Bioacoustics of Poyntonophrynus

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. 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