessory longitudinal tae sedis until they can be adequately studied bars of cartilage dorsal to ceratobranchialia or placed in a more densely sampled frame­ II and III present (Haas 120.1); (26) posterior work. Another group of genera (i.e., Kalomargin of ventral velum discontinuous (Haas phrynus, Synapturanus , Phrynomantis , Mi­ 129.1); (27) glottis position posterior (Haas cryletta) is left incertae sedis, as well, al­ 130.1); (28) nostrils closed in larval stages though the phylogenetic structure we ob­ (Haas 131.1); (29) branchial food traps di­ tained among these taxa is instructive and vided and crescentic (Haas 135.1); and (30) points to new questions for systematists to eggs floating (Haas 141.2). Although most of address. Nevertheless, our obtained structure these characters will survive denser taxon suggests that the biogeography of Microhysampling, the placement of some of them is lidae is complex and old. currently ambiguous inasmuch as some of Our data show that the former ‘‘Microthe characters listed could actually be sitting hylinae’’ (sensu lato) is heterogenous mixon branches from which Synapturanus and ture of basal taxa (e.g., Synapturanus , Mi­ Kalophrynus are derived. cryletta) and two distantly related clades with

Presence of palatal folds is optimization­ which we have associated the names Microdependent. Presence of palatal folds may be hylinae (Asia) and Gastrophryninae (Americonvergent in Microhylidae and Xenosyneu­ cas). There is no published evidence that nitanura, or a synapomorphy of Allodapan­ would allow us to allocate any of the unura and lost in Laurentobatrachia. studied Asian taxa to Microhylinae or to any

SYSTEMATIC COMMENTS: The obtained phy­ other position in our cladogram beyond their logenetic structure of Microhylidae surprised being microhylids. Similarly, although we us as we expected Scaphiophryninae to form assume that such taxa as Hypopachus are in the sister taxon of the remaining microhylids, Gastrophryninae , our molecular results are so because the scaphiophrynine tadpole mor­ incongruent with results from morphology phology (Blommers­Schlösser, 1975; Haas, (e.g., Zweifel, 1986; Donnelly et al., 1990; 2003), is annectant in many ways between Wild, 1995) that we hesitate to conjecture. the ranid and more typical microhylid con­ Morphological characters that are candidition. As in several other parts of the tree, dates as synapomorphies of [134] Dyscophithe density of our taxon sampling was inad­ nae 1 Asterophryninae 1 Scaphiophryninae equate to address all problems in microhylid 1 Microhylinae clade are (1) double­layered systematics, and we intend our results to dermis (Haas 13.1, also in Hemisus and Kasguide more thorough studies. Rafael de Sá sina); (2) anterior insertion of m. subarcualis and collaborators have begun such a study, rectus II–IV on ceratobranchial I (Haas and we anticipate further revision of micro­ 37.0); and (3) partes corpores forming medial

hylid systematics as a result. For this reason body (Haas 87.2). we leave several taxa unnamed and unad­ Because the nominal subfamilies of Midressed. As an initial step toward an entirely crohylidae are large and morphologically dis­

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 227

parate, we include separate accounts for the place completely within the egg capsule, alnominal subfamilies. though others (e.g., Cophylinae , some Gas­

trophryninae) are endotrophic and nidicolous

[135] SUBFAMILY: ASTEROPHRYINAE (Blommers­Schlösser, 1975) . (See appendix

GÜNTHER, 1858 5 for molecular synapomorphies.) Asterophrydidae Günther, 1858b: 346 . Type ge­

SYSTEMATIC COMMENTS: Former Genynus: Asterophrys Tschudi, 1838 . ophryninae is paraphyletic with respect to Xenorhinidae Mivart, 1869: 286. Type genus: Xe­ the old Asterophryinae , and for this reason norhina Peters, 1863. the two nominal taxa were synonymized in Genyophrynidae Boulenger, 1890: 326. Type ge­ ‘‘Results’’. Parker (1934) noted Genyophrynus: Genyophryne Boulenger, 1890 . New syn­ ninae (as Sphenophryninae) to be procoelous onym. and Asterophryinae as diplasiocoelous, and Sphenophryninae Noble, 1931: 531. Type genus: this clearly influenced later authors (e.g., Sphenophryne Peters and Doria, 1878 , by mon­ Zweifel, 1972) in retaining a distinction beotypy. New synonym.

tween the nominal subfamilies. The place­ Phrynomantini Burton, 1986: 405–450. Type genus: ‘‘ Phrynomantis Peters, 1867 ’’.

ment in our tree of Australo­Papuan AsterBarygenini Burton, 1986: 405–450. Type genus: ophryinae (sensu lato) as the sister taxon of Barygenys Parker, 1936 . the Madagascan Dyscophinae is a remark­ Callulopini Dubois, 1988a: 3. Type genus: Cal­ able biogeographic signature. lulops Boulenger, 1888. Burton (1986: 443) provided evidence that

Xenorhina is paraphyletic with respect to Xe­ IMMEDIATELY MORE INCLUSIVE TAXON: nobatrachus, the latter differing only in lack­ [134] unnamed taxon. ing large odontoids on the vomeropalatine. SISTER TAXON: Dyscophinae Boulenger, Zweifel (1972) provided no evidence for the 1882. monophyly of Xenorhina . On the basis of RANGE: Southern Philippines, Sulawesi, these works we consider them to be syno­ and Lesser Sunda Islands and Moluccas east­ nyms, with Xenorhina being the older name wards through New Guinea and satellite is­ (see appendix 7 for new combinations). lands to Australia. Burton (1986: 443) also noted that ‘‘ Manto­ CONTENT: Albericus Burton and Zweifel , phryne ’’ and ‘‘ Hylophorbus ’’ are dubiously 1995; Aphantophryne Fry, 1917 ‘‘1916’’; monophyletic, so we place these names in Asterophrys Tschudi, 1838 ; Austrochaperina quotation marks until their monophyly can Fry, 1912; Barygenys Parker, 1936 ; Callu­ be substantiated. Although Burton (1986) lops Boulenger, 1888; Choerophryne Kam­ provided a number of morphological charpen, 1914; Cophixalus Boettger, 1892 ; Cop­ acters and a character matrix, no one so far iula Méhely, 1901; Genyophryne Boulenger , has analyzed these data phylogenetically. 1890; Hylophorbus Macleay, 1878 ; Liophryne Boulenger, 1897 ; Mantophryne Boulen­ [118] SUBFAMILY: COPHYLINAE COPE, 1889 ger, 1897; Oreophryne Boettger, 1895 ; Oxydactyla Kampen, 1913 ; Pherohapsis Zwei­ Cophylidae Cope, 1889: 248. Type genus: Cophy­

la Boettger, 1880. fel, 1972; Sphenophryne Peters and Doria ,

Rhombophryninae Noble, 1931: 529. Type genus: 1878; Xenorhina Peters, 1863 (including Xe­ Rhombophryne Boettger, 1880 . nobatrachus Peters and Doria, 1878; see appendix 7 for new combinations). IMMEDIATELY MORE INCLUSIVE TAXON: CHARACTERIZATION AND DIAGNOSIS: None [116] unnamed taxon. of the morphological characters in our anal­ SISTER TAXON: [117] An unnamed taxon in ysis apply to this taxon because as direct de­ our analysis composed of our exemplars Hovelopers they were not part of the tadpole plophryne Barbour and Loveridge, 1928 study by Haas (2003). Among microhylids, ( Melanobatrachinae Noble, 1931 ) and Raonly Asterophryinae and Myersiella (Micro­ manella Rao and Ramanna, 1925 (formerly the hylinae ; Izecksohn et al., 1971; Zweifel, of ‘‘Microhylinae’’). Together these are 1972; Thibaudeau and Altig, 1999) exhibit sister taxon of [121] Gastrophryninae Fitzindirect development, the development taking ger, 1843.

228 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

RANGE: Madagascar. phryne (see appendix 7 for the species name

CONTENT: Anodonthyla Müller, 1892 ; Co­ changes that this causes). Andreone et al. phyla Boettger, 1880; Madecassophryne Gui­ (2004 ‘‘2005’’) hesitated to take this step bebe´, 1974; Platypelis Boulenger, 1882 ; Pleth­ cause they did not feel there was sufficient odontohyla Boulenger, 1882 (see Systematic statistical support for their maximum­likeli­ Comments); Rhombophryne Boettger, 1880 hood conclusion. They did, however, note (see Systematic Comments and appendix 7); that their parsimony tree arrived at the same Stumpffia Boettger, 1881 . conclusion. We therefore think that it is bet­

CHARACTERIZATION AND DIAGNOSIS: None ter to recognize two clades that might be of the morphological characters in our anal­ found to be each other’s closest relatives ysis optimizes on this branch; because our when more data are added to the analysis, morphological characters were largely de­ than to retain a taxon, ‘‘ Plethodontohyla ’’ rived from larvae, and cophylines (as tradi­ (sensu lato) for which the preponderance of tionally defined) are endotrophic and nidic­ data does not support its monophyly. There olous. Nevertheless, endotrophy is a syna­ are a number of species, nominally in Plethpomorphy at this level. Also, cophylines odontohyla, but not treated by Andreone et have unfused sphenethmoids, which appear al. (2004 ‘‘2005’’). We retain those in Plethas paired elements (Parker, 1934), otherwise odontohyla, although some of may be found found convergently in Dyscophus (Dysco­ to be members of Rhombophryne . phinae) and Calluella ( Microhylinae ). (See appendix 5 for molecular synapomorphies on SUBFAMILY: DYSCOPHINAE BOULENGER, 1882 this branch [118].) Dyscophidae Boulenger, 1882: 179 . Type genus:

SYSTEMATIC COMMENTS: The association by Dyscophus Grandidier, 1872 . our molecular data of Cophylinae (Madagas­ IMMEDIATELY MORE INCLUSIVE TAXON: car) with our exemplars Hoplophryne (East [134] unnamed taxon. Africa) and Ramanella ( India) is suggestive. SISTER TAXON: [135] Asterophryinae Gün­ Madagascar – India is a repeated pattern in ther, 1858. biogeography, as is an apparently later con­ RANGE: Madagascar. nection of India –Africa (e.g., Chiromantis in CONTENT: Dyscophus Grandidier, 1872 . Africa 1 Chirixalus in Asia [Rhacophori­ CHARACTERIZATION AND DIAGNOSIS: Haas dae]; Petropedetes 1 Arthroleptides in Af­ (2003) suggested the following larval charrica and Indirana in India [ Petropedetidae ]). acters that are presumed synapomorphies of The association of Gastrophryninae with this the taxon: (1) ramus mandibularis (cranial overall clade also speaks to a standard bio­ nerve V 3) runs through the m. levator mangeographic pattern, that of South America– dibulae externus group (Haas 65.1); and (2) Madagascar. free basihyal absent (Haas 105.0).

Andreone et al. (2004 ‘‘2005’’) provided SYSTEMATIC COMMENT : Our data reject the considerable DNA sequence evidence that association of Calluella with Dyscophinae Plethodontohyla is polyphyletic (not para­ (Blommers­Schlösser, 1976), which instead phyletic as suggested in the original publi­ place Calluella deeply within Microhylinae . cation; see fig. 33). As noted by Andreone et This is not surprising, inasmuch as the only al. (2004 ‘‘2005’’) the name Plethodontohyla characteristics suggested to ally Calluella Boulenger, 1882 (type species: Callula no­ with Dyscophinae are apparent plesiomortosticta Gunther, 1877) adheres to his Pleth­ phies (e.g., presence of teeth, diplasiocoelous odontohyla Group 1. Their second group of vertebral column, large vomer). The molec­ ‘‘ Plethodontohyla ’’ falls into a monophyletic ular synapomorphies supporting a relationgroup with Rhombophryne testudo . Rhom­ ship of this taxon to Asterophryinae (branch bophryne Boettger, 1880, is substantially old­ 134, appendix 5) is novel. er than the next older name for this taxon, Mantiphrys Mocquard, 1901 (type species: [121] SUBFAMILY: GASTROPHRYNINAE Mantiphrys laevipes Mocquard, 1895 ), and FITZINGER, 1843 to provide a monophyletic taxonomy, this in­ Gastrophrynae Fitzinger, 1843: 33. Type genus: clusive taxon should be known as Rhombo­ Gastrophryne Fitzinger, 1843 .

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 229

IMMEDIATELY MORE INCLUSIVE TAXON: SUBFAMILY: MELANOBATRACHINAE NOBLE , [115] unnamed taxon. 1931 SISTER TAXON: [116] unnamed taxon. Melanobatrachinae Noble, 1931: 538 . Type ge­ RANGE: Southern United States south to nus: Melanobatrachus Beddome, 1878 . Argentina. Hoplophryninae Noble, 1931: 538–539 . Type ge­ CONTENT: Ctenophryne Mocquard, 1904 ; nus: Hoplophryne Barbour and Loveridge , Dasypops Miranda­Ribeiro, 1924 ; Derma­ 1928. tonotus Méhely, 1904; Elachistocleis Parker, IMMEDIATELY MORE INCLUSIVE TAXON: 1927; Gastrophryne Fitzinger, 1843 ; Hamp­ [117] unnamed taxon. tophryne Carvalho, 1954; Nelsonophryne SISTER TAXON: Ramanella Rao and Ra­ Frost, 1987. manna, 1925. CHARACTERIZATION AND DIAGNOSIS: Opti­ RANGE: Montane Tanzania and southern mization is problematic because none of the India. direct­developing microhylids were sampled CONTENT: Hoplophryne Barbour and Lovin our morphological data set. Nevertheless eridge, 1928; Melanobatrachus Beddome , the following are candidates for being syna­ 1878; Parhoplophryne Barbour and Loverpomorphies of Gastrophryninae , although idge, 1928. they could be synapomorphies of Gastro­ CHARACTERIZATION AND DIAGNOSIS: Melanphryninae 1 Cophylinae or some subset of obatrachinae shares two synapomorphies Gastrophryninae inasmuch as the exemplars (Parker, 1934): (1) middle and outer ear abon which this supposition is built are Gas­ sent; (2) parasphenoid and sphenethmoid trophryne carolinensis, Hamptophryne boli­ fused. viana, and Elachistocleis ovalis ): (1) m. le­ SYSTEMATIC COMMENTS: Although we provator arcuum branchialium III split into two visionally retain Melanobatrachinae as an uncrossing bundles (Haas 41.1); (2) origin of tested taxon, the placement of Hoplophryne m. suspensoriohyoideus from otic capsule (our exemplar) in the general tree (see figs. (Haas 46.2); (3) posterolateral projections of 50 and 61) suggests that a more densely samthe crista parotica processus otobranchialis pled analysis will provide results that render

a Melanobatrachinae containing several more (Haas 67.2); (4) processus muscularis absent genera (such as Ramanella ) than as currently (Haas 79.0); (5) anterolateral base of proces­ composed. Hoplophryne and Parhoplophry­ sus muscularis bearing ventrolateral process ne were placed in Hoplophrynine by Noble (Haas 80.1); and (6) ligamentum mandibu­ (1931) on the basis of sharing the apomorphy losuprarostrale absent (Haas 127.0). of a greatly reduced first finger. (Noble also Molecular evidence (branch 121, appendix allied these genera with Brevicipitidae on the 5) is strong that the New World microhylids basis of retaining a complete clavicle, but (with the exception of Synapturanus , and this alliance is not supported by our data.) possibly several others for which we had no Parker (1934) placed Hoplophryninae in the tissues) form a clade that is most closely re­ synonymy of Melanobatrachinae ( India) be­ lated to the Madagascan Cophylinae . cause they share the absence of the auditory SYSTEMATIC COMMENTS: The exclusion of apparatus and fusion of the parasphenoid to Synapturanus from this taxon comes as the sphenenthmoid. We could not sample something of a surprise, inasmuch as both Melanobatrachus , but it remains possible Zweifel (1986) and Wild (1995) provided ev­ that it is the sister taxon of Hoplophryninae idence for its placement within a New World and that Hoplophryne and Parhoplophryne clade. Nevertheless, neither Zweifel (1986) are African outliers of a predominantly Innor Wild (1995) presented morphological ev­ dian group. This is conjecture, however, and idence for the monophyly of the New World only more data and denser sampling will remicrohylids (of which our Gastrophryninae solve the issue. is a part). We expect that further research will show the New World microhylids to be a [130] SUBFAMILY: MICROHYLINAE GÜNTHER , composite of gastrophrynines, some basal 1858 (1843) taxa (e.g., Synapturanus ), and possibly some Hylaedactyli Fitzinger, 1843: 33. Type genus: Hywith relations in Asia. laedactylus Duméril anbd Bibron, 1841.

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Micrhylidae Günther, 1858b: 346. Type genus: these are highly contingent on topological Micrhyla Duméril and Bibron, 1841 (an incor­ position of Scaphiophryne : (1) keratinized rect subsequent spelling of Microhyla Tschudi , jaw sheaths present (Haas 6.1; reversal from 1838). the microhylid condition); (2) eye position Kaloulinae Noble, 1931: 538. Type genus: Kal­ dorsolateral (Haas 11.0; reversal from the oula Gray, 1831. microhylid condition); (3) spiracle position IMMEDIATELY MORE INCLUSIVE TAXON: sinistral (Haas 18.1; reversal from the micro­ [129] unnamed taxon. hylid condition); (4) m. interhyoideus pos­ SISTER TAXON: Scaphiophryninae Laurent , terior absent (Haas 23.0; reversal from the 1946. phthanobatrachian condition); (5) m. subar­ RANGE: India, China, Japan, and Korea to cualis rectus II–IV represented by a single the Philippines and Greater Sunda Islands. flat tract of fibers (Haas 30.0; reversal from CONTENT: Calluella Stoliczka, 1872 ; the microhylid condition); (6) insertion of m. Chaperina Mocquard, 1892 ; Kaloula Gray , rectus cervicis on proximal ceratobranchialia 1831; Microhyla Tschudi, 1838 . III and IV (Haas 39.2; reversal from micro­ CHARACTERIZATION AND DIAGNOSIS: Haas hylid condition); (7) m. interhyoideus and m. (2003) examined only Kaloula within this intermandibularis well separated by a gap clade, so this is our only morphological ex­ (Haas 47.1; reversal from the microhylid emplar for this subfamily, but the following condition); (8) m. mandibulolabialis inserting are candidates for being synapomorphies of in soft tissue of lip (Haas 49.0; reversal from the Microhylinae : (1) vena caudalis dorsalis microhylid condition); (9) m. levator manabsent (Haas 14.0); (2) origin of m. suspen­ dibulae internus low (Haas 58.1; reversal soriohyoideus from otic capsule (Haas 46.2); from microhylid condition); (10) m. levator and (3) posterolateral projections of the crista mandibulae longus originates from posterior parotica expansive flat chondrifications palatoquadrate (Haas 60.1; reversal from mi­ (Haas 67.2). Nevertheless, the molecular ev­ crohylid condition); (11) ramus mandibularis idence is decisive for the recognition of this (cranial nerve V 3) anterior (dorsal) to the m. taxon (see appendix 5). levator mandibulae longus (Haas 64.2); (12) COMMENT: See Microhylidae account for processus suboticus quadrati absent (Haas comment on East Asian ‘‘microhylines’’ ex­ 76.0; reversal from microhylid condition); cluded from this taxon because of lack of (13) arcus subocularis with irregular margin evidence to place them. (Haas 81.1; reversal of microhylid condi­

tion); (14) cartilaginous roofing of the cavum SUBFAMILY: SCAPHIOPHRYNINAE LAURENT , cranii absent (Haas 96.0; reversal of predom­

1946 inant microhylid condition); and (15) glottis

position posterior (Haas 130.0; reversal of Scaphiophryninae Laurent, 1946: 337 . Type ge­ microhylid condition). nus: Scaphiophryne Boulenger, 1882 . Pseudohemisiinae Tamarunov, 1964a: 132. Type SYSTEMATIC COMMENTS: Ford and Cannagenus: Pseudohemisus Mocquard, 1895 . tella (1993: 94–117), found no evidence for

the monophyly of this taxon. Haas (2003: 50) IMMEDIATELY MORE INCLUSIVE TAXON: suggested on the basis of tadpole morphol­ [129] unnamed taxon. ogy that Paradoxophyla is more closely re­ SISTER TAXON: [130] Microhylinae Gün­ lated to Phrynomantis than to the remaining ther, 1858 (1843). Scaphiophryninae , rendering the latter non­ RANGE: Madagascar. monophyletic. On that basis alone, because CONTENT: Scaphiophryne Boulenger , we did not have tissues of Paradoxophyla , 1882. we transfer Paradoxophyla from Scaphio­ CHARACTERIZATION AND DIAGNOSIS: In our phryninae to incertae sedis under Microhytopology Scaphiophryne is deeply imbedded lidae. The association (branch 129, appendix within Microhylidae , requiring a remarkable 5) of Madagascan Scaphiophryninae with

number of reversals. Nevertheless, we sug­ Microhylinae may suggest an Indian origin gest these reversals are likely synapomor­ of Microhylinae . (See Systematic Comment phies of the taxon, while noting that most of under Cophylinae .)

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 231

[143] AFROBATRACHIA NEW TAXON which at this position in the general clado­

gram is a synapomorphy. Breviceps and ETYMOLOGY: Afro­ (Latin: of Africa) 1 Hemisus also share a single median thyroid batrachos (Greek: frog), in reference to the gland (Blommers­Schlösser, 1993), so we predominantly African range of this taxon. presume that this, too, is a synapomorphy IMMEDIATELY MORE INCLUSIVE TAXON: joining the two taxa. Breviceps and Hemisus [109] Allodapanura new taxon. also exhibit nasal plugs (De Villiers, 1931) SISTER TAXON: [110] Microhylidae Gün­ which may be homologous. (See also ‘‘Char­ ther, 1858 (1843). acterization and Diagnosis’’ under Hemiso­ RANGE: Sub­Saharan Africa, Madagascar, tidae for other characters that may optimize and the Seychelles. on this taxon.) Molecular synapomorphies CONCEPT AND CONTENT : Afrobatrachia is a are provided in appendix 5. monophyletic taxon composed of [144] Xenosyneunitanura new taxon and [148] Laurentobatrachia new taxon. [145] FAMILY: BREVICIPITIDAE BONAPARTE,

1850 CHARACTERIZATION AND DIAGNOSIS: Likely candidates for being synapomorphies are the Brevicipitina Bonaparte, 1850: 1 p. Type genus: larval characters: (1) m. transversus ventralis Breviceps Merrem, 1820 . IV present (Haas 22.1); (2) posterolateral Engystomidae Bonaparte, 1850: 1 p. Type genus: projections of the crista parotica forming Engystoma Fitzinger, 1826 . processus otobranchialis (Haas 67.3); (3) I:

MMEDIATELY MORE INCLUSIVE TAXON processus ascendens thin (Haas 72.1); (4) [144] Xenosyneunitanura new taxon. dorsal connection from processus muscularis S: Hemisotidae Cope, 1867 .

ISTER TAXON to ‘‘high’’ commissura quadrato­orbitalis R: Sub­Saharan East Africa and

ANGE (Haas 78.3); and (5) anterolateral base of southern Africa, from Ethiopia south to An­ processus muscularis bearing ventrolateral gola and South Africa. process (Haas 80.1). See characterisation of C: Balebreviceps Largen and

ONTENT Allodapanura for additional discussion of Drewes, 1989; Breviceps Merrem, 1820 ; possible synapomorphies. Callulina Nieden, 1911 ‘‘1910’’; Probrevi­ COMMENT: Our Afrobatrachia is identical ceps Parker, 1931; Spelaeophryne Ahl, 1924 . in content to the enlarged Brevicipitidae of C: Parker

HARACTERIZATION AND DIAGNOSIS Dubois (2005). (1934) noted that brevicipitids lack ossified

sphenethmoids, which is clearly a synapo­ [144] XENOSYNEUNITANURA NEW TAXON morphy at this level. In addition, the loss of ETYMOLOGY: Xeno­ (Greek: strange) 1 the pterygoid, palatoquadrate, and m. opersyneunitos (Greek: bed sharer) 1 anoura cularis (De Villiers, 1931) are likely syna­ (Greek: frog). In other words, the name pomorphies for this group. The extremely means ‘‘strange bedfellows’’ inasmuch as short head and direct development exhibited Hemisotidae and Brevicipitidae , although by this taxon (Parker, 1934) are also synacladistic nearest relatives, are dissimilar ani­ pomorphies. mals. SYSTEMATIC COMMENT : Loader et al. (2004) IMMEDIATELY MORE INCLUSIVE TAXON: suggested a phylogeny of Breviceps (Spe­ [143] Afrobatrachia new taxon. laeophryne 1 ( Callulina 1 Probreviceps )); SISTER TAXON: [148] Laurentobatrachia they, like us, did not include Balebreviceps new taxon. in their analysis. On the basis of our larger RANGE: Sub­Ssaharan Africa. amount of evidence but less dense sampling, CONCEPT AND CONTENT : Xenosyneunitanu­ we placed Probreviceps nearer to Breviceps ra new taxon is a monophyletic taxon con­ in our tree. Nevertheless, both arrangements taining Hemisotidae Cope, 1867 , and [145] conflict with the character of fusion of the Brevicipitidae Bonaparte, 1850 . urostyle and sacrum found in Probreviceps

ad­

CHARACTERIZATION AND DIAGNOSIS: Hemi­ and Breviceps but not in Spelaeophryne and sotidae and Brevicipitidae share the absence Callulina (Parker, 1934) , suggesting that of the palatine bones (De Villiers, 1931), ditional testing is warranted.

232 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

FAMILY: HEMISOTIDAE COPE, 1867 Hyperoliidae Laurent, 1943 , and [164] Ar­ Hemisidae Cope, 1867: 198 . Type genus: Hemisus throleptidae Mivart, 1869.

Günther, 1859 ‘‘1858’’. Emended to Hemisotina CHARACTERIZATION AND DIAGNOSIS: The by Günther, 1870: 119. characters (from Haas, 2003) 54.1 (larval m. levator manidbulae externus in two portion), IMMEDIATELY MORE INCLUSIVE TAXON: 111.0 (commissura proximalis III absent), [144] Xenosyneunitanura new taxon. and 151.0 (intercalary elements absent) are SISTER TAXON: [145] Brevicipitidae Bona­ likely synapomorphies of this group, alparte, 1850. though because of the low density of taxon RANGE: Sub­Saharan Africa. sampling this requires additional specimen CONTENT: Hemisus Günther, 1859 examination. In addition, claw­shaped ter­ ‘‘1858’’. minal phalanges appear to optimize on this CHARACTERIZATION AND DIAGNOSIS: All of branch, appearing convergently in Ptychadthe characters in our analysis (from Haas, ena and several of the hyloids (Liem, 1970), 2003) that optimize on Hemisus (our only although the distribution of this character is morphological exemplar in this clade) may complicated, and further work may show that be synapomorphies of this clade, the Hemi­ this optimization is mistaken. Drewes (1984) sotidae, or some subset of Hemisus : (1) dou­ suggested that thyrohyals borne on cartilagble­layered dermis in larvae (Haas 13.1); (2) inous stalks (his character 10.1) might be a posterior dorsal process of pars alaris ex­ synapomorphy, although this is optimizationpanded terminally, almost rectangular in lat­ dependent inasmuch as this character is not eral view (Haas 89.1); (3) larvae are guided in Leptopelis (Laurent, 1978) . The external by the female from the nest to pond (Haas metatarsal tubercle is absent or poorly de­ 137.1); and (4) amplexus absent (Haas veloped throughout Laurentobatrachia (Lau­ 139.0). Some of these may be synapomor­ rent, 1986), but the exact distribution of this phies at the level of Xenosyneunitanura in­ requires verification. Molecular synapomorasmuch as Brevicipitidae was not studied by phies for this taxon are summarized in ap­ Haas (2003) because they lack exotrophic pendix 5.

larvae, which were the focus of Haas’ study. SYSTEMATIC COMMENT : Vences and Glaw Hemisus lacks vomers, middle ear, and (2001) and Van der Meijden et al. (2005) recductus lacrimosus, and exhibits fusion of ver­ ognized this taxon as the epifamily Arthrotebrae 8 and 9 (De Villiers, 1931). Further, leptoidae, and originally Laurent (1951) con­ Hemisus burrows head­first (Channing, sidered this clade (with the possible inclusion 1995). All of these characters can safely be of Scaphiophryninae ) to be a single family, considered synapomorphies of Hemisotidae . and Dubois (2005) considered our Laurentobatrachia to be 4 of the 6 subfamilies of [148] LAURENTOBATRACHIA NEW TAXON his Brevicipitidae . We attempted to retain fa­ ETYMOLOGY: R.L. Laurent 1 batrachia miliar usage, with the exception of moving (Greek: batrachos, frog). This name cele­ Leptopelinae from Hyperoliidae to Arthrobrates the enormous contributions to amphib­ leptidae. Because we think that the diversity ian systematics by the father of central Af­ of this taxon has been greatly underestimatrican herpetology and a prominent figure in ed, our approach leaves considerable room Argentinian herpetology, Raymond L. Lau­ for more informative taxonomies as evidence rent. becomes available.

IMMEDIATELY MORE INCLUSIVE TAXON:

[143] Afrobatrachia new taxon. [149] FAMILY: HYPEROLIIDAE LAURENT, 1943 SISTER TAXON: [144] Xenosyneunitanura Hyperoliinae Laurent, 1943: 16 . Type genus: Hynew taxon. perolius Rapp, 1842.

RANGE: Sub­Saharan Africa, Madagascar, Kassinini Laurent, 1972: 201. Type genus: Kas­ and the Seychelles. sina Girard, 1853.

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 233

IMMEDIATELY MORE INCLUSIVE TAXON: taxa. We did not sample Chrysobatrachus or [143] Afrobatrachia new taxon. Callixalus and cannot guess into which

SISTER TAXON: [164] Arthroleptidae Mi­ group they would fall. Their association with vart, 1869. Acanthixalus in the tree of Drewes (1984)

RANGE: Sub­Saharan Africa and Madagas­ suggests that they might follow Acanthixalus car; Seychelles. into Kassininae, but this is merely conjecture

CONTENT: Acanthixalus Laurent, 1944 ; and a combined study of morphology and Afrixalus Laurent, 1944 ; Alexteroon Perret , molecules is ongoing by Drewes and collab­ 1988; Arlequinus Perret, 1988 ; Callixalus orators. Our results differ substantially from Laurent, 1950; Chlorolius Perret, 1988 ; the results of Vences et al. (2003d; figs. 28 , Chrysobatrachus Laurent, 1951 ; Cryptothy­ 29) with respect to the relative placement of lax Laurent and Combaz, 1950; Heterixalus several genera. This is presumably due to our Laurent, 1944; Hyperolius Rapp, 1842 (in­ application of much denser sampling and cluding Nesionixalus Perret, 1976 ); Kassina more evidence. Girard, 1853; Kassinula Laurent, 1940 ; Op­ The association by the molecular data of isthothylax Perret, 1966; Paracassina Per­ Tachycnemis ( Seychelles) and Heterixalus acca, 1907; Phlyctimantis Laurent and Com­ ( Madagascar) is of some biogeographic inbaz, 1950; Semnodactylus Hoffman, 1939 ; terest. We expected Alexteroon to be imbed­ Tachycnemis Fitzinger, 1843 . ded within Hyperolius , but our sampling of

CHARACTERIZATION AND DIAGNOSIS: One Hyperolius was insufficient to test this proplarval character in our analysis that may be osition adequately. On the basis of our limsynapomorphy of this group is (from Haas, ited exemplar selection, Alexteroon may be 2003): commissura proximalis II absent. Be­ the sister taxon of Hyperolius (sensu lato). yond that, hyperoliids are unique among However, we found, as did Drewes and Wilfrogs in having a distinctive gular gland kinson (2004), that Nesionixalus is clearly (Drewes, 1984). Drewes (1984) summarized deeply imbedded in Hyperolius , but also rep­ a character distribution suggesting that lack­ resents a monophyletic group. We suggest ing sphincter control of the vocal slits may that Nesionixalus be treated as a subgenus of also be a synapomorphy of Hyperoliidae . Hyperolius with no coordinate taxon to im­

The presence of intercalary phalangeal el­ ply that the remaining species of Hyperolius ements per se is not a synapomorphy of this are a monophyletic group (see appendix 7 for group (or at least not without making as­ new combinations). We expect that Chlorosumptions of character optimization), being lius and Arlequinus will also be found to be found also in the Leptopelinae of Arthrolep­ imbedded within Hyperolius , although at this tidae. Nevertheless, Drewes (1984) noted time no data can be brought to bear to test that hyperoliid and leptopeline intercalary el­ this proposition. ements are histologically quite different from each other. The latter does not accept either [164] FAMILY: ARTHROLEPTIDAE MIVART, 1869 Alizarin or Alcian Blue stain, suggesting that these elements may not be homologous. Arthroleptina Mivart, 1869: 294. Type genus: Ar­

throleptis Smith, 1849.

SYSTEMATIC COMMENTS: The position in

Astylosterninae Noble, 1927: 110 . Type genus: our tree of Acanthixalus is heterodox com­ Astylosternus Werner, 1898 . pared with previous studies (e.g., Drewes, Leptopelini Laurent, 1972: 201. Type genus: Lep­ 1984) and implies a number of reversals and topelis Günther, 1859. New synonym. convergences in the morphology of hyperoliid frogs. We considered recognizing sub­ IMMEDIATELY MORE INCLUSIVE TAXON: families within Hyperoliidae , corresponding [147] Laurentobatrachia new taxon. to the two major clades of exemplars, for SISTER TAXON: [149] Hyperoliidae Laurent , which the name Kassininae Laurent, 1972, is 1943. insee

available for the Kassina – Phlyctimantis – RANGE: Sub­Saharan Africa.

234 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Systematic Comments); Astylosternus Wer­ apomorphic should one be willing to make ner, 1898; Cardioglossa Boulenger, 1900 ; assumptions about character optimization Leptodactylodon Andersson, 1903 ; Lepto­ and that the phalangeal elements of leptopelis Günther, 1859; Nyctibates Boulenger , pelines and hyperoliids are homologous.

1904; Scotobleps Boulenger, 1900 ; Trichob­ Arthroleptis renders Schoutedenella paraatrachus Boulenger, 1900. phyletic, and we therefore consider them to CHARACTERIZATION AND DIAGNOSIS: Arthro­ be synonyms. Laurent and Fabrezi’s (1986 leptids are small frogs exhibiting forked ‘‘1985’’) contention that Schoutedenella and omosterna that, with the exception of Arthro­ Arthroleptis are not each other’s closest relleptis, have a typically biphasic life history. atives is rejected, although the position of Like many of the taxa within Afrobatrachia, Poynton (1964a) and Poynton and Broadley many of the arthroleptids have vertical pu­ (1967), that Schoutedenella are merely small pils, with the exceptions of Leptodactylodon Arthroleptis is also rejected. (Our tree sug­ (quadrangular) and Arthroleptini (horizontal, gests that if size were characteristic, we except for Scotobleps ). None of the morpho­ would have to say that Arthroleptis are big logical characters in our analysis optimize Schoutedenella .) Our molecular data support unambiguously to this branch [164]. Regard­ the notion that nominal Arthroleptis is imless, the molecular data are decisive in sup­ bedded within Schoutedenella and we place port of recognition of this group (see appen­ them in synonymy. (See appendix 7 for new dix 5). and revived combinations.)

Larval characters of Haas’ (2003) exem­ Perret (1966) suggested that Nyctibates is plar Leptopelis —a distinct medial ossifica­ a synonym of Astylosternus , but Amiet (1971 tion center of vertebral centra ventral to no­ ‘‘1970’’, 1973 ‘‘1972’’) resurrected Nyctitochord present (Haas 100.1)—may be syn­ bates on the basis of tadpole morphology beapomorphies of Arthroleptidae , of Leptope­ ing more similar to Leptodactylodon and Trilinae, or of some subset of Leptopelis . The chobatrachus. Our molecular data support direct development of Arthroleptis is subse­ recognition of Nyctibates .

quently derived.

SYSTEMATIC COMMENTS: We recognize two [180] NATATANURA NEW TAXON subfamilies within Arthroleptidae , [165]

Leptopelinae Laurent, 1972 , for Leptopelis , ETYMOLOGY: Natata­ (Greek: swim) 1 anformerly associated with Hyperoliidae (al­ oura (Greek: tailless, i.e., frog), referencing though shown to be phylogenetically distant that many of the frogs in this clade are semifrom them by Vences et al., 2003c), and aquatic.

[168] Arthroleptinae Mivart, 1869 , contain­ IMMEDIATELY MORE INCLUSIVE TAXON: ing two tribes, [169] Astylosternini Noble, [108] Ranoides new taxon.

1931 ( Leptodactylodon , Nyctibates , Trichob­ SISTER TAXON: [109] Allodapanura new atrachus, and Leptodactylodon ) and [172] taxon.

Arthroleptini Mivart, 1869 ( Arthroleptis [in­ RANGE: Worldwide temperate and tropical cluding Schoutedenella ], Cardioglossa , and habitats on all continents and major islands, Scotobleps ). Scotobleps formerly was asso­ except most of Australia and New Zealand.

ciated with Astylosterninae , so its transfer to CONCEPT AND CONTENT : Natatanura is a Arthroleptini is something of a surprise (on monophyletic group composed of [181] Ptythe basis of evidence shown in appendix 5). chadenidae Dubois, 1987 ‘‘1986’’, and [183] [165] Leptopelinae Laurent, 1972 , is dis­ Victoranura new taxon.

tinguished morphologically from its near CHARACTERIZATION AND DIAGNOSIS: Nataneighbors by the possession of an entire, tanura is identical to the epifamily Ranoidae rather than forked, omosternum and by his­ of Dubois (1992) and Ranidae (sensu lato) of tologically distinct intercalary phalangeal el­ Laurent (1986). Characters in our analysis ements (Drewes, 1984). (from Haas, 2003) that are likely synapo­

[168] Arthroleptinae Mivart, 1869 , is not morphies of this taxon are (1) anterior inserdiagnosable via morphology, although the tion of m. subarcualis rectus II–IV on ceraabsence of intercalary elements may be syn­ tobranchial III (Haas 37.2); (2) commissura

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 235

proximalis II absent (Haas 110.0); and (3) Lanzarana Clarke, 1982 ; Ptychadena Boucommissura proximalis III absent (Haas lenger, 1917.

111.0). CHARACTERIZATION AND DIAGNOSIS: In our J.D. Lynch (1973) and Laurent (1986) analysis, the morphological (larval) characsuggested that an ossified metasternal style is ters that attach to the only exemplar of this a synapomorphy at this level of universality, taxon, Ptychadena , are (1) m. subarcualis but this requires corroboration inasmuch as rectus I portion with origin from ceratobranseveral groups within Natatanura have carti­ chial III absent (Haas 35.0); (2) partes corlaginous metasterna (Laurent, 1986). pores medially separate (Haas 87.0); and (3) SYSTEMATIC COMMENT : Burton (1998a) eggs float as a surface film (Haas 141.2). Benoted that several genera of Natatanura share cause of our limited sampling for morpholthe presence of an extra slip of the m. flexor ogy, it is possible that these characters do not teres digiti IV, which is ventral to the m. apply to Hildebrandtia or Lanazarana; it is transversus metacarpus II: Altirana , Aubria , also possible that they apply only to a subset Ceratobatrachus , Conraua , Hildebrandtia , of Ptychadena . Only denser sampling will Mantella , Mantidactylus , Petropedetes , Pty­ tell.

chadena, Pyxicephalus , and Rana , but not in Other features that are likely synapomor­ Batrachylodes , Cacosternum , Discodeles , phies, although originally suggested in a Laliostoma , Meristogenys , Micrixalus , Nan­ somewhat different outgroup structure nophrys, Nanorana , Natalobatrachus , Nyc­ (Clarke, 1981), are (1) otic plate absent or tibatrachus, Occidozyga , Palmatorappia , rudimentary; (2) (neo)palatines absent; (3) Platymantis , or Strongylopus (with many point overlap of the medial ramus of the ptertaxa not examined). If this character is opti­ ygyoid and the anterior lateral border of the mized on our most parsimonious tree, the im­ parasphenoid ala in an anterior–posterior plication is that this character arose at least plane; (4) clavicles reduced and well­sepasix times, of which the following is one of rated at midline; (5) sternal style a short several equally parsimonious arrangements: compact bony element; (6) eight presacral (1) Ceratobatrachus ; (2) in the branch sub­ and sacral vertebrae fused (also in some Lithtending Conraua 1 Petropedetes , and there­ obates); and (7) dorsal protuberance on ilium fore likely to be in Indirana and Arthrolep­ not or only slightly differentiated from dorsal tides); (3) Ptychadenidae (Hildebrantia, Pty­ prominence, which is smooth surfaced and chadena, and presumably in Lanzarana ); (4) confluent with a well­developed ilial crest.

Pyxicephalini ( Pyxicephalus and Aubria ); (5) SYSTEMATIC COMMENT : See Systematic Altirana (5 part of Nanorana ); (6) Aglaioan­ Comments under Natatanura. Our association ura (Rhacophoroidea 1 Ranidae ). Neverthe­ of Hildebrandtia and Lanzarana with this less, considerably more specimens of more taxon rests on the morphological data analtaxa need to be examined before the opti­ ysis of Clarke (1981), who suggested a nummization of this feature can confidently be ber of synapomorphies for the group (see considered settled. above).

[181] FAMILY: PTYCHADENIDAE DUBOIS, 1987 [183] VICTORANURA NEW TAXON

‘‘1986’’ Ptychadenini Dubois, 1987 ‘‘1985’’: 55. Type ge­ ETYMOLOGY: Victor (Latin: conqueror) 1 nus: Ptychadena Boulenger, 1917 . anoura (Greek: tailless; i.e., frog), alluding to

the remarkable success of this taxon world­ IMMEDIATELY MORE INCLUSIVE TAXON: wide. [180] Natatanura new taxon. IMMEDIATELY MORE INCLUSIVE TAXON: SISTER TAXON: [183] Victoranura new tax­ [180] Natatanura new taxon. on. SISTER TAXON: [181] Ptychadenidae Du­ ex­

RANGE: Sub­Saharan tropical and subtrop­ bois, 1987 ‘‘1986’’. ical Africa; Seychelles and Madagascar. RANGE: Worldwide continents and major CONTENT : Hildebrandtia Nieden, 1907 ; islands in temperate and tropical regions, 236 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

cept southern Australia, the Seychelles, and lossini Anderson, 1871) is rejected by our New Zealand. evidence. CONCEPT AND CONTENT : Victoranura is a Dubois (1992) placed Batrachylodes outmonophyletic group composed of [184] Cer­ side of his Ceratobatrachini, because, unlike atobatrachidae Boulenger, 1884, and [189] the more typical members of Ceratobatrachi­ Telmatobatrachia new taxon. nae, it lacks a forked omosternum. Neverthe­ CHARACTERIZATION AND DIAGNOSIS: None less, Batrachylodes does have endotrophic of the morphological characters in our anal­ larvae (Thibaudeau and Altig, 1999), and our ysis diagnose on this taxon, although the mo­ molecular evidence places Batrachylodes lecular data are decisive (see appendix 5 for firmly within the ceratobatrachine clade. summary of molecular synapomorphies). Roelants et al. (2004) provided molecular

evidence suggesting that Ingerana is in Oc­

[184] FAMILY: CERATOBATRACHIDAE cidozyginae rather than Ceratobatrachinae ,

BOULENGER, 1884 but this is not corroborated by our denser

taxonomic sampling and larger amount of Ceratobatrachidae Boulenger, 1884: 212 . Type genus: Ceratobatrachus Boulenger, 1884 . data, which place Ingerana in the more con­ Platymantinae Savage, 1973: 354. Type genus: ventional location in Ceratobatrachidae and Platymantis Günther, 1859 . as the sister taxon of the remaining genera

within Ceratobatrachinae . Like Roelants et IMMEDIATELY MORE INCLUSIVE TAXON: al. (2004), we did not evaluate species of the [183] Victoranura new taxon. nominal subgenus Liurana , a taxon that Du­ SISTER TAXON: [189] Telmatobatrachia new bois (1987 ‘‘1985’’) erected as a subgenus of taxon. Ingerana , but subsequently was recognized RANGE: Western China (Xizang and Yun­ by some workers as a genus (Fei et al., 1997) nan); Myanmar, adjacent Thailand and pen­ and later (Dubois, 2005, without discussion) insular Malaysia; Philippines, Borneo; New as a synonym of Taylorana (5 Limnonectes ). Guinea; Admiralty, Bismarck, and Solomon Liurana is reported to be differentiated from Islands; Fiji; Palau. Ingerana by condition of the finger disc (ab­ CONTENT: Batrachylodes Boulenger, 1887 ; sent in Liurana , present in Ingerana ) and Ceratobatrachus Boulenger, 1884 ; Discode­ median lingual papilla (present in Liurana , les Boulenger, 1918; Ingerana Dubois, 1987 absent in Ingerana ; Dubois, 1987 ‘‘1985’’), ‘‘1986’’; Palmatorappia Ahl, 1927 ‘‘1926’’; but some species of Liurana possess small Platymantis Günther, 1858 . finger discs (Zhao and Li, 1984; Fei et al., CHARACTERIZATION AND DIAGNOSIS: None 2005), and the condition of the tongue is of the morphological characters in our anal­ known for only two of the five species of ysis optimize as synapomorphies of this tax­ Ingerana (Smith, 1930; Inger, 1954, 1966). on, although all ceratobatrachids are charac­ We treat Liurana as a synonym of Ingerana , terized by large eggs and direct development pending evidence being published to sub­ (Noble, 1931). Many of the species have ex­ stantiate Dubois’ (2005) assertion of its panded toe tips, but this is likely plesiom­ placement in Limnonectini ( Dicroglossidae ). orphic at this level of universality. Molecular synapomorphies for the clade are summa­ [189] TELMATOBATRACHIA NEW TAXON rized in appendix 5. SYSTEMATIC COMMENT : Dubois (1987 ETYMOLOGY: Telmato­ (Greek: of a marsh) ‘‘1985’’, 1992) placed his Ceratobatrachiini 1 batrachos (Greek: frog), referencing the Boulenger, 1884, within a larger Dicroglos­ preference of these frogs for wet microhabsinae Anderson, 1871. The subsequent im­ itats. plication of Dubois et al. (2001) that Cera­ IMMEDIATELY MORE INCLUSIVE TAXON: tobatrachidae (his Ceratobatrachinae ) is of [183] Victoranura new taxon. uncertain relationship to Dicroglossinae was SISTER TAXON: [184] Ceratobatrachidae

justified inasmuch as an inclusive Dicroglos­ Boulenger, 1884. sinae (including Ceratobatrachiini Boulenger, RANGE: Worldwide continents and major 1884, Conrauini Dubois, 1992 , and Dicrog­ islands in temperate and tropical environ­

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 237

ments, except for southern South America, cluding Madagascar, New Zealand, Sey­ Madagascar, New Zealand, and most of Aus­ chelles, and Australia except for the far tralia. north.

CONCEPT AND CONTENT : Telmatobatrachia CONCEPT AND CONTENT : Ametrobatrachia is is a monophyletic taxon composed of [190] a monophyletic taxon composed of [192] Af­ Micrixalidae Dubois, Ohler, and Biju, 2001 , ricanura new taxon and [220] Saukrobatra­ and [191] Ametrobatrachia new taxon. chia new taxon.

CHARACTERIZATION AND DIAGNOSIS: None CHARACTERIZATION AND DIAGNOSIS: None of the morphological characters in our anal­ of the morphological characters in our analysis optimize on the branch subtending this ysis optimize as synapmorphies of this taxon. taxon although our molecular data decisively Nevertheless, the molecular data are decisupport its recognition. (See appendix 5 for sive. (See appendix 5 for summary of molisting of molecular synapomorphies.) lecular synapomorphies for this taxon.)

[190] FAMILY: MICRIXALIDAE DUBOIS, OHLER , [192] AFRICANURA NEW TAXON

AND BIJU, 2001

Micrixalinae Dubois et al., 2001: 54. Type genus: ETYMOLOGY: Afric­ (Latin: of Africa) 1 Micrixalus Boulenger, 1888 . anoura (Greek: tailless, i.e., frog).

IMMEDIATELY MORE INCLUSIVE TAXON: IMMEDIATELY MORE INCLUSIVE TAXON: [191] Ametrobatrachia new taxon.

[189] Telmatobatrachia new taxon. SISTER TAXON: [220] Saukrobatrachia new SISTER TAXON: [191] Ametrobatrachia new taxon.

taxon. RANGE: Sub­Saharan Africa.

RANGE: India. CONTENT: [193] Phrynobatrachidae Lau­ CONTENT: Micrixalus Boulenger, 1888 . rent, 1941 ‘‘1940’’, and [200] Pyxicephalo­ CHARACTERIZATION AND DIAGNOSIS: None idea Bonaparte, 1850.

of the morphological characters in our anal­ CHARACTERIZATION AND DIAGNOSIS: None ysis optimize on this taxon and the decisive of the morphological characters in our analevidence for its recognition is entirely mo­ ysis optimize on this taxon. Nevertheless, lecular (see appendix 5 for summary). Unlike molecular data are decisive. (See appendix 5 Ptychadenidae , Ceratobatrachidae , and basal­ for summary of molecular transformation asly in Ametrobatrachia, the omosternum is un­ sociated with this taxon.)

forked in Micrixalidae (Dubois et al., 2001) , SYSTEMATIC COMMENT : The existence of which at this level of universality is a syna­ this taxon had not been suspected prior to the pomorphy of the group as is the low kera­ publication of Van der Meijden et al. (2005), todont formula 1/0 (Dubois et al., 2001). The although it certainly meets biogeographic expresence of digital discs in Micrixalinae is pectations.

likely a plesiomorphy at this level of univer­

sality. [193] FAMILY: PHRYNOBATRACHIDAE

LAURENT, 1941 ‘‘1940’’

[191] AMETROBATRACHIA NEW TAXON

Hemimantidae Hoffmann, 1878: 613 . Type genus: ETYMOLOGY: Ametros (Greek: beyond Hemimantis Peters, 1863 .

measure) 1 batrachos (Greek: frog), denot­ Phrynobatrachinae Laurent, 1941 ‘‘1940’’: 79. ing the enormity of this taxon in terms of Type species: Phrynobatrachus Günther, 1862 . species and with respect to the enormous

numbers of questions that remain about its IMMEDIATELY MORE INCLUSIVE TAXON: internal phylogenetic structure. [192] Africanura new taxon.

IMMEDIATELY MORE INCLUSIVE TAXON: SISTER TAXON: [200] Pyxicephaloidea Bon­ [189] Telmatobatrachia new taxon. aparte, 1850.

SISTER TAXON: [190] Micrixalidae Dubois, RANGE : Sub­Saharan Africa.

Ohler, and Biju, 2001. CONTENT: Ericabatrachus Largen, 1991 RANGE : Worldwide in temperate and trop­ (see Systematic Comments); Phrynobatraical continental areas and major islands, ex­ chus Günther, 1862 (including Dimorphog­

238 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

nathus Boulenger, 1906, and Phrynodon chus as currently arrayed, but at present we Parker, 1935; see Systematic Comments). cannot reject the possibility that it is the sis­

CHARACTERIZATION AND DIAGNOSIS: Phry­ ter taxon of Phrynobatrachus . We presume nobatrachids are small terrestrial and semi­ that Dubois’ (2005) association of Ericabaaquatic frogs with poorly understood species trachus with his Phrynobatrachinae is based boundaries and with a typically biphasic life on similar reasoning although he provided no history, with eggs laid in water. Like many justification for this inclusion. members of Ranoides, phrynobatrachids frequently have T­shaped terminal phalanges, [200] SUPERFAMILY: PYXICEPHALOIDEA although they lack digital discs. They usually BONAPARTE, 1850 retain an outer metatarsal tubercle (Laurent, IMMEDIATELY MORE INCLUSIVE TAXON: 1986) and are characterized by a tarsal tu­ [192] Africanura new taxon. bercle (Channing, 2001) that is distinctive SISTER TAXON: [193] Phrynobatrachidae and may be synapomorphic. Phrynobatra­ Laurent, 1941 ‘‘1940’’. chus species exhibit a median lingual tuber­ RANGE: Sub­Saharan Africa. cle (Grant et al., 1997), which may be syn­ CONTENT: [201] Petropedetidae Noble , apomorphic, although this needs to be care­ 1931, and [209] Pyxicephalidae Bonaparte , fully surveyed. Its presence also in Indirana, 1850. Arthroleptides , and Petropedetes suggests CHARACTERIZATION AND DIAGNOSIS: Althat it may be synapomorphic at a more gen­ though no morphological characters in our eral level. study optimize to this branch, our molecular

Nevertheless, none of the morphological data are decisive. See appendix 5 for sumcharacters in our analysis optimize on this mary of molecular synapomorphies. taxon, although the molecular data are deci­ COMMENT: This taxon is highy heterogesive in recognition of this taxon. (See appen­ nous morphologically, at least with respect to dix 5 for listing of molecular synapomor­ overall appearance. Nevertheless, the molecphies for this taxon.) ular evidence is strong, and the taxon should

SYSTEMATIC COMMENTS: Our data show survive additional testing. that Phrynobatrachus is paraphyletic with respect to Phrynodon and Dimorphognathus . [201] LFAMILY: PETROPEDETIDAE NOBLE, 1931 Surprisingly, Amiet (1981) suggested a close Petropedetinae Noble, 1931 : 520. Type genus: Perelationship of Phrynodon with Petropedetes tropedetes Reichenow, 1874 . ( Petropedetidae ) to the exclusion of Phry­ Ranixalini Dubois, 1987 ‘‘1985’’: 66. Type genus: nobatrachus. Our data do not support this re­ Ranixalus Dubois, 1986 . New synonym. lationship and because this nominal genus Conrauini Dubois, 1992: 314 . Type genus: Con­ and Dimorphognathus are both monotypic raua Nieden, 1908. New synonym. and imbedded within Phrynobatrachus , we Indiraninae Blommers­Schlösser, 1993: 211. Type place Phrynodon and Dimorphognathus into genus: Indirana Laurent, 1986. New synonym. the synonymy of Phrynobatrachus , which af­ IMMEDIATELY MORE INCLUSIVE TAXON: ter this action is monophyletic. Nevertheless, [200] Pyxicephaloidea Bonaparte, 1850. Phrynobatrachus remains one of the larger SISTER TAXON: [209] Pyxicephalidae Bontaxonomic problems in Africa in terms of aparte, 1850. species boundaries and infrageneric clades. It RANGE: South India; tropical West and will yield its secrets only with a considerable East Africa. amount of morphological, behavioral, and CONTENT: Arthroleptides Nieden, 1911 molecular work. (See appendix 7 for new ‘‘1910’’; Conraua Nieden, 1908 ; Indirana and revivied combinations caused by these Laurent, 1986; Petropedetes Reichenow , synonymies.) Our inclusion in Phrynoba­ 1874. trachidae of Ericabatrachus Largen, 1991 CHARACTERIZATION AND DIAGNOSIS: Petro­ (not studied by us) rests on the original pub­ pedetidae is heterogeneous morphologically,

lication, which suggests that Ericabatrachus with forked omosterna. No morphological is ‘‘ Phrynobatrachus ­like’’. Likely, it will be synapomorphies are evident to us, although found to be imbedded within Phrynobatra­ the molecular data are decisive. (See appen­

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 239

dix 5 for molecular synapomorphies for this Amiet and Perret, 1969; Inger et al., 1984; taxon.) Drewes et al., 1989).

SYSTEMATIC COMMENTS: The association of Adults of Arthroleptides, Indirana, and Indirana ( India), Conraua (tropical West Af­ Petropedetes also share characters whose porica; Ethiopia and Eritrea), and Arthrolep­ larity is less clear. Males of most Petropetides 1 Petropedetes (tropical West Africa; detes and Arthroleptides , and males of Indi­ Tanzania and Kenya) at first surprised us, rana (where they are known) share the preseven though we had expected the undiagnos­ ence of femoral glands of variable size and able Petropedetidae (sensu lato, now distrib­ the presence of spicules around the margins uted among Petropedetidae , Phrynobatrachi­ of jaw and/or chin in the pectoral area dae, and Dicroglossidae ) to be obliterated. (Amiet, 1973; Inger et al., 1984; Perret,

The stream­dwelling larvae of Arthrolep­ 1984; Dubois, 1986 ‘‘1985’’; Klemens, 1998; tides and stream­dwelling and arboreal tad­ however spicules are absent in Petropedetes poles of Indirana are amazingly similar parkeri [Amiet, 1983], and femoral glands (compare Altig and Johnston, 1989, and are absent in A. yakusini [Channing et al., Channing et al., 2002b, with Annandale and 2002b]). Note that spicules around the mar­ Rao, 1918) in having elongate tails with very gins of jaw and/or chin and pectoral area, low caudal fins, large bulging eyes, a dor­ occur also in Conraua and in at least several soventrally flattened body, and a laterally phrynobatrachids as redefined here (Perret, compressed jaw sheath with prominent lat­ 1966). Until this character can be widely aseral processes (Annandale, 1918; Rao, 1920; sessed its level of generality remains un­ Amiet and Perret, 1969; Inger et al., 1984; known.

Dubois, 1986 ‘‘1985’’; Drewes et al., 1989; Dubois (1987 ‘‘1985’’) proposed the rec­ Channing et al., 2002b). Only larvae of Pe­ ognition of the tribe Ranixalini (later treated tropedetes natator and P. palmipes have as a subfamily by Dubois, 1992), for the genbeen fully described (Lamotte and Zuber­Vo­ era Nannophrys , Nyctibatrachus , and Indigeli, 1954; Lamotte et al., 1959; Lamotte and rana on the basis of the presence of femoral Lescure, 1989), but some superficial refer­ glands in males of Nyctibatrachus and Indiences to morphology or behavior are avail­ rana (unknown in Nannophrys ), and the able for the larvae of P. cameronensis (Bou­ morphological proximity of Nannophrys and lenger, 1906 ‘‘1905’’; Lawson, 1993), P. Nyctibatrachus was noted by Clarke (1981). newtoni (Perret, 1966; Amiet and Perret, Nannophrys and Indirana further share the 1969; Lawson, 1993), and P. parkeri and P. modifications of larval morphology associjohnstoni (Amiet and Perret, 1969; Amiet, ated with semiterrestrial life that were men­ 1983; Lawson, 1993). Drewes et al. (1989) tioned earlier (Kirtisinghe, 1958). From a noted inconsistencies in the description of morphological perspective, the evidence supthe larva of P. palmipes . Regardless, from porting the monophyly of Nannophrys 1 Inthe comments or illustrations presented by dirana is the same as that favoring a relathe authors mentioned above, larvae of Pe­ tionship among Indirana, Arthroleptides , and tropedetes seem to have the same morpho­ Petropedetes . As discussed earlier, other logical peculiarities as do those of Arthrolep­ characters of still unclear polarity that could tides and Indirana. The only exception is the further support this hypothesis are the preslarva of P. natator , which has an abdominal ence of femoral glands and spicules around disc and an oral disc that is proportionally the margins of jaw and/or chin and pectoral larger, with conspicuous lateral folds, and area.

jaw sheaths that are not compressed laterally Petropedetes and Arthroleptides have (Lamotte and Zuber­Vogeli, 1954; Lamotte large digital discs, a long metasternal style, and Lescure, 1989). and T­shaped terminal phalanges. Indirana

In transforming larvae of Arthroleptides , has Y­shaped terminal phalanges (Laurent, Indirana, and Petropedetes , the hind legs are 1986), which may be synapomorphic with to­

of

large and seem to develop precociously, on the T­shaped terminal phalanges of Petro­ a different growth trajectory from the front pedetes 1 Arthroleptides although in our legs (Annandale, 1918; Lamotte et al., 1959; pology the simple terminal phalanges 240 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

Conraua presumably represent the apomor­ SYSTEMATIC COMMENTS: This morphologiphy. Roelants et al. (2004) suggested that In­ cally heterogeneous taxon is coherent geodirana would find its closest relatives in In­ graphically. Although the association of dia. However, inasmuch as these authors did these genera was only noted recently (Van not include any African taxa in their analysis, der Meijden et al., 2005), much of the earlier it was impossible for them to detect a rela­ taxonomy was based on very general notions tionship with African taxa. Van der Meijden of overall similarity, which are significantly et al. (2005) placed Indirana as the sister tax­ influenced by perceptions of body size. The on of our Dicroglossinae . They also placed association of Afrana and Strongylopus (for­ Conraua outside of a clade composed of Pe­ merly in Ranini of Dubois, 1992) with Antropedetes 1 Pyxicephalinae , in both cases hydrophryne, Arthroleptella , Cacosternum , on the basis of fewer data and more analyt­ and Natalobatrachus (formerly of Phrynoical assumptions. Additional data or denser batrachidae [ Petropedetidae ] of Dubois, taxon sampling may rearrange these taxa, but 1992), and with Pyxicephalus and Aubria (in at present our molecular data are decisive Pyxicephalinae of Dubois, 1992), was some­ and, as discussed earlier, they are consistent thing of a surprise (at least for us, as this was with the distribution of various larval and before Van der Meijden et al., 2005, apadult characteristics. peared), although no evidence beyond overall similarity ever supported the older tax­ [209] FAMILY: PYXICEPHALIDAE BONAPARTE , onomy. We still have three ‘‘flavors’’ of frogs 1850 in this group: those that look like Rana (Af­ Pyxicephalina Bonaparte, 1850: 1. Type genus: rana and Strongylopus ); those that are stocky Pyxicephalus Tschudi, 1838 . and big ( Pyxicephalus and Aubria ); and Phrynopsinae Noble, 1931: 518. Type genus: those that are generally small and have not Phrynopsis Pfeffer, 1893 . attracted from systematists the attention they Cacosterninae Noble, 1931: 540 . Type genus: Ca­ deserve (the remainder). The absence of a costernum Boulenger, 1887. median lingual process may be synapo­ Tomopternini Dubois, 1987 ‘‘1985’’: 56. Type ge­ morphic, as this feature is present in Petronus: Tomopterna Duméril and Bibron, 1841 . pedetidae and Phrynobatrachidae (Grant et New synonym. al., 1997). Dubois (2005), anticipating the IMMEDIATELY MORE INCLUSIVE TAXON: publication of Van der Meijden et al. (2005), [200] Pyxicephaloidea Bonaparte, 1850. recognized this taxon as a subfamily of Ran­ SISTER TAXON: [201] Petropedetidae Noble , idae, Pyxicephalinae , which we recognize as 1931. a family.

RANGE: Sub­Saharan Africa. Within Pyxicephalidae , we recognize two CONTENT: Amietia Dubois, 1987 ‘‘1986’’ subfamilies: [210] Pyxicephalinae Bonapar­ (including Afrana Dubois, 1992 , see Sys­ te, 1850 ( Pyxicephalus and Aubria ) and tematic Comments); Anhydrophryne Hewitt , [212] Cacosterninae Noble, 1931 (for the re­ 1919; Arthroleptella Hewitt, 1926 ; Aubria maining genera). Pyxicephalinae is united by Boulenger, 1917; Cacosternum Boulenger , the following synapomorphies: (1) skull ex­ 1887; Microbatrachella Hewitt, 1926 ; Na­ ostosis; (2) occipital canal present; (3) zytalobatrachus Hewitt and Methuen, 1912; gomatic ramus much longer than otic ramus, Nothophryne Poynton, 1963 ; Poyntonia articulating with the postorbital process of Channing and Boycott, 1989; Pyxicephalus the pars facialis of the maxilla; and (4) strong Tschudi, 1838; Strongylopus Tschudi, 1838 ; overlap of the medial ramus of the pterygoid Tomopterna Duméril and Bibron, 1841 . and the parasphenoid ala (Clarke, 1981). Ca­ CHARACTERIZATION AND DIAGNOSIS: Al­ costerninae in our sense is not united by any though we know of no morphological syna­ morphological feature that we can identify pomorphies for this group, the molecular ev­ with any certainty, although the molecular idence is decisive in support of this branch. data are decisive (see appendix 5).

(See appendix 5 for molecular synapomor­ We place Afrana Dubois, 1992 , into the phies of this taxon; also see Systematic Com­ synonymy of [218] Amietia Dubois, 1987 ments.) ‘‘1986’’, to resolve the paraphyly of Afrana .

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 241

No characteristics of ‘‘ Afrana ’’ or Amietia of the morphological characters in our analreject this placement. ysis optimize on this taxon although the mo­ Clearly, our data do not support the notion lecular data decisively support recognition of (Poynton, 1964a) that Cacosternum is close­ this taxon. (See appendix 5 for molecular ly related to Phrynobatrachus . Our associa­ transformations.) We recognized two subtion of Microbatrachella , Nothophryne , and families within Dicroglossidae , which are Poyntonia with this clade is provisional, discussed in separate accounts because of the based on the assertion by Blommers­Schlös­ size and complexity of discussion. ser (1993) that these genera are allied by reduced ossification of the omosternal style [225] SUBFAMILY: DICROGLOSSINAE and procoracoid clavicular bar. ANDERSON, 1871

Dicroglossidae J. Anderson, 1871: 38 . Type ge­ [220] SAUKROBATRACHIA NEW TAXON nus: Dicroglossus Günther, 1860 .

Limnonectini Dubois, 1992: 315. Type genus: ETYMOLOGY: Saukro­ (Latin: graceful, Limnonectes Fitzinger, 1843 . pretty) 1 batrachos (Greek: frog), referenc­ Paini Dubois, 1992: 317. Type genus: Paa Duing the beauty of many of the species in­ bois, 1975. cluded in this clade. IMMEDIATELY MORE INCLUSIVE TAXON: IMMEDIATELY MORE INCLUSIVE TAXON: [221] Dicroglossidae Anderson, 1871 . [191] Ametrobatrachia new taxon. SISTER TAXON: [222] Occidozyginae Fei, SISTER TAXON: [192] Africanura new tax­ Ye, and Huang, 1991 ‘‘1990’’. on. RANGE: Northwestern and sub­Saharan Af­ RANGE: Eurasia, Africa, and Madagascar, rica; southern Arabian Peninsula; Pakistan, to northern Australia; North and Central­ Afghanistan, India, Sri Lanka, and Nepal, America to central South America. through southern China (including part of CONCEPT AND CONTENT : Saukrobatrachia Xizang) and Indochina to the islands of the new taxon is a monophyletic taxon com­ Sunda Shelf; Japan. posed of [221] Dicroglossidae Anderson, CONTENT : Annandia Dubois, 1992 (see 1871, and [244] Aglaioanura new taxon. Systematic Comments); Eripaa Dubois, CHARACTERIZATION AND DIAGNOSIS: Al­ 1992 (see Systematic Comments); Euphlycthough no morphological characters that we tis Fitzinger, 1843; ‘‘ Fejervarya ’’ Bolkay, are aware of optimize on this branch, the mo­ 1915 (see Systematic Comments); Hoplobalecular data are decisive in support of this trachus Peters, 1863; Limnonectes Fitzinger , taxon. (See appendix 5 for listing of molec­ 1843 (including Taylorana Dubois, 1987 ular synapomorphies.) ‘‘1986’’); Minervarya Dubois, Ohler , and

Biju, 2001; Nannophrys Günther, 1869 [221] FAMILY: DICROGLOSSIDAE ANDERSON,

1871 ‘‘1868’’; Nanorana Günther, 1896 (includ­

ing Altirana Stejneger, 1927 ; Chaparana IMMEDIATELY MORE INCLUSIVE TAXON: Bourret, 1939; and Paa Dubois, 1975 ; see [220] Saukrobatrachia new taxon. Systematic Comments); Ombrana Dubois, SISTER TAXON: [244] Aglaioanura new tax­ 1992 (see Systematic Comments); Quasipaa on. Dubois, 1992; Sphaerotheca Günther, 1859 RANGE : Northwestern and sub­Saharan Af­ ‘‘1858’’. rica; southern Arabian Peninsula; Pakistan, CHARACTERIZATION AND DIAGNOSIS: Al­ Afghanistan, India, Sri Lanka, and Nepal, though the molecular evidence is decisive for through southern China (including part of the existence of Dicroglossinae , we are Xizang) and Indochina to Japan and the Phil­ aware of no morphological synapomorphies ippines; islands of the Sunda Shelf as far as that optimize to this branch. (See Systematic Flores. Comments.) Appendix 5 shows the molecu­ CONTENT: [225] Dicroglossinae Anderson , lar transformations associated with this tax­

1871, and [222] Occidozyginae Fei, Ye , and on.

Huang, 1991 ‘‘1990’’. SYSTEMATIC COMMENTS: Within Dicroglos­ CHARACTERIZATION AND DIAGNOSIS: None sinae Anderson, 1871, we recognize two

242 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

monophyletic tribes, [226] Limnonectini Du­ pending the publication of evidence, we rebois, 1992, for Limnonectes (including as gard these as monotypic genera of uncertain synonyms Elachyglossa Anderson, 1916 ; placement within Dicroglossidae (see appen­ Taylorana Dubois, 1987 ), and [232] Dicrog­ dix 7 for combinations).

lossini Anderson, 1871, for the remaining Previous authors (Dubois and Ohler, 2000; genera, Annandia , ‘‘ Fejervarya ’’ (see be­ Dubois et al., 2001; Grosjean et al., 2004) low), Nanorana (including Chaparana and demonstrated that Sphaerotheca and Fejer­ Paa ), Quasipaa , Sphaerotheca , Nannophrys , varya are closely related. Our data permit us Euphlyctis , and Hoplobatrachus . (Evidence to go further and suggest strongly that recfor both is listed in appendix 5.) This agrees ognition of Sphaerotheca (as well as Euwith several other phylogenetic analyses that phlyctis, Hoplobatrachus , and Nannophrys ) used DNA evidence (e.g., Bossuyt and Mil­ renders Fejervarya sensu Dubois and Ohler inkovitch, 2000; Emerson et al., 2000b; Mar­ (2000) paraphyletic, as does a group commayou et al., 2000; Vences et al., 2000c; Ko­ posed of Nannophrys , Euphlyctis , and Hosuch et al., 2001; Grosjean et al., 2004; Roe­ plobatrachus. J. M. Hoyos (in Dubois and lants et al., 2004; Jiang et al., 2005; Jiang Ohler, 2000) suggested that Fejervarya does and Zhou, 2005), although our expanded tax­ have a morphological synapomorphy: venon sampling and data altered some relation­ trolateral edge of the m. pectoralis pars abships within Dicroglossini. dominalis slightly attached to muscles that As noted in ‘‘Results’’, our results are are dorsal relative to it, which results in a strongly congruent with those of Jiang et al. dark ventrolateral line from axilla to groin, (2005), especially when the rooting point is especially visible in live specimens. This corrected by our larger outgroup sampling needs to be verified with reference to the (see fig. 64 ). Because their analysis provided condition in Sphaerotheca and the other sat­ DNA sequence evidence unrejected by mor­ ellite genera as well as to assure that this is phological synapomorphies, we take their re­ universal in Fejervarya and not just in some sults at face value: Nanorana as they viewed subset of the nominal genus. Serious systemit is imbedded within a paraphyletic ‘‘ Paa ’’, atic and nomenclatural issues impede reso­ and ‘‘ Chaparana ’’ is polyphyletic with the lution of this paraphyly. The most important two components both imbedded within is that there are many species of nominal Fe­ ‘‘ Paa ’’. Nevertheless, they provided evi­ jervarya that we did not study, and there may dence that their Group 1 (composed of nom­ be several species of frogs masquerading uninal Paa , Nanorana , and Chaparana , and ex­ der the name Fejervarya limnocharis (Ducluding Quasipaa ), is monophyletic. Group bois and Ohler, 2000). Because our exemplar 1 is characterized by paired patches of spines of Fejervarya limnocharis is from Vietnam on the chest (Jiang et al., 2005), which may and the type locality of this same nominal not be synapomorphic but distinguishes this taxon is Java, we are reluctant to assume too taxon morphologically from Quasipaa . The much about the phylogenetic placement of F. oldest name for Group 1 is Nanorana Gün­ limnocharis sensu stricto. Ongoing research ther, 1896. (See appendix 7 for the name by Dubois and Ohler (cited in Dubois and changes that extend from the synonymy of Ohler, 2000) should provide some resolution Chaparana Bourret, 1939 , and Paa Dubois , in the near future to this problem. In the in­ 1975, with Nanorana Günther, 1896 .) An­ terim we recommend using quotation marks nandia Dubois, 1992, and Ombrana Dubois , around the name ‘‘ Fejervarya ’’ to denote the 1992, were originally named as subgenera of paraphyly of this taxon.

Chaparana , and Eripaa Dubois, 1992 , was We reaffirm that placement of Limnonecoriginally named as a subgenus of Paa . None tes limborgi in the monotypic genus Taylorof these three taxa were included, discussed, ana renders Limnonectes paraphyletic and or even mentioned in the study of Jiang et therefore continue the synonymy of Tayloral. (2005). Without discussion, Dubois ana with Limnonectes , following Inger

(2005) transferred Annandia into Limnonec­ (1996) and Emerson et al. (2000a). Emerson tini. The placement of these taxa in Dicrog­ et al. (2000a) and Evans et al. (2004) prolossinae is presumably not controversial, so vided considerable evidence that Elachyglos­

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 243

sa (formerly Bourretia ) renders Limnonectes [244] AGLAIOANURA NEW TAXON paraphyletic as well. We therefore reject the ETYMOLOGY: Aglaio­ [Greek: splendid or use of subgenera—at least as currently for­ noble] anoura [Greek: tailless, i.e., frog]. 1

mulated—within Limnonectes , even though I: MMEDIATELY MORE INCLUSIVE TAXON some authors (e.g., Delorme et al., 2004) [220] Saukrobatrachia new taxon.

have retained their use even though they mis­ SISTER TAXON: [221] Dicroglossidae An­ lead about evolutionary relationship. derson, 1871.

Although Minervarya exhibits the ‘‘Fejer­ RANGE: Eurasia, Africa, and Madagascar, varyan line’’ (of Dubois and Ohler, 2000; see to northern Australia; the Americas exclud­ Dubois et al., 2001), it was not included in ing southern South America.

our study, so we are unable to make any CONCEPT AND CONTENT : Aglaioanura is a comments about its position in the tree. Our monophyletic group composed of [245] Rhainclusion of Minervarya in Dicroglossinae is cophoroidea Hoffman, 1932 (1858), and obviously provisional; additional study is [269] Ranoidea Rafinesque, 1814 .

needed. CHARACTERIZATION AND DIAGNOSIS: On the basis of our few exemplars for morphology [222] SUBFAMILY: OCCIDOZYGINAE FEI, YE , ( Chiromantis xerampelina , Rhacophorus AND HUANG, 1991 ‘‘1990’’ pardalis , Rana nigrovittata , and Rana temporaria ) the following characters are sug­ Occydozyginae Fei et al., 1991 ‘‘1990’’: 123. gested as possibly synapomorphies of this Type genus: Occidozyga Kuhl and Van Hasselt , group: (1) functional larval m. levator man­ 1822. dibulae lateralis absent (Haas 56.0); and (2) terminal phalanges bifurcated T­shape or Y­ IMMEDIATELY MORE INCLUSIVE TAXON: shaped (Haas 156.2; reversed in several lin­ [221] Dicroglossidae Anderson, 1871 . eages of Ranidae ). (Molecular synapomor­ SISTER TAXON: [225] Dicroglossinae An­ phies are provided in appendix 5.)

derson, 1871.

RANGE: Southern China (Guangxi, Yun­ [245] SUPERFAMILY: RHACOPHOROIDEA nan, and Hainan), Thailand, Indochina, Ma­ HOFFMAN, 1932 (1858)

laya, Greater and Lesser Sunda Islands as far

as Flores, and Philippines. IMMEDIATELY MORE INCLUSIVE TAXON: CONTENT: Occidozyga Kuhl and Hasselt , [244] Aglaioanura new taxon.

1822 (including Phrynoglossus Peters, 1867 ; SISTER TAXON: [269] Ranoidea Rafinesque , see Systematic Comments). 1814.

C: Al­ RANGE: Tropical sub­Saharan Africa; MadHARACTERIZATION AND DIAGNOSIS

though the molecular data are decisive (see agascar; South India and Sri Lanka; Japan; northeastern India to eastern China south appendix 5), Occidozyginae has other syna­ through the Philippines and Greater Sundas; pomorphies: (1) aquatic larvae with a kera­ Sulawesi.

todont formula of 0/0; and (2) a lateral line CONTENT: [246] Mantellidae Laurent , system that persists into adulthood (absent in 1946, and [253] Rhacophoridae Hoffman , Occidozyga lima ; Dubois et al., 2001; con­ 1932 (1858).

vergent in Euphlyctis : Dicroglossinae ). C: See HARACTERIZATION AND DIAGNOSIS SYSTEMATIC COMMENTS: Our data demon­ Rhacophoridae . One character in our analysis strate that Phrynoglossus (which retains the definitely optimizes on this taxon: intercalary lateral line system into adulthood) is para­ element present (Haas 151.1). Channing phyletic with respect to Occidozyga (which (1989) also suggested the following as syndoes not). We therefore agree with Inger apomorphies: (1) only one slip of the m. ex­ (1996) that Phrynoglossus is a synonym of tensor digitorum communis longus, inserting Occidozyga (the senior name), providing a on distal portion of fourth metatarsal; and (2)

in­

monophyletic Occidozyga . (See appendix 7 outermost slip of the m. palmaris longus for new and revived combinations resulting serting on the proximolateral rim of the apofrom this synonymy.) neurosis palmaris. Ford and Cannatella

244 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

(1993) also suggested that bifurcate terminal and Vences, 1994). They share with their sisphalanges are a synapomorphy of this taxon, ter taxon, Rhacophoridae , intercalary phalanalthough this character may optimize at a geal elements.

more general level inasmuch as expanded toe Laurent (1986: 764) distinguished manteltips seem to optimize on or near Aglaioan­ lids from rhacophorids solely on basis of the ura. third carpal being fused with the fourth and SYSTEMATIC COMMENTS: Our study puts to fifth in rhacophorids, but being free in manrest whether mantellids and rhacophorids are tellids (this feature is likely synapomorphic sister taxa (e.g., Emerson et al., 2000b) or at this level of universality). Nevertheless, mantellids are imbedded in some way within this feature has not been adequately assayed, the rhacophorids (Liem, 1970). Whether they so at present the molecular evidence is parshould be considered mutual subfamilies of ticularly decisive in distinguishing this as a a larger Rhacophoridae (5 Rhacophoroidea monophyletic group that forms the sister taxin our use) is not a scientific proposition. We on of Rhacophoridae . None of the morphofollow the usage of Glaw and Vences (e.g., logical characters in our analysis optimize on Vences et al., 2002; Vallan et al., 2003; this taxon. (Molecular transformations are Vences et al., 2003a; Vences and Glaw, listed in appendix 5.)

2004). SYSTEMATIC COMMENTS: Vences and Glaw (2001) recognized three subfamilies on the [246] FAMILY: MANTELLIDAE LAURENT, 1946 basis of molecular data arranged phylogenetically: Laliostominae ( Boophinae 1 Mantel­ Mantellinae Laurent, 1946: 336 . Type genus: linae). We consider Mantellinae and Lalios­ Mantella Boulenger, 1882 . tominae of Vences and Glaw (2001) to be Boophinae Vences and Glaw, 2001: 88 . Type ge­

nus: Boophis Tschudi, 1838 . tribes within a larger subfamily [248] Man­ Laliostominae Vences and Glaw, 2001: 88 . Type tellinae, this subfamily forming the sister taxgenus: Laliostoma Glaw, Vences, and Böhme , on of [247] Boophinae . Aglyptodactylus and 1998. Laliostoma are in [249] Laliostomini, and within Boophini, only Boophis , and [252] IMMEDIATELY MORE INCLUSIVE TAXON: Mantella and [251] ‘‘ Mantidactylus ’’ are in [245] Rhacophoroidea Hoffman, 1932 [250] Mantellini. Although ‘‘ Mantidactylus ’’ (1858). is clearly paraphyletic with respect to Man­ SISTER TAXON: [253] Rhacophoridae Hoff­ tella (e.g., Vences and Glaw, 2001), our limman, 1932 (1858). ited taxon sampling did not reveal this. It RANGE: Madagascar. should be noted that there are many nominal CONTENT: Aglyptodactylus Boulenger , subgenera that require reformulation as well 1919 ‘‘1918’’; Boophis Tschudi, 1838 ; Lal­ (Raxworthy, Grant, and Faivovich, in prepiostoma Glaw, Vences, and Böhme, 1998; aration). For instance, Vences et al. (2002) Mantella Boulenger, 1882 ; ‘‘ Mantidactylus ’’ revised the species of the ‘‘ Mantidactylus ’’ Boulenger, 1895. subgenus Laurentomantis and presented ev­ CHARACTERIZATION AND DIAGNOSIS: Man­ idence in their resulting tree of the paraphyly tellids are small to medium­size terrestrial or of ‘‘ Mantidactylus ’’ with respect to Mantella , arboreal frogs, predominantly found in semi­ the paraphyly of the subgenus Brygoomantis , arid to wet forested habitats. Although most and the polyphyly of Guibemantis and Geare drab or cryptically colored, species of phyromantis, as well as a lack of evidence Mantellini in particular are brightly colored. for either paraphyly or monophyly of Pan­ Life history is varied, from the usual biphasic danusicola. Much remains to be done, and life history with aquatic eggs and feeding we cannot recommend the use of subgenera tadpoles ( Boophis ) to nidicolous larvae (e.g., within ‘‘ Mantidactylus ’’ until the inconsismany Mantidactylus ). At least some (e.g., tency of taxonomy with phylogeny is ad­ Mantidactylus eiselti ) have direct develop­ dressed within that group.

ment. Most species lay eggs away from wa­ Pseudophilautus Laurent, 1943 , was ter, in some cases in a suspended nest from placed in the synonymy of Philautus by R.F. which the tadpoles drop into water (Glaw Inger (In Frost, 1985). This was accepted by

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 245

Dubois (1999b: 5) although the assignment dialis only (Haas 96.5); (5) free basihyal abto Mantellidae by Laurent (1986) has not sent (Haas 105.0); (6) commissura proximalbeen directly challenged through discussion is II present (Haas 110.1); and (7) commisof evidence. A second look is warranted. sura proximalis III present (Haas 111.1).

SYSTEMATIC COMMENTS: Taxonomic deci­ [253] FAMILY: RHACOPHORIDAE HOFFMAN , sions taken here are guided by our results

1932 (1858) (figs. 50, 65), the DNA sequence study of Polypedatidae Günther, 1858b: 346. Type genus: J.A. Wilkinson et al. (2002; fig. 48 ) and the Polypedates Tschudi, 1838 . essentially data­free tree of Delorme et al. Rhacophoridae Hoffman, 1932: 581 . Type genus: (2005; fig. 49 ), which was presented along Rhacophorus Kuhl and Van Hasselt, 1822 . with a system of morphological differentia Philautinae Dubois, 1981: 258. Type genus: Phi­ that delimited a number of monophyletic and lautus Gistel, 1848. paraphyletic groups, seemingly without ref­ Buergeriinae Channing, 1989 . Type genus: Buer­ erence to the tree itself. Results of the three geria Tschudi, 1838.

have basic agreements.

IMMEDIATELY MORE INCLUSIVE TAXON: Buergeriinae Channing, 1989 , may be rec­ [244] Rhacophoroidea. ognized for Buergeria and Rhacophorinae SISTER TAXON: [246] Mantellidae . Hoffman, 1932 (1858), for the remaining RANGE: Tropical sub­Saharan Africa; rhacophorines, as was suggested by Chan­ South India and Sri Lanka; Japan; northeast­ ning (1989) and as diagnosed by J.A. Wilern India to eastern China south through the kinson et al. (2002). We cannot subscribe to Philippines and Greater Sundas; Sulawesi. the tribal taxonomy of Delorme et al. (2005) CONTENT: Aquixalus Delorme, Dubois , because their Philautini is not monophyletic Grosjean, and Ohler, 2005 (see Systematic on their own figure ( fig. 49 ), and because the Comments); Buergeria Tschudi, 1838 ; Chi­ evidence in support of their tree was largely romantis Peters, 1854 (including Chirixalus undisclosed.

Boulenger, 1893; see Systematic Comments); On the basis of our results, and the studies Feihyla new genus (see Systematic Com­ of J.A. Wilkinson et al. (2002) and Delorme ments); Kurixalus Ye, Fei, and Dubois, 1999 et al. (2005), two problems of generic delim­ (see Systematic Comments); Nyctixalus itation appear to persist in the taxonomy. The Boulenger, 1882; Philautus Gistel, 1848 ; Po­ first of these, the paraphyly/polyphyly of lypedates Tschudi, 1838; Rhacophorus Kuhl ‘‘ Rhacophorus ’’ is beyond the scope of this and Hasselt, 1822; Theloderma Tschudi , paper; more taxa need to be analyzed before 1838. this problem can be addressed. The second CHARACTERIZATION AND DIAGNOSIS: Al­ problem is that nominal ‘‘ Chirixalus ’’ seemthough a few groups are primarily terrestrial, ingly falls into four generic units. We can rhacophorids are predominantly treefrogs, help correct the problems surrounding the sharing with basal ranids expanded digital polyphyly/paraphyly ‘‘ Chirixalus ’’, although pads and with mantellids the characteristic of the phylogenetic position of many species of intercalary phalangeal elements. Most spe­ both ‘‘ Chirixalus ’’ and nominal Philautus cies have T­shaped terminal phalanges. Sev­ needs to be evaluated.

eral larval characters that optimized on this (1) Kurixalus Fei, Ye, and Dubois (in Fei, branch may actually be synapomorphies of 1999). As noted in ‘‘Results’’, we apply this Rhacophoroidea, or some part of Rhaco­ name to a taxon that includes K. eiffingeri phoridae: (1) anterior insertion of m. subar­ and K. idiootocus , which is diagnosed by our cualis rectus II–IV on ceratobranchial II molecular evidence (see appendix 5, branch (Haas 37.1); (2) larval m. levator mandibulae 256). We provisionally include K. verrucoexternus present as two portions (profundus sus, which Delorme et al. (2005), without ev­ and superficialis; Haas 54.1); (3) posterior idence or discussion, figured as the sister taxdorsal process of pars alaris expanded ter­ on of Kurixalus eiffingeri 1 K. idiootocus . as

minally, almost rectangular in lateral view (These authors included idiootocus and ver­ (Haas 89.1); (4) cartilaginous roofing of the rucosus without discussion in their new polycavum cranii composed of taeniae tecti me­ phyletic/paraphyletic ‘‘ Aquixalus ’’, even 246 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

they illustrated these species as being in an other species with this taxon will require exclusive monophyletic group with Kurixal­ considerable additional work.

us eiffingeri ). Under this concept, there are Although ‘‘ Chirixalus ’’ palpebralis has currently no identified morphological syna­ been demonstrated to be phylogenetically pomorphies of Kurixalus , because the pur­ distinct (J.A. Wilkinson et al., 2002; Delorme ported synapomorphies associated with Ku­ et al., 2005) and deserving a new generic rixalus eiffingeri (well­developed prepollex name, the status of presumably closely relat­ and oophagus tadpoles) are not exhibited in ed species ‘‘ Chirixalus ’’ romeri and ‘‘ C. ’’ K. idiootocus or K. verrucosus (Kuramoto ocellatus of the ‘‘ Philautus ’’ palpebralis and Wang, 1987; Ziegler and Vences, 2002; group of Fei, 1999) remains an open ques­ Matsui and Orlov, 2004). Excluding ‘‘ Aquix­ tion, although no evidence so far has sugalus ’’ idiootocus and ‘‘ A. ’’ verrucosus from gested that these species form a monophy­ ‘‘ Aquixalus ’’, we suggest, renders Aquixalus letic group. Morphological evidence provid­ (sensu stricto) monophyletic (see below), if ed by Delorme et al. (2005) differentiating we assume that the tree of Delorme et al. their Rhacophorini (including ‘‘ Chirixalus ’’ (2005) survives testing by evidence. palpebralis on their tree) and Philautini (a (2) Feihyla new genus (type species: Phi­ paraphyletic group that on their tree includes lautus palpebralis Smith, 1924 . Etymology: ‘‘ Philautus ’’ gracilipes [5 Aquixalus graci­ Fei Liang 1 hyla [Greek: vocative form of lipes]), suggests that Aquixalus (including Hylas, a traditional generic root for treefrogs] ‘‘ Chirixalus ’’ gracilipes ) is not close to Feihto commemorate the extensive contributions yla (see discussion below under Aquixalus ). to Chinese herpetology by Fei Liang). J.A. (3) Chiromantis Peters, 1854 , and Chirix­ Wilkinson et al. (2002) found his exemplar alus Boulenger, 1893. A third unit is the cluster of species paraphyletic with respect of the ‘‘ Philautus ’’ palpebralis group of Fei to Chiromantis . The paraphyly of Chirixalus (1999), ‘‘ Chirixalus ’’ palpebralis , to be the (sensu stricto) with respect to Chiromantis sister taxon of a group composed of all rha­ was not a surprise to us. J.A. Wilkinson et cophorids except Buergeria . Delorme et al. al. (2002) had suggested that Chirixalus do­ (2005) placed ‘‘ Chirixalus ’’ palpebralis in riae is the sister taxon of Chiromantis , and their Rhacophorini, which otherwise corre­ that Chirixalus vittatus is close to Polypesponds to a monophyletic group recovered dates (compare their results with ours, which by us and by J.A. Wilkinson et al. (2002). In are based on substantially more data). We fact, this is the major point of disagreement place Chirixalus Boulenger, 1893 , into the between J.A. Wilkinson et al. (2002) and De­ synonymy of Chiromantis Peters, 1854 , to lorme et al. (2005). What is clear is that correct this paraphyly. (See appendix 7 for ‘‘ Chirixalus ’’ palpebralis is not in a mono­ new combinations that extend from this phyletic group with Chirixalus (sensu stric­ change and appendix 5 for molecular synato), nor obviously associated closely with pomorphies.)

any other generic grouping. For this reason (4) Aquixalus Delorme, Dubois, Grosjean , we have named Feihyla to recognize its dis­ and Ohler, 2005. We recognize a monophytinctiveness. We cannot construe Feihyla to letic Aquixalus (i.e., Aquixalus sensu Delorthe ‘‘ Philautus ’’ palpebralis group of Fei me et al., 2005, but excluding ‘‘ Aquixalus ’’ (1999) because the diagnosis of this group is idiootocus and ‘‘ Aquixalus ’’ verrucosus ; that insufficient to distinguish it from many other is, without the molecular synapomorphies of species outside of China (i.e., Fei, 1999, branch 256—see above). Delorme et al diagnosed his ‘‘ Philautus ’’ palpebralis group (2005) diagnosed this taxon (although we do as ‘‘ Philautus ’’ from China, with an X not know which of the listed species they or)(shape on the dorsum and lacking vo­ actually evaluated for these characters), but merine teeth), such as Aquixalus gracilipes our exclusion of Kurixalus idiootocus (and and A. supercornutus ; see discussion below). provisionally K. verrucosus ) from Aquixalus

We therefore diagnose Feihyla by the char­ on the basis of the molecular synapomoracters for the species ‘‘ Philautus ’’ palpe­ phies that place Kurixalus distant from bralis provided by Fei (1999). Association of Aquixalus should render Aquixalus mono­

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 247

phyletic if the tree provided by Delorme et rixalus verrucosus , so this diagnosis must be al. (2005) is correct. We suggest, on the basis largely or entirely based on plesiomorphies, of the tree provided by Delorme et al (2005), with the nominal subgenus Aquixalus being that the morphological similarities shared by those members of Aquixalus that do not share Kurixalus and Aquixalus are plesiomorphic. the apomorphies of Gracixalus . Detailed We follow the recognition by Delorme et analysis of disclosed evidence is necessary.

al. (2005) of a putatively monophyletic sub­

genus Gracixalus for ‘‘ Philautus ’’ gracilipes [269] SUPERFAMILY: RANOIDEA RAFINESQUE, Bourret, 1937 , and ‘‘ Philautus ’’ supercor­ 1814

nutus Orlov, Ho, and Nguyen, 2004 (not

IMMEDIATELY MORE INCLUSIVE TAXON: studied by us). The morphological diagnosis

[244] Aglaioanura new taxon.

of Gracixalus (spines on the upper eyelid,

S: [245] Rhacophoroidea ISTER TAXON

rictal gland connected to the mouth, foot

Hoffman, 1932 (1858).

very thin, two outer palmar tubercles, white

RANGE: Worldwide temperate and tropical spot on snout tip of tadpole, five pairs of pre­

environments, except for southern Australia, lingual papillae on the tadpole, crescent­

New Zealand, Seychelles, and southern shaped crest on the tadpole) purportedly sep­

South America.

arates it from the nominate subgenus Aquix­

C: [270] Nyctibatrachidae BlomONTENT

alus, but the absence of adequate published

mers­Schlösser, 1993, and [272] Ranidae Raftadpole descriptions suggest that this diag­

inesque, 1814.

nosis should be treated as provisional (Bain

C: Mor­ HARACTERIZATION AND DIAGNOSIS

and Nguyen, 2004; Matsui and Orlov, 2004;

phological synapomorphies for Ranidae (see Delorme et al., 2005). Although Gracixalus

below) may actually optimize at this level. can be separated from Feihyla palpebralis

Regardless, the molecular data are decisive (the latter in parentheses): snout triangularly

in support of this taxon (appendix 5).

pointed (obtusely pointed); skin translucent

(not translucent); small white tubercles along

[270] FAMILY: NYCTIBATRACHIDAE

the head, anal region, and large conical tu­ BLOMMERS­SCHLÖSSER, 1993

bercles on upper eyelid (all absent), these

characters do not unambiguously separate Nyctibatrachinae Blommers­Schlösser, 1993: 211 . Gracixalus from ‘‘ P. ’’ romeri, ‘‘ P. ’’ ocella­ Type genus: Nyctibatrachus Boulenger, 1882 .

Lankanectinae Dubois and Ohler, 2001: 82 . Type tus, the other members of the ‘‘ P. ’’ palpe­

genus: Lankanectes Dubois and Ohler, 2001 . bralis group of Fei (1999). The placement of New synonym.

these two species, as well as higher level re­

lationships will be dependent upon a rigorous IMMEDIATELY MORE INCLUSIVE TAXON: phylogenetic analysis. [269] Ranoidea Rafinesque, 1814 .

Although we cannot reject the putative SISTER TAXON: [272] Ranidae Rafinesque , monophyly of the subgenus Aquixalus (in­ 1814.

cluding the type species A. odontotarsus , as RANGE: Sri Lanka and India.

well as A. ananjevae , A. baliogaster , A. bis­ CONTENT: Nyctibatrachus Boulenger , acculus, A. carinensis , and A. naso ; modified 1882; Lankanectes Dubois and Ohler, 2001 . from Delorme et al., 2005), we do not see CHARACTERIZATION AND DIAGNOSIS: None any reason to recognize it, either, until the of our analyzed morphology optimizes on relevant phylogenetic data are published by this branch, although the molecular data are the original authors. According to Delorme decisive. See appendix 5 for list of unambig­ et al. (2005), the morphological diagnosis of uous molecular synapomorphies.

Aquixalus (webbing on feet not extending to SYSTEMATIC COMMENTS: Nyctibatrachidae toes, rictal gland not connected to mouth, in our sense brings two genera together, Nycfoot very thick, one outer palmar tubercle, tibatrachus, with a median lingual process concavity on tadpole snout in lateral view, (unknown polarity), digital discs present not

),

four pairs of prelingual papillae in tadpole, (plesiomorphic), femoral glands present (unmedian crest in tadpole triangular shaped, known polarity), and lateral line system 180– 240 eggs per clutch) also applies to Ku­ persisting into adulthood (plesiomorphic 248 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

and Lankanectes , with no median lingual Amerana Dubois, 1992 ; Aurorana Dubois , process, digital discs absent, femoral glands 1992; Pseudoamolops Jiang, Fei, Ye, Zeng , absent, and lateral line system persisting into Zhen, Xie, and Chen, 1997; and Pseudorana adulthood (Dubois et al., 2001). They are ar­ Dubois, 1992); Sanguirana Dubois, 1992 ; ranged in a single family to avoid the taxo­ Staurois Cope, 1865 ; Sylvirana Dubois , nomic redundancy of having monotypic (and 1992 (including Papurana Dubois, 1992 , therefore uninformative) family­group and Tylerana Dubois , 199232). (See Systemnames. atic Comments.)

CHARACTERIZATION AND DIAGNOSIS: Al­ [272] FAMILY: RANIDAE RAFINESQUE, 1814 though Haas (2003) included only two ranids

in his study, Sylvirana nigrovittata and Rana Ranaridia Rafinesque, 1814: 102 . Type genus: temporaria , characters that optimize on their Ranaridia Rafinesque, 1814

Limnodytae Fitzinger, 1843: 31. Type genus: Lim­ subtending branch are candidates as synanodytes Duméril and Bibron, 1841. pomorphies for Ranidae : (1) posterolateral Amolopsinae Yang, 1991a: 172. Type genus: projections of the crista parotica absent (Haas Amolops Cope, 1865 . 67.0); and (2) processus branchialis closed

(Haas 114.1). Denser sampling should test IMMEDIATELY MORE INCLUSIVE TAXON: this proposition. These characters may actu­ [269] Ranoidea Rafinesque, 1814 . ally optimize on Ranoides. Regardless, the SISTER TAXON: [270] Nyctibatrachidae molecular data are decisive (see appendix 5). Blommers­Schlösser, 1993. SYSTEMATIC COMMENTS: As noted in ‘‘Re­ RANGE: Temperate and tropical Africa and sults’’, Batrachylodes is transferred defini­ Eurasia through Indonesia to northern Aus­ tively to Ceratobatrachidae and Amietia (intralia, North America, Central America, and cluding Afrana ) and Strongylopus are transnorthern South America. ferred to Pyxicephalidae . For discussion of CONTENT: Amolops Cope, 1865 (including these taxa see those familial accounts.

Amo Dubois, 1992 ); Babina Thompson, As noted in the ‘‘Review of Current Tax­ 1912 (including Nidirana Dubois, 1992 ); onomy’’, the sections and subsections of Clinotarsus Mivart, 1869 ; Glandirana Fei , ‘‘ Rana ’’ (sensu lato) provided by Dubois Ye, and Huang, 1991 ‘‘1990’’ 32 (including (1992) do not inform about evolutionary re­ Rugosa Fei, Ye, and Huang, 1991 ‘‘1990’’); lationships, so for this discussion and the tax­ Hydrophylax Fitzinger, 1843 (including Am­ onomic remedies we suggest, we will focus nirana Dubois, 1992, and Chalcorana Du­ on genera and subgenera. The discussion that bois, 1992); Hylarana Tschudi, 1838 ; Huia follows addresses the generic taxonomy that Yang, 1991 (including Eburana Dubois , we recommend (moving from top to bottom 1992; Bamburana Fei, Ye, Jiang, Xie, and of Ranidae [new taxonomy] in figure 71 , al­ Huang, 2005; Odorrana Fei, Ye, and Huang , though addressing other genera and problems 1991 ‘‘1990’’); Humerana Dubois, 1992 ; in passing).

Lithobates Fitzinger, 1843 (including Aquar­ Staurois Cope, 1865 : We accept Staurois ana Dubois, 1992 ; Pantherana Dubois , as a genus, although we note that evidence 1992; Sierrana Dubois, 1992 ; Trypheropsis for this taxon’s monophyly is equivocal and Cope, 1868; Zweifelia Dubois, 1992 ); Mer­ requires testing. The traditional diagnosis of istogenys Yang, 1991; Nasirana Dubois , Staurois —digital discs broader than long; T­ 1992; Pelophylax Fitzinger, 1843 ; Pterorana shaped terminal phalanges with horizontal Kiyasetuo and Khare, 1986; Pulchrana Du­ arm longer than longitudinal arm; outer bois, 1992; Rana Linnaeus, 1758 (including metatarsals separated to base but webbed;

nasals small separated from each other and 32 Dubois (1999a: 91) considered Glandirana Fei, Ye , frontoparietal; omosternal style not forked and Huang, 1991, to have priority over Rugosa Fei, Ye , (Boulenger, 1918); and lacking a raised ab­ and Huang, 1991, and Sylvirana Dubois, 1992 , to have dominal sucker disc on larva (Inger, 1966)—

priority over Papurana Dubois, 1992 , and Tylerana Du­

are plesiomorphic for Ranidae . Although 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 249

with deep, cup­like labial parts; upper lip of the large number of ranid species whose oral disc with two continuous rows of papil­ adults are morphologically similar to those of

lae; lower lip with one broad continuous Staurois , but whose larvae remain undeband of papillae; Inger, 1966), the diagnostic scribed.

value of these characters is unknown due to [274] Hylarana Tschudi, 1838 : We asso­

250 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

ciate our exemplars of Hylarana Tschudi , which are reported to not bear humeral 1838 ( H. erythraea , the type species, and H. glands 33) is done on the assumption that taipehensis ), as well as of ‘‘ Sylvirana ’’ some of the molecular apomorphies of this guentheri , with the generic name Hylarana . taxon are synapomorphies of Hydrophylax in Although these two units were assigned, re­ the sense of including the species that we did spectively, to the no­humeral­gland (Hylar­ not study. On the basis of evidence presented ana ) and humeral­gland subsections (Hydro­ by Matsui et al. (2005), we place Chalcorana phylax ) of Dubois (1992), our data suggest hosii in our Huia . Chalcorana is likely strongly that the humeral gland is convergent broadly polyphyletic, but without evidence in ‘‘ S. ’’ guentheri and Sylvirana (sensu stri­ of the remainder’s placement we provisionco) or that the presence of the structure has ally regard them as close to Chalcorana been missed in a widespread way because of chalconota , the type­species of Chalcorana . the lack of detailed morphological study (in­ We could have retained Chalcorana as a gecluding dissections). Hylarana (including nus, but it is clear that, as data emerge, the ‘‘ Sylvirana ’’ guentheri and H. macrodactyla , species in this nominal taxon will be asthe third species of Hylarana sensu Dubois , signed to Hydrophylax , Sylvirana , and likely 1992) lacks dermal glands in the larvae, a others as well. This is not a satisfactory socharacter that appears to optimize on the sis­ lution to the problem of trying to sort ter branch of Hylarana . The vocal sac con­ through this morass, but it is the only pracdition is variable among species of Hylarana , tical solution available to us at present. with ‘‘ S. ’’ guentheri possessing gular pouch­ We retain Humerana Dubois, 1992 , and es and H. taipehensis and H. erythraea lack­ Pulchrana Dubois, 1992 , as nominal genera ing gular pouches. This character is highly only because we did not study these humeralhomoplastic throughout the ranid portion of gland­bearing genera. Future work should our tree. We take the molecular apomorphies test the hypothesis that the remaining species for branch 274 (appendix 5) to be synapo­ of the ‘‘humeral­gland group’’ constitute a morphies of Hylarana . monophyletic unit. The results of Matsui et

We are unable to diagnose Hylarana on al. (2005; fig. 46 ) suggest that Humerana ulthe basis of morphology. We did not study, timately will be assigned to Hylarana . and so cannot document, the phylogenetic [280] Sylvirana Dubois, 1992 : Our results position of H. macrodactyla . Thus, our as­ demonstrate the polyphyly of nominal Sylsociation of this species with Hylarana re­ virana (see discussion of ‘‘ S. ’’ guentheri unquires testing. Similarly, we do not know der discussion of Hylarana ) and the parawhich other species may be included in this phyly of the major group of nominal Sylvirhistorically ambiguously diagnosed genus. ana (including its type species, S. nigrovit­

[278] Hydrophylax Fitzinger, 1843 (in­ tata). To remedy the demonstrated polyphyly cluding Amnirana Dubois, 1992 , and Chal­ of Sylvirana , we transfer ‘‘ S. ’’ guentheri into corana Dubois, 1992): We associate our ex­ Hylarana Tschudi, 1838 (see above). To reemplars of humeral­gland­bearing genera lieve the paraphyly of remaining Sylvirana , ( Hydrophylax and Amnirana ), as well as the we place Papurana Dubois, 1992 , and Tyimbedded Chalcorana , with the generic lerana Dubois, 1992, into the synonymy of name Hydrophylax Fitzinger, 1843 . Chan­ Sylvirana Dubois, 1992 . Although it is clear ning (2001) had already considered the Af­ on the basis of molecular data that ‘‘ S. ’’ rican member of Hydrophylax (H. galamen­ guentheri is not in the clade containing S. sis ) to be in Amnirana , along with other Af­ nigrovittata (the type species of Sylvirana ), rican Hylarana ­like frogs. Our association of it is also not clear how many species of nomthe type species of Hydrophylax , H. malabarica (unstudied by us), with the clade of 33 Possession of humeral glands can be a difficult studied terminals requires testing, of course, characteristic to assess due to level of development, and as does the association of the unstudied their presence may be apparent only on dissection.

Therefore, any statement that humeral glands are absent members of these nominal taxa. The associ­ really requires that a dissection has been made. Dubois ation of unstudied members of Amnirana , (1992) did not mention whether he had made such dis­ Hydrophylax , and Chalcorana (some of sections.

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 251

inal Sylvirana are associated with ‘‘ S. ’’ Kiyasetuo and Khare, 1986: We provisionguentheri. We take the most falsifiable po­ ally retain Sanguirana Dubois, 1992 , and sition—that only ‘‘ S. ’’ guentheri is far from Pterorana Kiyasetuo and Khare, 1986 (both Sylvirana nigrovittata —and suggest that unstudied by us) as genera, owing to the amcareful study is needed. biguous nature of their putative synapomor­

Meristogenys Yang, 1991 , Clinotarsus Mi­ phies (both genera are Hylarana ­like forms). vart, 1869, and Nasirana Dubois, 1992 : Our Sanguirana sanguinea (type species of Sanresults place Meristogenys as the sister taxon guirana) has a tadpole with characters shared of Clinotarsus (as found by Roelants et al., with Meristogenys , Clinotarsus , and Altir­ 2004; fig. 35), and far from both Amolops ana : a moderate to high number of labial ker­ and Huia , to which it was considered to be atodont rows (4–6/4–5); upper lip with diclosely related by Yang (1991b) and Dubois vided keratodont rows; and dermal glands on (1992). Besides the molecular evidence, Cli­ the head and body; and ventral portions of notarsus shares several larval characters with the body and tail fins (Alcala and Brown, Meristogenys : (1) dermal glands on the flank; 1982). Pterorana khare (tadpole unknown) is (2) increased numbers of rows of labial ker­ distinguished from other ranid frogs by the atodonts (5–9/ 5–10 in Meristogenys and 6– large, fleshy folds on the flanks and thighs 8/ 6–8 in Clinotarsus ; over 1–5/ 2–8 in Amo­ and over the vent that extend away from the lops and Huia ; Boulenger, 1920: 132–133; body when the frog is under water (Kiyase­ Chari, 1962; Yang, 1991b; Hiragond et al., tuo and Khare, 1986). 2001); and (3) upper labial keratodont rows Amolops Cope, 1865 , and Amo Dubois , with a medial gap. Unlike Clinotarsus , but 1992: The phylogenetic association of Amolike Amolops , Huia , and (superficially) Pseu­ lops, Meristogenys , and Huia (Yang, 1991b; doamolops, Meristogenys have a raised ab­ Dubois, 1992), as noted in ‘‘Results’’ and in dominal sucker in the larvae (Kuramoto et the discussion above of Meristogenys , was al., 1984; Yang, 1991b; Jiang et al., 1997). rejected. Further, the association of Pseu­

Clinotarsus lacks the obvious synapomor­ doamolops Jiang et al., 1997, suggested by phies associated with Meristogenys (a raised, Kuramoto et al. (1984) and Fei et al. (2000) sharply defined abdominal sucker in the lar­ is also rejected, suggesting that in each case vae, ribbed jaw sheaths, and upper or both the ventral sucker on the larvae is nonhojaw sheaths divided (Yang, 1991b). Because mologous and should be considered indepenmost of the species of Meristogenys , like dently apomorphic in each lineage. Kuramost Hylarana ­like species (sensu lato), moto et al. (1984) provided morphological have not been sampled and may be involved evidence that the ventral sucker disc on the with this group, we retain both Clinotarsus larvae of Amolops is not homologous with and Meristogenys as genera. that of ‘‘ Pseudorana ’’ sauteri : the edge of

Nasirana alticola (not studied by us) may the disc is sharply defined in Amolops (not be allied with Clinotarsus , as their larvae so in sauteri ); the m. diaphragmatobranchialshare two possible synapomorphies: (1) large is medialis engages the floor of the sucker to size; and (2) supracaudal glands (Grosjean et generate negative pressure in Amolops (musal., 2003). Furthermore, Nasirana shares cle does not communicate with sucker in with Meristogenys and Clinotarsus other lar­ sauteri ); and inframarginal U­shaped band of val characters of uncertain polarity: multiple keratinized material on the sucker in Amo­ (3–7) medially divided upper labial kerato­ lops (absent in sauteri ). Regardless, Kuradont rows; high numbers of labial keratodont moto et al. (1984) suggested a close relationrows (7–8: 7–8); and presence of dermal ship of sauteri to Amolops . glands on the flanks of the body (Yang, The status of Amo Dubois, 1992 (not stud­ 1991b; Hiragond et al., 2001; Grosjean et al., ied by us), is arguable. Dubois (1992) sug­ 2003). Nasirana can be distinguished from gested that Amo is unique among Amolops in all other frogs by a fleshy prominence on the having axillary glands in both sexes and an the

snout of the male. As with Clinotarsus , we outer metatarsal tubercle (a character pleprovisionally retain Nasirana as a genus. siomorphic at the base of the ranids), but Sanguirana Dubois, 1992 , and Pterorana outer metatarsal tubercle is nevertheless pre­

252 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

sent in Amolops nepalicus 34 and A. torrentis low glands; axillary glands and distal femo­ (after Yang, 1991b). Amo lacks the charac­ ral glands densely packed, forming a roll; teristics of both Huia and Meristogenys (tibia and intermittent longitudinal ridges, densely elongate; having lateral dermal glands on the covered with small tubercles on the dorsum larvae; high number of larval keratodont (Fei et al., 1991 ‘‘1990’’). It shares with Perows on the lower lip) but otherwise shares lophylax a very low number of labial keraone apomorphy with Amolops (sensu stricto) todont rows in larvae (likely plesiomorphic in our topology: first metacarpal greater than on our topology). Jiang and Zhou (2005; half the length of the second. So, rather than their fig. 1), with different taxon sampling, suggest that a sucker developed on the venter found Glandirana to be the sister taxon of of the larvae five times in ranids (rather than Rugosa (not studied by us, but placed by Duthe four events currently required by our to­ bois, 1992, in his section Pelophylax ), and pology) we regard Amo as a synonym of phylogenetically distant from their samples Amolops . of Pelophylax ( P. hubeiensis and P. nigro­ We found nominal Amolops to be poly­ maculata ). phyletic (figs. 50, 65). In this case, the larva Glandirana and Rugosa share the followof Amolops chapaensis is unknown (Yang, ing characteristics that appear to be synapo­ 1991b), and that species had been assigned morphic (on our tree and on that of Jiang and to Amolops on the basis of having an adult Zhou, 2005): entire body of tadpole covered morphology more similar to Amolops than to in glands; digital discs absent in adults; and Hylarana (i.e., no humeral glands and pres­ dorsum densely covered with longitudinal, ence of gular pouches in males; after Inger, tubercular skin ridges in adults (Stejneger, 1966: 257), rather than its having the larval 1907: 123–126; Okada, 1966; Ting and T’sai, synapomorphies of Amolops . We transfer this 1979; Fei et al., 1991 ‘‘1990’’; Fei et al., species out of Amolops and into another ge­ 2005: 132–138). There are morphological nus below. (See discussion of Huia , Odor­ differences between the two genera (Okada, rana , and Eburana ). Although we obtain 1966; Fei et al., 1991 ‘‘1990’’; Fei et al., Amolops as the sister taxon of Pelophylax , 2005: 132–138; Stejneger, 1907: 123–126; we are unaware of any morphological syna­ Ting and T’sai, 1979): sternal cartilage pomorphy uniting these groups (see appen­ forked posteriorly in Glandirana [deeply dix 5, branch 287). notched in Rugosa ]; toes half­webbed, reach­ [288] Pelophylax Fitzinger, 1843 : We re­ ing the second subarticular tubercle on toe strict the generic name Pelophylax to the IV in Glandirana [fully webbed to beyond subgenus Pelophylax of Dubois (1992). We second subarticular tubercle on toe IV in Ruare unaware of any morphological synapo­ gosa]; skin densely covered in granular yelmorphy for this group, although the molec­ low glands, as well as axillary and distal ular data are seemingly decisive (see appen­ femoral glands densely packed, forming a dix 5, branch 288). roll in Glandirana [prominent glands only Glandirana Fei, Ye, and Huang, 1991 behind tympanum in Rugosa ]). However, ‘‘1990’’, and Rugosa Fei, Ye, and Huang , none of these characters is obviously in con­ 1991 ‘‘1990’’: Glandirana minima is the sole flict with Glandirana 1 Rugosa forming a species in its nominal genus (formerly a sub­ monophyletic group. In light of this evigenus of the section Hylarana , subsection dence, we recognize this clade as one genus, Hylarana : Dubois, 1992). It is diagnosed by Glandirana , placing Rugosa into synonomy. having skin densely covered in granular yel­ Rugosa rugosa , the type species of Rugosa ,

should be included in subsequent phyloge­ 34 Dubois (2000: 331; 2004a: 176) suggested, on the netic analysis to test this hypothesis. basis of examination of the holotype, this taxon is syn­ [291] Babina Thompson, 1912 , and Nidionymous with Amolops formosus but did not provide rana Dubois, 1992 : Nidirana Dubois, 1992 , any discussion regarding the differences itemized in the has been associated with Babina Thompson ,

original description or the diagnostic differences noted

1912 (unstudied by us) on the basis of two 2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 253

deposition in water­filled nests of terrestrial raised, sharply defined abdominal sucker in burrows or open puddles (Pope, 1931: 536– the tadpole (Yang, 1991b; see discussion of 538; C.­C. Liu, 1950: 258–260; Kuramoto, Meristogenys and Amolops above). Beyond 1985; Dubois, 1992: 154–156; Chou, 1999: this structure, the only characters uniting 398–399). Babina is further diagnosable Huia with Amolops and Meristogenys are from Nidirana on the basis of the male hav­ ventral and postorbital glands of the larvae. ing a spine on the prepollex (absent in Ni­ None of these characters is present in Odordirana; Okada, 1966; Kuramoto, 1985; rana grahami , the only other member of this Chou, 1999). Nidirana , however, has no clade whose tadpole is known.

characters that suggest that it is monophylet­ We know of no morphological synapoic with respect to Babina (Dubois, 1992; morphy that unites this clade (branch 292), Chou, 1999). For this reason, although a sub­ but our molecular data are decisive for its genus Babina (the group with the large pre­ being a monophyletic group (see appendix pollical spine) could be employed, the name 5). We therefore apply a single generic name. Nidirana applies to no monophyletic group The oldest available name from this group of that can be identified at this time. We there­ species is Huia Yang, 1991b (published 18 fore transfer all members of Dubois’ subge­ February, 1991; the publication containing nus Nidirana to the genus Babina . Odorrana did not appear until at least March [292] Huia Yang, 1992 , Odorrana Fei, Ye , of 1991; Fei et al., 1991 ‘‘1990’’). We there­ and Huang, 1991 ‘‘1990’’, Bamburana Fei et fore place ‘‘ Amolops ’’ chapaensis ; Eburana al., 2005, ‘‘ Amolops ’’ chapaensis , and Ebur­ Dubois, 1992; and Odorrana Fei, Ye , and ana Dubois, 1992 : Although our molecular Huang, 1991 ‘‘1990’’, into the synonymy of evidence capturing this clade of Himalayan Huia Yang, 1991 .

and Southeast Asian cascade­dwelling spe­ We recognize that this taxonomy is probcies is unambiguous (see appendix 5, branch lematic for two reasons. First, we did not in­ 292), insufficient sampling, the lack of mor­ clude any of the types of the nominal genera phological data, and the concomitant taxo­ in this study. Thus, the assigned name may nomic confusion surrounding these taxa pre­ be inappropriate. Indeed, Huia nasica may sented us with a significant taxonomic chal­ not be closely related to Huia cavitympanum lenge. ‘‘ Amolops ’’ chapaensis is embedded Boulenger, 1893 (the type species of Huia in our Huia – Eburana – Odorrana clade, but and not studied by us). The association with its assignment to Amolops was done on the Huia nasica of a tadpole with a raised, sharpbasis of overall similarity (see discussion in ly defined abdominal sucker and ventral and Amolops section), and it is clearly not part of postorbital glands of the larvae was based on that genus. There is no known morphological one specimen (C.­C. Liu and Hu, 1961). synapomorphy linking species of Odorrana, Yang (1991b) cast doubt on this assignment as its purported synapomorphy, colorless when he reported that a ‘‘tadpole from Menspines on chest of the male, is also known in yang assigned to H. nasica by Liu and Hu Huia nasica (B.L. Stuart and Chan­ard, (1961), is certainly Huia even if not larval 2005) and species of at least two other gen­ H. nasica ’’. Our grouping of H. nasica withera (i.e., some Chalcorana and at least Ba­ in a clade of Odorrana and Eburana might bina caldwelli [R. Bain, personal obs.]), and be evidence that nasica is not a member of is absent in many species of Odorrana sensu Huia . And second, our small sample size (4 Fei et al. (1991 ‘‘1990’’; see discussion in species, only 2 of which have known tad­ ‘‘Review of Current Taxonomy’’). Similarly, poles) from this large, undiagnosed group of there is no evidence suggesting that Eburana species (minimum 36 species; Frost, 2004) is monophyletic, because its putative syna­ may speak to an oversimplification of the repomorphy, unpigmented eggs, is shared by lationships among these taxa.

at least some species of three other genera Whereas both of these problems are real (e.g., Chalcorana , Odorrana , Amolops ; see concerns, this decision, as with all of our taxon

, discussion in ‘‘Review of Current Taxono­ onomic decisions, is a hypothesis based my’’). Huia (sensu stricto) represents a third the preponderance of the available evidence example in our tree of convergence of a which we prefer to taxonomic decisions

254 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

based on similarity groupings. As this entire the abdominal suction cup on the larvae. section of former Rana seems to have avoid­ This structure is found in sauteri alone, al­ ed detailed study, we suggest that a concerted though in a less­developed form than in effort to amass the necessary comparative Amolops , Meristogenys , and Huia (sensu morphological and molecular data is needed, stricto; Jiang et al., 1997). Tanaka­Ueno et and we interpret our results as identifying al. (1998a) suggested on the basis of 587 bp key areas for further study and not as a de­ of mtDNA that sauteri is imbedded within cisive resolution of these problems. the brown frog clade (Dubois’ subgenus

[296] Rana Linnaeus, 1758 (including Au­ Rana ). Our results corroborate this. Unlike rorana Dubois, 1992, Amerana Dubois , Amolops , Meristogenys , and Huia , both 1992, Pseudoamolops Jiang, Fei, Ye, Zeng , Pseudorana and Pseudoamolops lack dermal Zhen, Xie, and Chen, 1997, and Pseudorana glands on the larvae, which might be a syn­ Fei, Ye, and Huang, 1991 ‘‘1990’’): To ren­ apomorphy, although we do not know the der a monophyletic grouping, we place Pseu­ condition of this feature in the Rana tempordorana and Pseudoamolops as junior syno­ aria group. For our taxonomy, we relegate nyms of Rana , because they are both embed­ Pseudoamolops and Pseudorana to the synded within the same clade as Rana tempor­ onymy of Rana , which is decisively diagaria (the type species of Rana ). The nosable on the basis of molecular data (apabdominal sucker disc of the tadpole of pendix 5, branch 296). Pseudoamolops is not homologous with [301] Lithobates Fitzinger, 1843 (includthose of Amolops , Huia , and Meristogenys , ing Aquarana Dubois, 1992 , Pantherana Duall of which are distant from each other in bois, 1992, Sierrana Dubois, 1992 , Trypherour tree. opsis Cope, 1868, and ‘‘ Rana ’’ sylvatica ):

Because Amerana 1 Aurorana form the Because of the phylogenetic propinquity of sister taxon of our exemplars of a clade with Aquarana Dubois, 1992 , Lithobates Fitzin­ Rana temporaria , we also place both of these ger, 1843, Pantherana Dubois, 1992 , Siergenera as junior synonyms of Rana (sensu rana Dubois, 1992 , Trypheropsis Cope , stricto) to render a monophyletic group. 1868, ‘‘ Rana ’’ sylvatica , and Zweifelia Du­ These frogs are unusual among American bois, 1992 (the latter not studied by us, but ‘‘ Rana ’’, but otherwise similar to members placed phylogenetically in this group by Hilof Rana (sensu stricto) in retaining an outer lis and Wilcox, 2005; fig. 44 ), we place these metatarsal tubercle. taxa into their own genus, for which the old­

Dubois (1992) recognized Pseudorana as est available name is Lithobates Fitzinger , including Rana sangzhiensis , Rana sauteri , 1843. Therefore, we consider Lithobates to and R. weiningensis , characterized as lacking be a genus, within which we place Aquardermal glands in the larvae (likely a syna­ ana , Trypheropsis , Sierrana , Zweifelia , and pomorphy at this level of universality) and Pantherana as junior synonyms. Absence of having a labial keratodont row formula of 4– an outer metatarsal tubercle is a morpholog­ 7/5–8, an abdominal sucker in the larvae (al­ ical synapomorphy. (For species affected by though not as well­developed as in Amo­ this nomenclatural change see Frost, 2004, lops), digit I longer than digit II (likely ple­ and appendix 7). siomorphy), toe pads present on digit I and We considered recognizing Aquarana for toe IV; metatarsal tubercle present (plesiom­ the former R. clamitans / R. catesbeiana orphy), dorsolateral folds present; no gular group; Lithobates for the former R. palmipes pouches in males; and a chevron­shaped group; Pantherana for the R. pipiens group; mark on the anterior dorsum. Subsequently, and Zweifelia for the former R. pustulosa /R. Jiang et al. (1997) partitioned Pseudorana , tarahumarae group. However, this would with P. weiningensis staying in Pseudorana have necessitated naming a new monotypic along with johnsi and sangzhiensis , but sau­ genus for Rana sylvatica . Hillis and Wilcox teri being transferred to Pseudoamolops on (2005) also suggested, on the basis of a gen­

the basis of several features. The most dis­ erally more limited study, but much more tinctive feature is that Pseudorana (contra densely sampled within ‘‘ Rana ’’ than ours, the diagnosis of Dubois, 1992) actually lacks that ‘‘ Rana ’’ sylvatica is the sister taxon of

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 255

Aquarana . We found it to be the sister taxon thanks: Lisa Gugenheim and Merrily Sterns of the (old) Pantherana – Sierrana –Lithoba­ (Office of Government Relations) and Diane tes–Typheropsis clade. However, this result Bynum and Barbara Green (Office of Grants is weakly corroborated (due to the variable and Fellowships) were unstinting in their placement of ‘‘ R. ’’ sylvatica ; this branch has support. Eleanor Sterling (Center for Biodi­ a Bremer value of 1 and jackknife frequency versity and Conservation), initiated and proof 52%), and the results of Hillis and Wilcox moted fieldwork in Vietnam and Bolivia that (2005) therefore deserve further careful con­ resulted in the acquisition of many of the sideration. What does seem to be highly cor­ valuable tissue samples used in this study. roborated by both our data and those of Hillis Angelique Corthals and Julie Feinstein and Wilcox (2005) is that, excluding the spe­ (AMNH Monell Cryo­Collection) cooperatcies formerly assigned to Amerana and Au­ ed in last­minute tissue requests and accesrorana, all North American species currently sions. Leo Smith provided advice and supassigned to Rana form a clade. To recognize port regarding the vagaries of POY. Ho Ling this and to underscore the fact that the spe­ Poon (Center for Biodiversity and Consercies on the West Coast are more closely re­ vation) provided timely assistance with translated to Eurasian species than to other North lations of Chinese literature. Mary DeJong American species, we recognize the com­ was invaluable in providing library assispletely American group as Lithobates . (See tance. In the AMNH Herpetology Departappendix 7 for new combinations and con­ ment, Iris Calderon and Dawn Skala dealt tent.) Hillis and Wilcox (2005) provided sev­ skillfully with the large demands placed on eral new names for various clades within them by this and related projects. Denny Di­ Lithobates , but inasmuch as these were not veley provided extensive editorial and library associated with organismal characteristics support. that purport to delimit them, they are nomina Enormous assistance and encouragement nuda. was also provided from formal and informal reviewers. Maureen Donnelly, David Gower, ACKNOWLEDGMENTS Robert F. Inger, Roy W. McDiarmid, Joseph Mendelson III, Jay M. Savage, and Tom A. We thank the National Aeronautic and Titus read the entire manuscript, caught Space Administration (NASA) for significant many errors, and provided invaluable insight support of computational and molecular bi­ and suggestions; their efforts are deeply apology at the American Museum of Natural preciated. Paul Chippindale read the sala­ History. This support (NASA grants NAG5­ mander sections, caught errors, and provided 12333 and NAG5­8443 to Frost and NAG5­ timely advice. Jeffery A. Wilkinson, provid­ 13028 to Wheeler) allowed the continued de­ ed welcome advice and comments on the velopment of necessary algorithms, the soft­ various sections relevant to rhacophorid sys­ and hardware for massively parallel compu­ tematics. Richard Mayden was a great source tation of large phylogenetic trees, the of counsel and encouragement during the delarge­scale acquisition of molecular data that velopment of this study. elucidate our understanding of the evolution Grant and Frost acknowledge NSF grant and distribution of life on planet Earth, as DEB­0309226, which allowed development well as the student involvement so necessary of many of the primers used in this study and to the success of this venture. This support many of the sequences used both in the supwent far to assuring the success of this in­ ported dendrobatid research as well as in this ternational community project and we are study. During the course of this study Grant deeply grateful. Regardless, any opinions, was supported by an AMNH Graduate Stufindings, and conclusions or recommenda­ dent Fellowship, a Columbia University Centions expressed in this material are those of ter for Environmental Research and Conserthe authors and do not necessarily reflect the vation Faculty Fellowship, and NASA grant

views of the National Aeronautics and Space NAG5­13028.

Administration. Faivovich and Frost acknowledge NSF Many people in the AMNH deserve grant DEB­0407632 which supported devel­

256 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 297

opment of primers and sequences used in this Haas acknowledges support by the Deutstudy as well as the supported hylid research. sche Forschungsgemeinschaft Grant Ha Faivovich acknowledges the timely support 2323/2­1. of a Theodore Roosevelt Memorial grant. De Sá acknowledges NSF grants DEB­ During the course of this study Faivovich 0342918 and 9815787 which provided supwas supported by an AMNH Graduate Stu­ port for field work and Leptodactylus redent Fellowship and NASA grant NAG5­ search that concomitantly furthered the de­ 13028. Faivovich and Blotto thank Santiago velopment of this study. Nenda, Guido Corallo, Andres Sehinkman, Campbell gratefully acknowledges the and Diego Baldo for field assistance. support of NSF grants DEB­0102383 and Bain acknowledges NSF grant DEB­ 9705277, which allowed the acquisition of 9870232 to the Center for Biodiversity and many of the Middle American and tissue Conservation (CBC/AMNH) for financial samples used in this paper, as well as field support as well as the generous support of and collection assistance by Dwight Law­ The John D. and Catherine T. MacArthur son, Brice Noonan, Eric Smith, and Paul Us­ Foundation. Collecting and export permits tach. for Vietnam amphibians were granted by the Channing acknowledges the Tanzania Forestry Protection Department, Ministry of Commission for Science and Technology Re­ Agriculture and Rural Development, Viet­ search Permit 2002­319­ER­99­40, which nam (export permit numbers 31–98, 102–98, provided field support for the collection of 340–99, 341–99, and 174–00). Thanks are genetic samples in Tanzania. also due to Le Xuan Canh, Nguyen Quang Donnellan thanks Michael Mahony (Uni­ Truong, Ho Thu Cuc, and Khuat Dang Long versity of Newcastle), Steve Richards (South of the Institute of Ecology and Biological Australian Museum), Allen Allison (Bernice Resources, Hanoi, and Melina Laverty, CBC / P. Bishop Museum), Dale Roberts (Univer­ AMNH, for cooperation and assistance in all sity of Western Australia), Michael Tyler aspects of Vietnam fieldwork. Tissues of Bo­ (University of Adelaide), and Ken Aplin livian amphibians were collected on expedi­ (Western Australian Museum) for access to tions supported by the CBC/AMNH and the critical tissues, field support, and courtesies Center for Environmental Research and Con­ extended to him that furthered this study. servation at Columbia University, New York, Raxworthy acknowledges NSF grant in collaboration with the Museo de Historia DEB­9984496, National Geographic Society Natural Noel­Kempff Mercado, Santa Cruz, grant 5396­94, and grants from Earthwatch Bolivia, and Colección Boliviana de la Fauna, La Paz. Collection permits for Bolivian which provided field support for acquisition

of important genetic samples from Madagasmaterial were granted by el Ministerio de Desarrollo Sostenible y Planificacion de Bo­ car. livia. Nussbaum and Raxworthy gratefully ac­ Haddad gratefully acknowledges Biota­ knowledge support from NSF grants DEB­ FAPESP (01/13341­3) and CNPq for finan­ 9024505, 9322600, and 9625873, which procial support. Exportation permits of Brazilian vided funds for field research and acquisition samples were issued by CITES Lic. 081968 of tissues. BR; Autorizações de Acesso e de Remessa Nussbaum acknowledges NSF grants de Amostras de Componentes do Patrimônio DEB­0070485, 9625873, and 9917453, Genético numbers 02001002851/2004; which provided funds for the acquisition of 02001.002669/2004; permits for collection genetic samples from Madagascar and the were issued by IBAMA/RAN, licenças 057/ Seychelles. Nussbaum also thanks Greg 03 and 054/05. Haddad thanks the following Schneider for efforts in developing and for field support in the acquisition of relevant maintaining the tissue collections at UMMZ; tissues: Antonio P. Almeida, João L. Gapar­ and Ronn Altig, Michael J. Pfrender, and Ed­

ini, José P. Pombal, Jr., Luis O.M. Giasson, mund D. Brodie II, Jr. for help with field Marília T.A. Hartmann, and Paulo C.A. work in China, Madagascar, São Tome´, and Garcia. Seychelles.

2006 FROST ET AL.: AMPHIBIAN TREE OF LIFE 257

Moler thanks Barry Mansell for field as­ São Paulo, Brazil), José Núñez N. (Instituto sistance. de Zoología, Universidad Austral de Chile, Drewes thanks the NSF for grant DBI­ Valdivia, Chile), Wade Ryberg (Washington 9876766, that helped support the CAS frozen University, St. Louis), Elizabeth Scott tissue collection, which proved to be an in­ (Transvaal Museum, Pretoria, South Africa), valuable resource for this study. Tom A. Titus (University of Oregon, Eu­ Green thanks for funding the National Sci­ gene), Jens V. Vindum (California Academy ences and Engineering Research Council of Sciences, San Francisco), David B. Wake (NSERC) of Canada. (Museum of Vertebrate Zoology , University For access to critical tissues and other of California, Berkeley), and Jorge Williams courtesies with respect to this study and (Museo de la Plata, Buenos Aires, Argenticlosely related ones we thank Stevan J. Ar­ na). nold (Oregon State University, Corvallis), J.W. Arntzen (National Museum of Natural REFERENCES History, Leiden), Christopher Austin, Robb Abel, O. 1919. Die Stämme der Wirbeltiere. Klas­ T. Brumfield, Donna Dittman, and Frederick se Amphibia. Berlin and Leipzig: Walter de Sheldon (Louisiana State University Muse­ Gruyter. um of Zoology , Baton Rouge), Boris Blotto Abourachid, A., and D.M. Green. 1999. Origins (Museo Argentino de Ciencias Naturales of the frog kick? Alternate­leg swimming in ‘‘Bernardino Rivadavia’’, Buenos Aires, Ar­ primitive frogs, families Leiopelmatidae and gentina), Rafe Brown, William E. Duellman, Ascaphidae . Journal of Herpetology 33: 657– and John Simmons (University of Kansas 663. Museum of Natural History and Biodiversity Adler, K.A. 1989. Herpetologists of the past. In K.A. Adler (editor), Contributions to the his­

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Figures

Fig. 28. Maximum­likelihood tree of hyperoliid, arthroleptid, and astylosternid frogs provided by Vences et al. (2003c). A, Maximum­likelihood analysis of 12S rRNA molecule (187 informative sites) analyzed under a GTR substitution model (cost functions reported) suggested by Modeltest (Posada and Crandall, 1998). Initial alignments under Clustal software, costs not disclosed, and subsequently adjusted manually. Highly variable regions and gaps were excluded as evidence. B, Maximum­likelihood trees based on 138 informative sites of 16S rRNA molecule under a GTR substitution model (cost functions reported) for hyperoliids, arthroleptids, and astylosternids. Initial alignments were made under Clustal, costs not disclosed, and subsequently adjusted manually. Highly variable regions and gaps sites were excluded as evidence.

Fig. 64. A, Original tree of Jiang et al. (2005; from fig. 42) of Paini and (on right) its equivalent undirected network; B, Tree rerooted and with augmented resolution as implied by our general results,

Fig. 48. Consensus of weighted parsimony trees of Rhacophoridae suggested by J.A. Wilkinson et al. (2002), with their subfamily taxonomy on right. (This is Mantellidae and Rhacophoridae of other authors.) The tree was based on 2kb (of 12S and 16S mt rRNA as well as tRNAVal). Alignment was manual, guided by models of secondary structure with ambiguously aligned segments discarded. In analysis, transversions were weighted twice transitions. Whether reatment of gaps were treated as evidence of relationship or as missing data was not stated. Chirixalus eiffingeri was placed in Kurixalus by Ye, Fei, and Dubois (In Fei, 1999), and Chirixalus idiootocus was transferred into an explicitly polyphyletic/paraphyletic Aquixalus by Delorme et al. (2005). The tree was rooted on Nidirana adenopleura and Aquarana catesbeiana.

Fig. 49. Delorme et al.’s (2005) dendrogram of rhacophorids, based on undisclosed molecular and morphological data (although characters were summarized for some genera and suprageneric groups), redrawn to illuminate the paraphyly of groupings.

Fig. 71. Generic changes suggested for ranid taxa that we studied. This is not exhaustive and the Systematic Comments under Ranidae in ‘‘A Taxonomy of Living Amphibians’’ should be consulted for additional taxonomic changes.

Fig. 46. Maximum­likelihood tree of Matsui et al. (2005) for East Asian ranids, based on mitochondrial 12S and 16S rRNA sequences (total of 1,283 bp). Sequence alignment was done under ClustalX (Thompson et al., 1997) with cost functions not disclosed and subsequently adjusted manually, guided by secondary structure models as suggested by Kjer (1995). Modeltest 3.06 (Posada and Crandall, 1998) was used to select nucleotide evolutionary model (GTR) assumed for analysis. Fejervarya and Buergeria were used to root the tree.

Fig. 44. Maximum­likelihood tree of Holarctic Rana of Hillis and Wilcox (2005). The underlying data are ca. 2kb of mtDNA of the 12S–16S region (spanning the tRNAVal gene). Sequence alignment was done initially using Clustal W (Thompson et al., 1994), costs not disclosed, and manually adjusted, guided by assumed secondary structure, ambiguously aligned sequences discarded. It was not stated whether gaps were treated as data, but we presume not. Substitution model GTR 1 G1 PINVAR was assumed for the maximum­likelihood analysis. On the basis of previous research, the root was assumed