Multilocus phylogeny and a new classification for Southeast … · 2015. 4. 21. · Multilocus...

Multilocus phylogeny and a new classification forSoutheast Asian and Melanesian forest frogs(family Ceratobatrachidae)

RAFE M. BROWN1*, CAMERON D. SILER2, STEPHEN J. RICHARDS3,ARVIN C. DIESMOS4 and DAVID C. CANNATELLA5

1Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas,Lawrence, KS 66045-7561, USA2Sam Noble Oklahoma Museum of Natural History and Department of Biology, University ofOklahoma, Norman, OK 73072-7029, USA3Herpetology Department, South Australian Museum, North Terrace, Adelaide, SA 5000, Australia4National Museum of the Philippines, Rizal Park, Padre Burgos Avenue, Ermita 1000, Manila,Philippines5Department of Integrative Biology and Texas Biodiversity Collections, University of Texas at Austin,1 University Station, C0990 Austin, TX 78712, USA

Received 28 May 2014; revised 2 November 2014; accepted for publication 25 November 2014

We present a near comprehensive, densely sampled, multilocus phylogenetic estimate of species relationships withinthe anuran family Ceratobatrachidae, a morphologically and ecologically diverse group of frogs from the island archi-pelagos of Southeast Asia and the South-West Pacific. Ceratobatrachid frogs consist of three clades: a small clade ofenigmatic, primarily high-elevation, semi-aquatic Sundaland species currently assigned to Ingerana (for which weerect a new genus), which is the sister taxon of two large, monophyletic radiations, each situated on islands on eitherside of Wallace’s Line. One radiation is composed of Philippine species of Platymantis and the other contains all taxafrom the eastern Indonesian, New Guinean, Solomon, Bismarck, and Fijian archipelagos. Several additional genera(Batrachylodes, Discodeles, Ceratobatrachus, and Palmatorappia) are nested within Platymantis, and of theseBatrachylodes and Discodeles are nonmonophyletic. To address the widespread paraphyly of the genus Platymantisand several additional nomenclatural issues, we undertook a wholesale nomenclatural reorganization of the family.Given our partially unresolved phylogeny, and in order to impart a conservative, stable taxonomy, involving a minimalnumber of genus-species couplet changes, we propose a conservative classification representing a few compromises.These changes are designed to preserve maximally the presumed original intent of taxonomy (widely used group namesassociated with morphological and ecological diversity of particular species or groups of species) while implementinga hierarchical system that is consistent with the estimate of phylogeny based on new molecular data.

© 2015 The Linnean Society of London, Zoological Journal of the Linnean Society, 2015, 174, 130–168.doi: 10.1111/zoj.12232

ADDITIONAL KEYWORDS: admiralty webbed frogs – evolutionary radiation – new genus – New Guineanwrinkled frogs – Philippine forest frogs – phylogenetic taxonomy – Solomon horned frogs – Solomon palm frogs– Solomon sticky-toed frogs – Sundaland mountain frogs – subgenera – taxonomy.

The Cornuferinae have arisen from Rana in different parts ofits range. They represent a very uniform group. Some of thegenera apparently grade into others, making the limits of thesegroups almost impossible to define. (Noble, 1931: 521).

Platymantis probably evolved within the Philippines in the lateTertiary and subsequently dispersed southwards into New Britain,the Solomon Islands, and Fiji by rafting. The direction of thesecondary radiation is a reflection of the demonstrable phylogeneticaffinities of the extant species. (Tyler, 1979: 78–79).

The Philippine fauna includes lineages with clear Papuanaffinities, Platymantis and Oreophryne. The presence of these*Corresponding author. E-mail: [email protected]

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Zoological Journal of the Linnean Society, 2015, 174, 130–168. With 3 figures

© 2015 The Linnean Society of London, Zoological Journal of the Linnean Society, 2015, 174, 130–168130

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two genera in the Philippines (but not in Palawan) may datefrom either pre-Tertiary or Oligocene. . .when the easternPhilippines-Halmahera arc was closest to New Guinea and theMelanesian Islands. (Inger, 1999: 462).

It is very likely that Platymantis arose from Rana and hasno relationship to Micrixalus. . .Palmatorappia of the Solomonsseems to be a case of parallel evolution in a different stock,namely Cornufer or an allied genus. (Noble, 1931: 522–523).

Rather than think of Platymantis as territory that you needto ‘divide up’ why not just see how much you can achieve to-gether in collaboration? [W. C. Brown (deceased), 1998, per-sonal communication with R. M. B.].

INTRODUCTION

The frog family Ceratobatrachidae (currentlyPlatymantis, Batrachylodes, Discodeles, Ceratobatrachus,Palmatorappia, and portions of the genus Ingerana)is a remarkable assemblage of amphibians distribut-ed throughout the Philippines, Palau, eastern Indo-nesia, New Guinea, the Solomon−Bismarck−Admiraltyarchipelagos, and the islands of Fiji (Brown, 1952;Zweifel, 1960, 1969; Brown & Tyler, 1968; Edgar &Lilley, 1993; Allison, 1996; Brown, 1997; Günther, 1999;Alcala & Brown, 1999; Inger, 1999; Tyler, 1999).Ceratobatrachids are noted for conspicuous character-istics of morphology (Boulenger, 1886, 1887; Brown,1952; Norris, 2002), larval direct development (Alcala,1962; Brown & Alcala, 1982), including unique struc-tures and patterns of embryonic growth (Thibaudeau& Altig, 1999; Narayan et al., 2011), and the abilityto colonize habitats that otherwise conspicuously lackranoid frogs (small, arid islands, dry limestone habi-

tats, and high-elevation mossy rain forests with nostanding water; Menzies, 2006; Pikacha, Morrison &Richards, 2008). This ability to persist and reproducein environments lacking standing fresh water has beenhypothesized to represent a key innovation that hasfacilitated dispersal and colonization across the South-West Pacific, and in the literature this life-history traitis associated with the presence of Platymantis on distantoceanic islands such as Palau (Crombie & Pregill, 1999)and Fiji (Gorham, 1965, 1968; Tyler, 1979; Ryan, 1984;Gibbons, 1985; Kuramoto, 1985, 1997; Ota & Matsui,1995; Narayan, Christi & Morley, 2008; Zug, 2013).

Whatever the combination of developmental, lifehistory, ecological characteristics or history, and cir-cumstances of colonization that led to the diversifica-tion of ceratobatrachid frogs in Southeast Asia and theSouth-West Pacific, the systematic relationships andpatterns of insular distributions of this group are ofinterest to biogeographers (Noble, 1931; Tyler, 1979;Inger, 1999). No other group of amphibians comes closeto exhibiting a similar distribution pattern with near-equivalent species diversity on either side of Wal-lace’s Line (Brown, 1952, 1997; Tyler, 1979, 1999; Inger,1999; Fig. 1). Furthermore, this radiation is unique inhaving such an appreciable portion of its diversity ondistant islands of the South-West Pacific (Allison, 1996;Brown, 1997; Inger, 1999).

Recent interest in species diversity of Philippineceratobatrachids has resulted in a sharp increase in de-scriptions of new species (Brown, Brown & Alcala, 1997a;Brown et al., 1997b, 1999a; Brown, Alcala & Diesmos,1997c, 1999b; Alcala & Brown, 1998, 1999; Brown, 2007;Brown & Gonzalez, 2007; Siler et al., 2007, 2009, 2010)

Figure 1. Distribution of the frog family Ceratobatrachidae. Numbers of species per major region are included inparentheses.

PHYLOGENY OF CERATOBATRACHIDAE 131


and 35–40 new species await description (Brown, 2004,2009; Brown, Diesmos & Alcala, 2008; Brown et al., 2013a).Known Melanesian ceratobatrachid diversity has in-creased as well, with new species described from main-land New Guinea (Günther, 1999, 2006), New Ireland(Brown & Menzies, 1979; Allison & Kraus, 2001), Manus(Richards, Mack & Austin, 2007; Kraus & Allison, 2009;Richards, Oliver & Brown, 2014), New Britain (Foufopoulos& Brown, 2004; Brown, Foufopoulos & Richards, 2006;Brown et al., 2006; Kraus & Allison, 2007, 2009; Brown,Richards & Broadhead, 2013), and the Solomon Islands(Brown & Richards, 2008).

To date, Platymantis (sensu lato) lacks an explicitphylogenetic definition (sensu de Queiroz & Gauthier,1990). Based on limited taxon sampling, Platymantisis clearly paraphyletic with respect to the morpho-logically derived non-Platymantis genera (Bossuyt et al.,2006; Wiens et al., 2009). Because of their consider-able species diversity (approximately 90 species;AmphibiaWeb, 2014), their curious distribution (Noble,1931; Brown, 1952, 1997; Tyler, 1979, 1999), their strik-ing array of morphological variation (Boulenger, 1884,1918a; Günther, 1859; Brown, 1952; Gorham, 1965;Brown et al., 1997a; Norris, 2002), and complex taxo-nomic history (Boulenger, 1918b; Brown, 1952; Dubois,1981, 1987, 1992; Inger, 1996; Frost, 2014), we under-took a phylogenetic analysis of the family, which hasonly been represented in previous systematic studiesby few species and sequences.

This study includes most Platymantis species diver-sity from both sides of Wallace’s Line (i.e. the Philip-pines vs. Solomon−Bismarck−Admiralty archipelagos;Fig. 1), representatives of the other four ceratobatrachidgenera (Ceratobatrachus, Palmatorappia, Batrachylodes,and Discodeles; AmphibiaWeb, 2014; Frost, 2014), afew species of Southeast Asian Ingerana (= Micrixalusof earlier authors; Inger, 1954, 1966; Inger & Tan, 1996a,b; now known to be allied to Ceratobatrachidae: Bossuytet al., 2006), and representative ranid outgroups fromAsia and Papuan faunal regions (Wiens et al., 2009;Blackburn & Wake, 2011). Here we provide aphylogenetic estimate of relationships amongst the frogsof the family Ceratobatrachidae (species of the generaPlatymantis, Palmatorappia, Ceratobatrachus,Discodeles, Batrachylodes, and some members of thegenus Ingerana) with particular attention to themonophyly and validity of the genera Platymantis andCornufer. We also address long-standing nomenclatu-ral problems with respect to generic taxonomy, andprovide a new comprehensive classification scheme tofacilitate future studies.

TAXONOMIC HISTORY OF CERATOBATRACHIDAE

The genus Platymantis has one of the most confus-ing histories and lengthy synonymy of any group of

ranoid frogs (Dubois, 1981, 1987, 1992; Ford &Cannatella, 1993; Frost, 2014). The unusual distribu-tion of the Ceratobatrachidae (Fig. 1), coupled with un-certainty about their systematic affinities (Noble, 1931)and a particularly unstable nomenclatural history, hasled to the current state in which relationships in thefamily are poorly understood (Norris, 2002; Brown, 2004;Frost et al., 2006; Köhler et al., 2008; Pyron & Wiens,2011). Biologists have indiscriminately referred a centuryof new species discoveries to the paraphyletic taxonPlatymantis and, to date, no comprehensive efforts tounderstand the group’s diversity or utilize phylogenyto inform classification have been undertaken. Theseactions of convenience have compromised attempts tounderstand the evolutionary relationships of the group(W. C. Brown, pers. comm.) and have prevented theempirical test of hypotheses regarding the biogeogra-phy and phylogenetic affinities of this evolutionary ra-diation (Allison, 1996; W. Brown, 1997; Inger, 1999;R. Brown, 2004; Bossuyt et al., 2006). Below, we sum-marize the taxonomic history of the family to eluci-date the nomenclatural issues that need to be addressedin order to implement a new classification (Dubois, 1981,1987, 1992; Inger, 1996; ICZN, 1999).

The genus Cornufer was named by Tschudi (1838)based on a single specimen from an uncertain local-ity (Zweifel, 1966). In subsequent years approximate-ly 20 species from the Philippines, New Guinea, NewBritain, New Ireland, the Solomons, and the Fijis weredescribed and assigned to Cornufer, Halophila, andHylodes on the basis of osteological and external mor-phological characters (Peters, 1863; Boulenger, 1886,1918a; Taylor, 1920, 1922a, b, 1923, 1925; Schmidt,1932; Parker, 1939, 1940; Brown, 1949, 1952; Gorham,1965). Meanwhile, several similar species were as-signed to the genus Platymantis (Günther, 1859), dif-fering from species of the genus Cornufer primarily onthe basis of narrowly or non-expanded terminal toe discs.Advocates of the validity of both Cornufer (wide discs)and Platymantis (narrow discs) included Boulenger(1918b), Barbour (1923), Van Kampen (1923; who rec-ognized Cornufer and Rana, with the subgenusPlatymantis), Noble (1931), Mertens (1934), Brown &Myers (1949), Brown (1952), and Gorham (1965). Inger(1954) considered the range of morphological vari-ation in the two genera to be a natural continuum ofvariation between the two extreme states of wide vs.narrowly expanded finger and toe discs. He proposedsynonymizing Platymantis with Cornufer, and thus ren-dering species with both wide and narrow terminalfinger and toe discs members of a single genus,Platymantis, a change followed by Alcala (1962) andmost others (but see Gorham, 1965).

Later, when it was determined that the type speciesof Cornufer was in fact a Neotropical frog in the genusEleutherodactylus, Zweifel (1966) proposed to the ICZN

132 R. M. BROWN ET AL.


that the name Cornufer be suppressed (Anonymous,1978); Zweifel (1967) summarized his reasoning andused Platymantis in subsequent publications (Zweifel,1969, 1975). However, the ICZN committee failed torule on Zweifel’s proposal for nearly ten years(Anonymous, 1978), and when it did, ruled againstZweifel’s proposition, which left Cornufer an avail-able name, unknown to the systematics community(Anonymous, 1978; ICZN, 1999). For the following 35years, systematists have referred all Southeast Asianand Melanesian forest frogs to Platymantis, of whichCornufer was considered a subjective synonym ofPlatymantis (Dubois, 1981; Frost, 1985, 2014).

The lengthy literature debate surrounding this taxo-nomic confusion discouraged investigators (notablyZweifel, 1967; Gorham, 1965; both assumed Cornuferwas unavailable) from coining a new generic name forspecies with wide discs to distinguish them from thespecies with narrow discs. This appears to have beenan admirable attempt to avoid further taxonomic in-stability but, as noted by Dubois (1981: 248): ‘. . .thisis a case where purely nomenclatural reasons haveimposed upon systematists a unanimity which purelytaxonomic arguments had not allowed them to reach’(translation from original French by M. Berson, Cali-fornia Academy of Sciences).

MATERIAL AND METHODSTAXON SAMPLING

We conducted fieldwork in the Philippines, easternIndonesia, the Admiralty Islands and Bismarck Archi-pelago of Papua New Guinea, and the Solomon Islands.This sampling was augmented by contributions of tissuesfrom these same areas, plus Palau, Borneo, and Fiji(see Specimens examined and Acknowledgements). Frogswere captured by hand, over-anaesthetized inchlorobutanol (KU IACUC no. 158-01), and dissectedfor liver and muscle; tissues were preserved by im-mersion in liquid nitrogen, 95% ethanol, high-saltdimethyl sulphoxide tissue preservation buffer, orRNAlater (Life Technologies). Specimens were fixedin buffered 10% formalin and stored in 70% ethanol.Voucher specimens are deposited in collections at theNational Museum of the Philippines (PNM), The Cin-cinnati Museum of Natural History (CMNH), Louisi-ana State University Museum of Natural Science(LSUMZ), the Texas Natural History Collections ofthe University of Texas at Austin (TNHC), the UnitedStates National Museum of Natural History (USNM),The Field Museum of Natural History (FMNH), theSouth Australian Museum (SAMA) the Western Aus-tralian Museum (WAM), the Bishop Museum (BPBM),and the University of Kansas Biodiversity Institute(KU).

DATA COLLECTION

We extracted total genomic DNA from liver or musclesamples with a Qiagen DNeasy kit or Fujita’s Guani-dine Thyocyanate protocol (Esselstyn et al., 2008). Ingroupsampling included 120 individuals representing the di-versity of the family Ceratobatrachidae, includingmembers of all six currently recognized genera(Batrachylodes, Ceratobatrachus, Discodeles, Ingerana,Palmatorappia, and Platymantis). Fifteen species wereincluded as outgroup taxa, representing a broad spec-trum of anuran diversity amongst the familiesDicroglossidae (Ingerana, Limnonectes, andHoplobatrachus), Microhylidae (Kaloula), and Ranidae(Amolops, Huia, Hylarana, and Rana) (Fig. 2; Appen-dix 1). Data for Ingerana tenasserimensis were down-loaded from GenBank (accession nos: DQ347030,AY322308). Each extraction was amplified for the genesof interest (Table 1) through standard PCR protocols(Palumbi, 1996).

We targeted a ∼2500-bp region of the 12S + tRNAVal

and 16S rRNA mitochondrial gene fragments usingvarious primers adopted or modified from publishedstudies (Goebel, Donnelly & Atz, 1999; Evans et al.,2003; Darst & Cannatella, 2004; Hillis & Wilcox, 2005;Table 1) in eight pairs to amplify segments via PCR;however, not all amplifications were successful. Addi-tionally, we sequenced portions of three nuclear loci:recombinase activating gene 1 (RAG1; ∼750 bp), tyrosinase(Tyr; ∼535-bp portion of exon 1), and proopiomelanocortin(POMC; ∼580 bp), using the primers and protocols ofWiens et al. (2005) and Bossuyt et al. (2006) (Appendix2). The nuclear genes were sampled for a subset of taxafor which mtDNA sequence was obtained.

We purified PCR product with QIAquick Gel Ex-tractions or used ExoSAPit (USB Corp.) with a 20%dilution of stock ExoSAPit, incubated for 30 min at 37 °Cand then 80 °C for 15 min. Cycle sequencing was carriedout with the following cycling conditions for 25 cycles:10 s at 96 °C; 5 s at 50 °C; and 4 min at 60 °C.

Cleaned PCR products were dye-labelled using Big-Dye terminator 3.1 (Applied Biosystems), purified usingSephadex (NC9406038, Amersham Biosciences,Piscataway, NJ), and sequenced on an ABI 3100 or3730xl automated capillary sequencer (AppliedBiosystems Inc.). Raw sequence data were processedusing SEQUENCING ANALYSIS software (AppliedBiosystems). Individual sequence chromatograms wereexamined in SEQUENCHER v. 4.3 (GeneCodes) andindividual single-stranded fragments were assem-bled into contiguous consensus reads, after checkingfor sequencing error, for subsequent analysis.

ALIGNMENT AND PHYLOGENETIC ANALYSIS

Initial alignments were produced in MUSCLE (Edgar,2004) and minor manual adjustments were made in



MacClade v. 4.08 (Maddison & Maddison, 2000). Formitochondrial gene regions, we defined ambiguouslyaligned regions as character sets using MacClade andexcluded regions of uncertain positional homologies fromfurther analyses after determining that doing so yieldedno difference in tree topology and unappreciable changesin nonparametric bootstrap values for parsimonysearches (not shown).

Preliminary analyses exploring the impact of missingdata on inferred outgroup relationships resulted insimilar relationships inferred with and without theinclusion of individual nuclear data partitions. To assesseffects of missing data, preliminary analyses of indi-

vidual genes and combinations of gene partitions wereconducted. We found that relationships recoveredamongst clades N, O, P, and Q (Fig. 2) varied betweenthe nuclear genes and mtDNA only. MrBayes analy-sis of the nuclear genes yielded only the topology(O,(N,P),(Q,R)). The nodes supporting these relation-ships had posterior probabilities (PP) = 1, except forthe clade O + N + P, which was 0.98. By contrast, analy-sis of the mtDNA resulted in (O,((N,P),(Q,R))); for allnodes the PP = 1.0. Thus, the nuclear and mtDNAtrees are strongly incongruent (Supporting Informa-tion Fig. S1). Because of this incongruence the com-bined tree (Fig. 2) lacks support for the relationships

Figure 2. Molecular phylogenetic estimate of major ceratobatrachid relationships based on maximum likelihood analy-sis of two mitochondrial gene partitions (12S–16S) and three nuclear genes (proopiomelanocortin, recombinase activat-ing gene 1, and tyrosinase; 11-partition model: Table 2). Maximum likelihood bootstrap and Bayesian posterior probabilityvalues are included. Boxed letters denoting selected nodes of interest are discussed in the text. Node B is Ceratobatrachidae.As illustrated, the tree is unrooted, and to save space the outgroups (Node A) are shown as if they form a clade, whichthey do not. The root of the tree lies on the branch between Kaloula and all other taxa. Photographs of selected speciesare included (approximately to scale), with current taxonomy summarized at tree tips (compare with revised taxonomy,summarized in Fig. 3). Nodal support: black dots ≥ 0.95 and ≥ 70 maximum likelihood bootstrap support (MLBS); greydots ≥ 0.75, posterior probabilities (PP) ≤ 0.95, and ≥ 50 MLBS ≤ 70. Support values provided (as MLBS/PP) for weaklysupported nodes and nodes with disparate levels of support between analyses.

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Table 1. Oligonucleotide primer sequences used in this study

Locus Primer name Sequence: 5′–3′ Citation

tRNAVal–16S MVZ59 ATAGCACTGAAAAYGCTDAGATG Goebel et al. (1999)tRNAVal GGTGTAAGCGAGAGGCTT Darst & Cannatella (2004)12L1 AAAAAGCTTCAAACTGGGATTAGATACCCCACTAT Hillis & Wilcox (2005)16Sa ATGTTTTTGGTAAACAGGCG Hillis & Wilcox (2005)12Sm GGCAAGTCGTAACATGGTAAG Hillis & Wilcox (2005)16Sh GCTAGACCATKATGCAAAAGGTA Hillis & Wilcox (2005)16Sc GTRGGCCTAAAAGCAGCCAC Darst & Cannatella (2004);

Hillis & Wilcox (2005)16Sd CTCCGGTCTGAACTCAGATCACGTAG Darst & Cannatella (2004)

POMC POMC-1 GAATGTATYAAAGMMTGCAAGATGGWCCT Wiens et al. (2005, 2009)POMC-2 TAYTGRCCCTTYTTGTGGGCRTT Wiens et al. 2005, 2009)POMC-3 TCTGCMGARTCWCCYGTGTTTCC Wiens et al. (2005, 2009POMC-4 TGGCATTYTTGAAAAGAGTCAT Wiens et al. (2005, 2009)

RAG1 Amp-RAG1 F AGCTGCAGYCARTACCAYAARATGTA Mauro et al. (2004)RAG1-R GCAAAGTTTCCGTTCATTCTCAT Fu, Weadick & Bi (2007)

Tyr Tyr1A AGGTCCTCTTRAGCAAGGAATG Bossuyt & Milinkovitch (2000);Bossuyt et al. (2006)

Tyr1B AGGTCCTCYTRAGGAAGGAATG Bossuyt & Milinkovitch (2000);Bossuyt et al. (2006)

Tyr1C GGCAGAGGAWCRTGCCAAGATGT Bossuyt & Milinkovitch (2000);Bossuyt et al. (2006)

Tyr1D TCCTCCGTGGGCACCCARTTCCC Bossuyt & Milinkovitch (2000);Bossuyt et al. (2006)

POMC, proopiomelanocortin; RAG1, recombinase activating gene 1.



amongst N, O, P, and Q + R; essentially these fourclades form a polytomy. However, this does not affectour taxonomy because we did not name any nodeswith PP < 0.98. The individual clades M, N, O, P, Q,and R are also each supported by PP > 0.98.

Therefore, we chose to include all data in a concat-enated data set. However, we urge careful considera-tion of incongruence between these partitions beforethe phylogeny is used for biogeographical inference orcomparative analyses. Our final concatenated matrix(deposited in Dryad at: doi:10.5061/dryad.4fd0k) con-sisted of 4416 nucleotide positions with variable numbersof taxa sequenced for 12S (N = 52), 16S (128), RAG1(102), Tyr (98), and POMC (76).

Partitioned Bayesian analyses were conducted inMrBayes v. 3.2.1 (Huelsenbeck & Ronquist, 2001;Ronquist & Huelsenbeck, 2003). All nuclear gene datasets were partitioned by codon position for protein-coding regions, and the mitochondrial genes 12S-tRNAVal and 16S were each treated as individualpartitions, for a total of 11 sequence partitions (Table 2).The Akaike information criterion (AIC), as implement-ed in jModeltest v. 2.1.4 (Guindon & Gascuel, 2003;Darriba et al., 2012), was used to select the best modelof nucleotide substitution for each partition (Table 2).We set the ratepr (rate multiplier) parameter to ‘vari-able’ to allow substitution rates to vary amongst subsets,and set a dirichlet process prior (1,1,1,1) on the statefrequency parameter. Default priors were used for allother model parameters. We ran four independentMarkov chain Monte Carlo analyses, each with four

Metropolis-coupled chains, an incremental heating tem-perature of 0.02, and an exponential distribution witha rate parameter of 25 as the prior on branch lengths.All analyses were run for 15 000 000 generations, withparameters and topologies sampled every 3000 gen-erations. We assessed stationarity with TRACER v. 1.4(Rambaut & Drummond, 2007) and confirmed conver-gence with AWTY (Wilgenbusch, Warren & Swofford,2004; Nylander et al., 2007). We conservatively dis-carded the first 20% of samples as burn-in, resultingin a total of 4000 topologies from the posterior distri-bution for each of four runs.

Partitioned maximum likelihood (ML) analyses wereconducted in RAxMLHPC v. 7.0 (Stamatakis, 2006) onthe concatenated data set using the same partition-ing strategy and sets of deleted characters as the Bayes-ian analysis. The General Time Reversible model withvariable sites modeled according to the Gamma dis-tribution was selected via AIC and used for all subsets(Table 2), with ML analyses performed using the rapidhill-climbing algorithm (Stamatakis et al., 2007). Eachinference was initiated with a random starting treeand nodal support was assessed with 1000 bootstrappseudoreplicates employing the rapid hill-climbingalgorithm (Stamatakis, Hoover & Rougemont, 2008).All new sequences were deposited in GenBank(Appendix 1).

RESULTSTAXON SAMPLING AND PHYLOGENETIC ANALYSES

The aligned matrix contains 135 samples (Appendix1). Similar to other high-level phylogenetic studies (Wienset al., 2009; Pyron & Wiens, 2011) Ceratobatrachidaewas found to be monophyletic, except for some speciesof Ingerrana (see below). To economize on space wepresent the tree (Fig. 2) as if it were rooted betweenthe outgroup and ingroup. The numbers of variablecharacters are: 996 of 1632 (12S); 627 of 909 (16S); 246of 588 (POMC); 233 of 534 (Tyr); 187 of 753 (RAG1).

With a few exceptions, all analyses result in topolo-gies with moderate to high ML bootstrap support (MLBS)and PP amongst species and major clades within thefamily Ceratobatrachidae (Fig. 2). General topologicalpatterns amongst the major clades of outgroup speciesare congruent with published studies (Bossuyt et al.,2006; Frost et al., 2006; Wiens et al., 2009; Pyron &Wiens, 2011) and are not discussed further. Inferredrelationships from Bayesian and ML analyses werebroadly similar; however, a few differences were ob-served. The sample of Ingerana tenasserimensis, thetype species of Ingerana, was recovered by all analysesas part of a clade of outgroup samples (Clade A) withstrong support (MLBS = 100; PP = 0.98; Supporting In-formation Fig. S1). No analyses support the monophyly

Table 2. Models of evolution selected by Akaike informa-tion criterion (AIC; as implemented in jModeltest) and thoseapplied in partitioned, model-based, analyses of mitochondrial(12S, 16S, tRNAVal) and nuclear [proopiomelanocortin (POMC),recombinase activating gene 1 (RAG1), tyrosinase (Tyr)] data

Partition AIC modelNumber ofcharacters

12S + tRNAVal GTR + Γ* 162416S GTR + Γ 909POMC, first codon position GTR + Γ 196POMC, second codon position GTR + Γ 196POMC, third codon position HKY + Γ 196RAG1, first codon position HKY + Γ 251RAG1, second codon position GTR + Γ 251RAG1, third codon position GTR + Γ 251Tyr, first codon position GTR + Γ 178Tyr, second codon position GTR + Γ 178Tyr, third codon position JC 178

*GTR + Γ, General Time Reversible Model with variablesites modeled according to the Gamma distribution; JC,Jukes-Cantor.



of Ingerana as currently defined (Fig. 2), and Ingeranamariae, Ingerana baluensis, and Ingerana rajae wererecovered as a well-supported clade that is the sistergroup of all remaining ceratobatrachid taxa (Clade C).Amongst ingroup samples (Clade B), all analyses re-covered three primary clades with high support (CladesC, E, M). Apart from Ingerana (Clade C), the remain-ing ceratobatrachids were recovered as part of two clades(Clades E, M). The first is composed of members ofDiscodeles, Ceratobatrachus, Palmatorappia,Batrachylodes, and Melanesian species of Platymantis(Clade E). A large, well-supported clade of PhilippinePlatymantis (Clade M) is the sister group of Clade E.

Focusing solely on strongly supported clades withinthe Philippine and non-Philippine ceratobatrachids (ex-cluding Ingerana), several relationships are note-worthy: (1) species of Discodeles are not recovered asa clade; Discodeles malukuna and Discodelesbufoniformis are supported as the sister group of allother non-Philippine taxa in Clade E, whereas Discodelesguppyi is nested within a group of Pacific species ofPlatymantis (Clades F and H); (2) Bayesian analysessupport the monophyly of the genus Batrachylodes,albeit with weak support for the inclusion ofBatrachylodes minutus as the sister taxon of all othersampled members of the genus (PP = 0.69; Support-ing Information Fig. S1), whereas ML analyses recovera clade of Batrachylodes to the exclusion of B. minutus(Clade L), with the placement of B. minutus weaklysupported (MLBS = 48); (3) the majority of Pacific speciesof the genus Platymantis are inferred to be membersof a single clade (Clade J); and (4) within the well-supported Philippine radiation (Clade M), all analy-ses support five major clades (Clades N–R).

A NEW CLASSIFICATION OF THE FAMILYCERATOBATRACHIDAE

OVERVIEW

Our phylogenetic analyses and those of others (Bossuytet al., 2006; Frost et al., 2006; Wiens et al., 2009; Pyron& Wiens, 2011) unequivocally support two large clades(Clades E and M) that are together the sister groupof the new genus described below (including Sundalandand Palawan Island species formerly assigned toIngerana). Given that these two clades arephylogenetically and biogeographically well circum-scribed (one is endemic to the Philippines, west of Wal-lace’s Line; and the other is widely distributed eastof Wallace’s Line throughout eastern Indonesia, NewGuinea, Palau, the Bismarcks, Admiralty archipela-gos, Solomon Islands, and Fiji), we assign to them theavailable generic names Platymantis (with restrictedcontent, see below), and Cornufer (with expandedcontent, see below), together within a new unranked

clade, which we define and name below. Our recogni-tion of three genera, Alcalus, Cornufer, and Platymantis,rather than an extensive splitting of Ceratobatrachidaeinto numerous genera, also maintains a desirable degreeof stability of content of Platymantis.

NEW TAXA AND ALLOCATION OF SPECIES TO EXISTING

SUPRASPECIFIC NAMES

We present parallel ranked and phylogenetic taxono-mies. Whereas traditional ranked taxonomy isagnostic with respect to phylogenetic relationships andfocuses on the content or the concept of the taxon,phylogenetic taxonomies associate a name with a cladeand are based on phylogenetic trees (de Queiroz &Gauthier, 1990, 1992, 1994). The phylogenetic defini-tions of taxon names follow the general recommenda-tions of the draft PhyloCode versions 4c and 5a1(Cantino & de Queiroz, 2014); we provide traditionaldiagnoses for most of the same names following therequirements of the ICZN so that these will be avail-able in the sense of the ICZN (1999).

Some explanation of terms is needed; these are takenfrom Cantino & de Queiroz (2014, particularly Article9.3 and the Glossary). A specifier is a species, speci-men, or apomorphy that serves as a reference pointto specify a clade of interest; here we use type speciesas specifiers. A crown clade is a node-based clade thatoriginates with the last common ancestor of two or moreextant species (or organisms); crown clades are de-limited by extant and not extinct taxa, although a crownclade may include extinct taxa. A node-based clade origi-nates with a particular node on a tree, rather than abranch (stem). By contrast, a branch-based (stem-based) clade originates with a specific branch. A branch-based clade might include fossils as the most basalbranches.

Maximum crown-clade definitions are formed as ‘thelargest crown clade containing A but not Z’ or the crownclade originating in the most recent common ances-tor of A and all extant organisms or species that sharea more recent common ancestor with A than with Z(or X, or Y, as needed), where A is an extant internalspecifier and Z is an external specifier (Article 9.9,Cantino & de Queiroz, 2014). In other words, it is themost inclusive crown clade including A but not Z (andother specifiers as needed). Maximum crown-clade defi-nitions are particularly useful when basal relation-ships are not well resolved and when it is desirableto include newly discovered species under the exist-ing taxon name, rather than proposing a new cladename or redefining the clade name to include the newspecies that lie outside of the clade. By contrast, if onewishes to stabilize the content of a taxon (sayCeratobatrachidae) such that the concept ofCeratobatrachidae is not expanded to include a newly



discovered sister group, then a node-based definitionof Ceratobatrachidae is preferable.

Converted clade names (CCNs) are also defined usingphylogenetic conventions. New clade names (NCNs) arenewly coined names. All unranked phylogenetic namesare italicized. We have generally used type species asspecifiers in phylogenetic definitions. The taxonomicauthority (author and date) for ranked taxon namesis included in Table 3. The authors and date for allNCNs are considered to be Brown, Siler, Richards,Diesmos, and Cannatella 2014.

In some cases we have coined NCNs to refer to thesame group denoted by existing ranked names, ratherthan convert (in the sense of the PhyloCode) the rankedname to a clade name. We have done this to avoid con-verting names that imply a rank because under thePhyloCode suffixes such as -idae or -ini do not indi-cate rank. Note that the Phylocode neither encour-ages nor discourages the use of ranks.

For example, Ceratobatrachidae has been various-ly ranked as a subfamily (Bossuyt et al., 2006), a family(Boulenger, 1884), or a tribe (Dubois, 1992). IfCeratobatrachidae were converted to a clade name, andif in the future Ceratobatrachidae were treated as thesubfamily Ceratobatrachinae, then the clade nameCeratobatrachidae and the ranked subfamily nameCeratobatrachinae would refer to the same clade,causing confusion.

For phylogenetic definitions of Alcalus, Cornufer, andPlatymantis we use maximum crown-clade defini-tions because the relationships of these taxa to eachother are well supported. Similarly we have usedmaximum crown-clade definitions for those subcladesof Platymantis that are strongly supported. However,relationships amongst the subclades within Cornuferare weakly supported in places, and several speciesare not assigned to a named subclade of Cornufer. Forgenera that typically have been named based onapomorphies we use apomorphy-based names to re-strict the content of these clades to species that possessthese apomorphies. An example is Discodeles, whichis unique amongst ceratobatrachids in having exten-sively webbed feet.

CERATOBATRACHIDAE BOULENGER, 1884

Type genusCeratobatrachus Boulenger, 1884.

DiagnosisFrogs of the family Ceratobatrachidae differ from theirclose relatives by the possession of (1) direct develop-ment; and (2) T-shaped terminal phalanges with as-sociated expanded finger and toe discs.

Phylogenetic definitionCeratobatrachia (NCN) is a node-based name that refersto the clade arising from the most recent common an-

cestor of Alcalus mariae (type species of Alcalus),Cornufer vitiensis (type species of Cornufer), andPlatymantis pliciferus (type species of Platymantis; cur-rently a junior synonym of Platymantis corrugatus).

ContentThe genera Alcalus (three or four species), Cornufer(58 species), and Platymantis (31 described species).

CommentWe define Ceratobatrachia using a node-baseddefinition, rather than a maximum crown-cladedefinition, because the closest relative of Ceratobatrachia(= Ceratobatrachidae) from amongst the ranoidsis not clear (e.g. Bossuyt et al., 2006; Pyron & Wiens,2011). The node-based name ensures that future useof the Ceratobatrachia refers to the same node,regardless of whether that node name isCeratobatrachidae or Ceratobatrachinae; i.e. its useis independent of any particular ranked taxonomy.We have not converted the ranked nameCeratobatrachidae to a phylogenetic name, but ratherwe have named Ceratobatrachia to avoid confusionbetween the homonymous ranked name and convert-ed clade name.

We apply the ranked name Ceratobatrachidae (Fig. 2,Clade B) to the node usually identified asCeratobatrachidae or Ceratobatrachinae. Several family-group names are available for clades within theCeratobatrachidae, including Cornuferinae Noble 1931,Ceratobatrachinae Boulenger, 1884, and PlatymantinaeLaurent, 1986. Ceratobatrachidae Boulenger, 1884, isnot nomenclaturally problematic. Cornuferinae wasnamed by Noble (1931) to include the generaBatrachylodes, Ceratobatrachus, Cornufer, Discodeles,Hylarana, Micrixalus, Palmatorappia, Platymantis, andStaurois (including Simomantis). Savage (1973: 354)later coined Platymantinae as a subfamily of Ranidae.However, he did not explicitly provide a list of char-acters that diagnose the taxon as required by the In-ternational Code of Zoological Nomenclature (ICZN1999; hereafter, the Code). Thus, the namePlatymantinae Savage, 1973, is not available (Article13.1; ICZN, 1999) and is a nomen nudum. Laurent(1986) diagnosed the same taxon and made the nameavailable as Platymantinae Laurent, 1986. Dubois (1992)listed Cornuferinae Noble, 1931 and PlatymantiniLaurent, 1986, as junior synonyms of CeratobatrachidaeBoulenger, 1884.

By contrast, Frost (2014) listed Cornuferinae Noble,1931, as a synonym of Eleutherodactylidae Lutz, 1954,stating ‘synonymy by implication of synonymy ofCornufer with Eleutherodactylus by Zweifel (1966).’ Thenomenclatural history of Cornufer is discussed in detailunder the Cornufer account, but relevant to the issueis that the International Commission on Zoological



Tab

le3.

Cla

ssifi

cati

onof

the

fam

ily

Cer

atob

atra

chid

aeba

sed

onph

ylog

enet

ices

tim

ate

from

two

mit

och

ondr

ial

gen

efr

agm

ents

(12S

+tR

NA

Val,

16S

)an

dth

ree

nu

clea

rge

nes

(pro

opio

mel

anoc

orti

n,

reco

mbi

nas

eac

tiva

tin

gge

ne

1,an

dty

rosi

nas

e).

Taxa

mar

ked

wit

han

aste

risk

(*)

wer

en

otin

clu

ded

inth

eph

ylog

enet

ican

alys

is;

som

eof

thes

ew

ere

assi

gned

tocl

ades

onth

eba

sis

ofph

enot

ypic

sim

ilar

ity

and

pres

um

edcl

ose

phyl

ogen

etic

affin

ity;

taxa

mar

ked

wit

ha

dagg

er(†

)ar

eex

tin

ct.

Nod

eO

rigi

nal

desi

gnat

ion

(au

thor

,da

te)

Pre

viou

sge

ner

icpl

acem

ent

Cu

rren

tge

ner

icpl

acem

ent

Su

bgen

us

Cla

den

ame

Spe

cies

Not

es

CIn

gera

na

Alc

alu

sge

n.

nov

.A

lcal

us

Inge

ran

am

aria

eIn

ger,

1954

Inge

ran

aA

lcal

us

non

en

one

mar

iae

Cor

nu

fer

balu

ensi

sB

oule

nge

r,18

96In

gera

na

Alc

alu

sn

one

non

eba

luen

sis

Ran

asa

riba

Sh

elfo

rd,

1905

Inge

ran

aA

lcal

us

non

en

one

sari

ba*

Inge

ran

ara

jae

Iska

nda

r,B

ickf

ord

&A

rifi

nIn

gera

na

Alc

alu

sn

one

non

era

jae

EC

orn

ufe

rC

orn

ufe

rF

Ran

abu

fon

ifor

mis

Bou

len

ger,

1884

Dis

cod

eles

Cor

nu

fer

Pot

amor

ana

subg

en.

nov

.P

otam

oran

an

ewcl

ade

nam

ebu

fon

ifor

mis

com

b.n

ov.

New

com

bin

atio

n

Dis

cod

eles

mal

uku

na

Bro

wn

&W

ebst

er,

1969

Dis

cod

eles

Cor

nu

fer

Pot

amor

ana

Pot

amor

ana

mal

uku

na

com

b.n

ov.

New

com

bin

atio

nR

ana

opis

thod

onB

oule

nge

r,18

84D

isco

del

esC

orn

ufe

rP

otam

oran

aP

otam

oran

aop

isth

odon

com

b.n

ov.*

New

com

bin

atio

nR

ana

vogt

iH

edig

er,

1934

(rep

lace

men

tn

ame

for

Ran

ave

ntr

icos

us

Vog

t,19

12)

Dis

cod

eles

Cor

nu

fer

Pot

amor

ana

Pot

amor

ana

vogt

ico

mb.

nov

.*N

ewco

mbi

nat

ion

TH

alop

hil

avi

tien

sis

Gir

ard,

1853

Pla

tym

anti

sC

orn

ufe

rC

orn

ufe

rYa

nu

boto

new

clad

en

ame

viti

ensi

sTr

ansf

erre

dto

Cor

nu

fer,

type

spec

ies

for

gen

us

Cor

nu

fer

Hyl

odes

viti

anu

sD

um

éril

,18

53P

laty

man

tis

Cor

nu

fer

Cor

nu

fer

Cor

nu

fer

viti

anu

sTr

ansf

erre

dto

Cor

nu

fer

Pla

tym

anti

sm

egab

oton

ivit

i†W

orth

y,20

01P

laty

man

tis

Cor

nu

fer

Cor

nu

fer

Cor

nu

fer

meg

abot

oniv

iti†

*Tr

ansf

erre

dto

Cor

nu

fer

SC

erat

obat

rach

us

guen

ther

iB

oule

nge

r,18

84C

erat

obat

rach

us

Cor

nu

fer

Cer

atob

atra

chu

sC

erat

obat

rach

us

guen

ther

ico

mb.

nov

.N

ewco

mbi

nat

ion

HC

orn

ufe

rgu

ppyi

Bou

len

ger,

1884

Dis

cod

eles

Cor

nu

fer

Dis

cod

eles

Dis

cod

eles

gupp

yico

mb.

nov

.N

ewco

mbi

nat

ion

UH

ylel

laso

lom

onis

Ste

rnfe

ld,

1920

(syn

.H

ypsi

ran

ah

effe

rnan

iK

ingh

orn

,19

28)

Pal

mat

orap

pia

Cor

nu

fer

Pal

mat

orap

pia

Pal

mat

orap

pia

hef

fern

ani

com

b.n

ov.

New

com

bin

atio

n,

resu

rrec

tion

ofC

o.h

effe

rnan

ias

asu

bsti

tute

nam

efo

rso

lom

onis

(see

text

)L

Bat

rach

ylod

esel

egan

sB

row

n&

Par

ker,

1970

Bat

rach

ylod

esC

orn

ufe

rB

atra

chyl

odes

Bat

rach

ylod

esel

egan

sco

mb.

nov

.*N

ewco

mbi

nat

ion

Bat

rach

ylod

esgi

gas

Bro

wn

&P

arke

r,19

70B

atra

chyl

odes

Cor

nu

fer

Bat

rach

ylod

esB

atra

chyl

odes

giga

sco

mb.

nov

.*N

ewco

mbi

nat

ion

Bat

rach

ylod

esm

edio

dis

cus

Bro

wn

&P

arke

r,19

70B

atra

chyl

odes

Cor

nu

fer

Bat

rach

ylod

esB

atra

chyl

odes

med

iod

iscu

sco

mb.

nov

.N

ewco

mbi

nat

ion

Bat

rach

ylod

esm

onta

nu

sB

row

n&

Par

ker,

1970

Bat

rach

ylod

esC

orn

ufe

rB

atra

chyl

odes

Bat

rach

ylod

esm

onta

nu

sco

mb.

nov

.*N

ewco

mbi

nat

ion

Bat

rach

ylod

esm

onta

nu

sB

row

n&

Mye

rs,

1949

Bat

rach

ylod

esC

orn

ufe

rB

atra

chyl

odes

Bat

rach

ylod

estr

ossu

lus

com

b.n

ov.

New

com

bin

atio

nB

atra

chyl

odes

vert

ebra

lis

Bou

len

ger,

1887

Bat

rach

ylod

esC

orn

ufe

rB

atra

chyl

odes

Bat

rach

ylod

esve

rteb

rali

sco

mb.

nov

.N

ewco

mbi

nat

ion

Sph

enop

hry

ne

wol

fiS

tern

feld

,19

20B

atra

chyl

odes

Cor

nu

fer

Bat

rach

ylod

esB

atra

chyl

odes

wol

fico

mb.

nov

.N

ewco

mbi

nat

ion

JP

laty

man

tis

adia

stol

us

Bro

wn

,R

ich

ards

,S

uku

mar

an&

Fou

fopo

ulo

s,20

06P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

rasu

bgen

.n

ov.

Aen

igm

anu

ran

ewcl

ade

nam

ead

iast

olu

sco

mb.

nov

.N

ewco

mbi

nat

ion

Pla

tym

anti

sad

mir

alti

ensi

sR

ich

ards

,M

ack

&A

ust

in,

2007

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

raad

mir

alti

ensi

sco

mb.

nov

.*N

ewco

mbi

nat

ion

Pla

tym

anti

sak

arit

hym

us

Bro

wn

&T

yler

,19

68P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

akar

ith

ymu

sco

mb.

nov

.N

ewco

mbi

nat

ion

Pla

tym

anti

sbo

ule

nge

riP

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

bou

len

geri

Tran

sfer

red

toC

orn

ufe

rP

laty

man

tis

citr

inos

pilu

sB

row

n,

Ric

har

ds&

Bro

adh

ead,

2013

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

raci

trin

ospi

lus

com

b.n

ov.

New

com

bin

atio

n

Pla

tym

anti

scu

stos

Ric

har

ds,

Oli

ver

&B

row

n,

2014

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

racu

stos

com

b.n

ov.

New

com

bin

atio

nP

laty

man

tis

des

tica

ns

Bro

wn

&R

ich

ards

,20

08P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

des

tica

ns

com

b.n

ov..

New

com

bin

atio

nP

laty

man

tis

gill

iard

iZ

wei

fel,

1960

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

ragi

llia

rdi

Tran

sfer

red

toC

orn

ufe

rP

laty

man

tis

gupp

yiB

oule

nge

r,18

84P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

hed

iger

iTr

ansf

erre

dto

Cor

nu

fer,

nom

enn

ovu

mfo

rC

o.gu

ppyi

(see

text

)P

laty

man

tis

latr

oR

ich

ards

,M

ack

&A

ust

in,

2007

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

rala

tro

com

b.n

ov.

New

com

bin

atio

nC

orn

ufe

rm

acro

psB

row

n,

1965

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

ram

acro

psTr

ansf

erre

dto

Cor

nu

fer



Tab

le3.

Con

tin

ued

Nod

eO

rigi

nal

desi

gnat

ion

(au

thor

,da

te)

Pre

viou

sge

ner

icpl

acem

ent

Cu

rren

tge

ner

icpl

acem

ent

Su

bgen

us

Cla

den

ame

Spe

cies

Not

es

Pla

tym

anti

sm

acro

scel

esZ

wei

fel,

1975

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

ram

acro

scel

esco

mb.

nov

.N

ewco

mbi

nat

ion

Pla

tym

anti

sm

agn

us

Bro

wn

&M

enzi

es,

1979

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

ram

agn

us

com

b.n

ov.

New

com

bin

atio

nP

laty

man

tis

mam

usi

oru

mB

row

n&

Fou

fopo

ulo

s,20

04P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

mam

usi

oru

mco

mb.

nov

.N

ewco

mbi

nat

ion

Pla

tym

anti

sn

akan

aior

um

Bro

wn

,F

oufo

pou

los

&R

ich

ards

,20

06P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

nak

anai

oru

mco

mb.

nov

.N

ewco

mbi

nat

ion

Pla

tym

anti

sn

ecke

riB

row

n&

Mye

rs,

1949

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

ran

ecke

riTr

ansf

erre

dto

Cor

nu

fer

Pla

tym

anti

sn

exip

us

Zw

eife

l,19

75P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

nex

ipu

sTr

ansf

erre

dto

Cor

nu

fer

Pla

tym

anti

sco

rru

gatu

sva

r.pa

puen

sis

Mye

r,18

75(‘1

874’

)P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

papu

ensi

sTr

ansf

erre

dto

Cor

nu

fer

Pla

tym

anti

spa

rili

sB

row

n&

Ric

har

ds,

2008

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

rapa

rili

sco

mb.

nov

.N

ewco

mbi

nat

ion

Pla

tym

anti

spe

lew

ensi

sP

eter

s,18

67P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

pele

wen

sis

Tran

sfer

red

toC

orn

ufe

rP

laty

man

tis

papu

ensi

ssc

hm

idti

Bro

wn

&T

yler

,19

68P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

sch

mid

tico

mb.

nov

.N

ewco

mbi

nat

ion

Cor

nu

fer

solo

mon

isB

oule

nge

r,18

84P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

solo

mon

isTr

ansf

erre

dto

Cor

nu

fer

Pla

tym

anti

ssu

lcat

us

Kra

us

&A

llis

on,

2007

Pla

tym

anti

sC

orn

ufe

rA

enig

man

ura

Aen

igm

anu

rasu

lcat

us

com

b.n

ov.

New

com

bin

atio

nP

laty

man

tis

web

eri

Sch

mid

t,19

32P

laty

man

tis

Cor

nu

fer

Aen

igm

anu

raA

enig

man

ura

web

eri

Tran

sfer

red

toC

orn

ufe

rP

laty

man

tis

acro

chor

du

sB

row

n,

1965

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rac

roch

ord

us

Tran

sfer

red

toC

orn

ufe

r

Pla

tym

anti

sac

ule

odac

tylu

sB

row

n,

1952

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rac

ule

odac

tylu

s*Tr

ansf

erre

dto

Cor

nu

fer

Pla

tym

anti

sba

tan

tae

Bro

wn

&Z

wei

fel,

1969

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rba

tan

tae

com

b.n

ov.

New

com

bin

atio

n

Pla

tym

anti

sbi

mac

ula

tus

Gü

nth

er,

1999

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rbi

mac

ula

tus

com

b.n

ov.

New

com

bin

atio

n

Pla

tym

anti

sbr

own

iA

llis

on&

Kra

us,

2001

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rbr

own

ico

mb.

nov

.*N

ewco

mbi

nat

ion

Pla

tym

anti

sbu

fon

ulu

sK

rau

s&

All

ison

,20

07P

laty

man

tis

Cor

nu

fer

Not

assi

gned

tosu

bgen

us

Cor

nu

fer

bufo

nu

lus

com

b.n

ov..

New

com

bin

atio

n

Pla

tym

anti

sca

esio

psK

rau

s&

All

ison

,20

09P

laty

man

tis

Cor

nu

fer

Not

assi

gned

tosu

bgen

us

Cor

nu

fer

caes

iops

com

b.n

ov.

New

com

bin

atio

n

Pla

tym

anti

sch

eesm

anae

Par

ker,

1940

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rch

eesm

anae

Tran

sfer

red

toC

orn

ufe

r

Pla

tym

anti

scr

ypto

tis

Gü

nth

er,

1999

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rcr

ypto

tis

com

b.n

ov.*

New

com

bin

atio

n

Pla

tym

anti

sm

anu

sK

rau

s&

All

ison

,20

09P

laty

man

tis

Cor

nu

fer

Not

assi

gned

tosu

bgen

us

Cor

nu

fer

man

us

com

b.n

ov.

New

com

bin

atio

n

Pla

tym

anti

sm

imic

us

Bro

wn

&T

yler

,19

68P

laty

man

tis

Cor

nu

fer

Not

assi

gned

tosu

bgen

us

Cor

nu

fer

mim

icu

sco

mb.

nov

.N

ewco

mbi

nat

ion

Bat

rach

ylod

esm

inu

tus

Bro

wn

&P

arke

r,19

70P

laty

man

tis

Cor

nu

fer

Not

assi

gned

tosu

bgen

us

Cor

nu

fer

min

utu

sco

mb.

nov

.R

emov

edfr

omB

atra

chyl

odes

,n

ewco

mbi

nat

ion

Pla

tym

anti

sm

yers

iB

row

n,

1949

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rm

yers

iTr

ansf

erre

dto

Cor

nu

fer

Cor

nu

fer

park

eri

Bro

wn

,19

65P

laty

man

tis

Cor

nu

fer

Not

assi

gned

tosu

bgen

us

Cor

nu

fer

park

eri

Tran

sfer

red

toC

orn

ufe

r

Pla

tym

anti

spu

nct

atu

sP

eter

s&

Dor

ia,

1878

Pla

tym

anti

sC

orn

ufe

rN

otas

sign

edto

subg

enu

sC

orn

ufe

rpu

nct

atu

sTr

ansf

erre

dto

Cor

nu

fer

Pla

tym

anti

sw

uen

sch

eoru

mG

ün

ther

,20

06P

laty

man

tis

Cor

nu

fer

Not

assi

gned

tosu

bgen

us

Cor

nu

fer

wu

ensc

heo

rum

com

b.n

ov.

New

com

bin

atio

n

MP

laty

man

tis

Pla

tym

anti

sP

laty

man

tis



Tab

le3.

Con

tin

ued

Nod

eO

rigi

nal

desi

gnat

ion

(au

thor

,da

te)

Pre

viou

sge

ner

icpl

acem

ent

Cu

rren

tge

ner

icpl

acem

ent

Su

bgen

us

Cla

den

ame

Spe

cies

Not

es

NP

laty

man

tis

corr

uga

tus

Du

mér

il,

1853

Pla

tym

anti

sP

laty

man

tis

Pla

tym

anti

sT

agom

ukh

us

new

clad

en

ame

corr

uga

tus

(syn

.P

l.pl

icif

era

Gu

enth

er,

1854

)

P.pl

icif

era

(syn

P.co

rru

gatu

s)is

the

type

spec

ies

for

gen

us

Pla

tym

anti

sO

Pla

tym

anti

sba

nah

aoB

row

n,A

lcal

a,D

iesm

os&

Alc

ala,

1975

bP

laty

man

tis

Pla

tym

anti

sL

ahat

nan

guri

subg

en.

nov

.L

ahat

nan

guri

new

clad

en

ame

ban

ahao

Pla

tym

anti

sbi

akS

iler

,D

iem

sos,

Lin

kem

,D

iesm

os&

Bro

wn

,20

10P

laty

man

tis

Pla

tym

anti

sL

ahat

nan

guri

subg

en.

nov

.L

ahat

nan

guri

new

clad

en

ame

biak

*

Cor

nu

fer

corn

utu

sTa

ylor

,19

22P

laty

man

tis

Pla

tym

anti

sL

ahat

nan

guri

Lah

atn

angu

rico

rnu

tus

Pla

tym

anti

sin

sula

tes

Bro

wn

&A

lcal

a,19

70P

laty

man

tis

Pla

tym

anti

sL

ahat

nan

guri

Lah

atn

angu

riin

sula

tus

Pla

tym

anti

sle

viga

tus

Bro

wn

&A

lcal

a,19

74P

laty

man

tis

Pla

tym

anti

sL

ahat

nan

guri

Lah

atn

angu

rile

viga

tus

Pla

tym

anti

spy

gmae

us

Bro

wn

,Alc

ala

&D

iesm

os,

1988

Pla

tym

anti

sP

laty

man

tis

Lah

atn

angu

riL

ahat

nan

guri

pygm

aeu

sP

Ph

ilau

tus

haz

elae

Tayl

or,

1920

Pla

tym

anti

sP

laty

man

tis

Tir

ahan

ula

psu

bgen

.n

ov.

Tir

ahan

ula

pn

ewcl

ade

nam

eh

azel

ae

Pla

tym

anti

sis

arog

Bro

wn

,B

row

n,A

lcal

a&

Fro

st(r

epla

cem

ent

nam

efo

rP

laty

man

tis

reti

cula

tus

Bro

wn

,B

row

n&

Alc

ala,

1997

a)

Pla

tym

anti

sP

laty

man

tis

Tir

ahan

ula

pT

irah

anu

lap

isar

og

Pla

tym

anti

sla

wto

ni

Bro

wn

&A

lcal

a,19

74P

laty

man

tis

Pla

tym

anti

sT

irah

anu

lap

Tir

ahan

ula

pla

wto

ni

Cor

nu

fer

mon

tan

us

Tayl

or,

1922

Pla

tym

anti

sP

laty

man

tis

Tir

ahan

ula

pT

irah

anu

lap

mon

tan

us

Pla

tym

anti

spa

nay

ensi

sB

row

n,

Bro

wn

&A

lcal

a,19

97P

laty

man

tis

Pla

tym

anti

sT

irah

anu

lap

Tir

ahan

ula

ppa

nay

ensi

sP

hil

autu

spo

lill

ensi

sTa

ylor

,19

22P

laty

man

tis

Pla

tym

anti

sT

irah

anu

lap

Tir

ahan

ula

ppo

lill

ensi

sP

laty

man

tis

sier

ram

adre

nsi

sB

row

n,A

lcal

a,O

ng,

&D

iesm

os,

1999

Pla

tym

anti

sP

laty

man

tis

Tir

ahan

ula

pT

irah

anu

lap

sier

ram

adre

nsi

s

Cor

nu

fer

subt

erre

stri

sTa

ylor

,19

22P

laty

man

tis

Pla

tym

anti

sT

irah

anu

lap

Tir

ahan

ula

psu

bter

rest

ris

QP

laty

man

tis

baya

ni

Sil

er,A

lcal

a,D

iesm

os&

Bro

wn

,20

08P

laty

man

tis

Pla

tym

anti

sT

ahan

anpu

no

subg

en.

nov

.T

ahan

anpu

no

new

clad

en

ame

baya

ni

Pla

tym

anti

sd

iesm

osi

Bro

wn

&G

onza

lez,

2007

Pla

tym

anti

sP

laty

man

tis

Tah

anan

pun

oT

ahan

anpu

no

die

smos

iC

orn

ufe

rgu

enth

eri

Bou

len

ger,

1882

Pla

tym

anti

sP

laty

man

tis

Tah

anan

pun

oT

ahan

anpu

no

guen

ther

iP

laty

man

tis

luzo

nen

sis

Bro

wn

,Alc

ala,

Die

smos

&A

lcal

a,19

97P

laty

man

tis

Pla

tym

anti

sT

ahan

anpu

no

Tah

anan

pun

olu

zon

ensi

s

Pla

tym

anti

sn

egro

sen

sis

Bro

wn

,Alc

ala,

Die

smos

&A

lcal

a,19

97P

laty

man

tis

Pla

tym

anti

sT

ahan

anpu

no

Tah

anan

pun

on

egro

sen

sis

Pla

tym

anti

sra

bori

Bro

wn

,Alc

ala,

Die

smos

&A

lcal

a,19

97P

laty

man

tis

Pla

tym

anti

sT

ahan

anpu

no

Tah

anan

pun

ora

bori

RP

laty

man

tis

caga

yan

ensi

sB

row

n,A

lcal

a&

Die

mso

s,19

99P

laty

man

tis

Pla

tym

anti

sL

upa

colu

ssu

bgen

.n

ov.

Lu

paco

lus

new

clad

en

ame

caga

yan

ensi

s

Cor

nu

fer

dor

sali

sD

um

éril

,18

53P

laty

man

tis

Pla

tym

anti

sL

upa

colu

sL

upa

colu

sd

orsa

lis

Pla

tym

anti

sin

dep

ren

sus

caga

yan

ensi

sB

row

n,A

lcal

a&

Die

mso

s,19

99P

laty

man

tis

Pla

tym

anti

sL

upa

colu

sL

upa

colu

sin

dep

ren

sus

Pla

tym

anti

sm

imu

lus

Bro

wn

,Alc

ala

&D

iem

sos,

1997

Pla

tym

anti

sP

laty

man

tis

Lu

paco

lus

Lu

paco

lus

mim

ulu

sP

laty

man

tis

nao

mia

eA

lcal

a,B

row

n&

Die

smos

,19

88P

laty

man

tis

Pla

tym

anti

sL

upa

colu

sL

upa

colu

sn

aom

iae

Pla

tym

anti

spa

engi

Sil

er,

Lin

kem

,D

iesm

os&

Alc

ala,

2007

Pla

tym

anti

sP

laty

man

tis

Lu

paco

lus

Lu

paco

lus

paen

gi

Pla

tym

anti

sps

eud

odor

sali

sB

row

n,A

lcal

a&

Die

mso

s,19

99P

laty

man

tis

Pla

tym

anti

sL

upa

colu

sL

upa

colu

sps

eud

odor

sali

s

Pla

tym

anti

ssp

elae

us

Bro

wn

&A

lcal

a,19

82P

laty

man

tis

Pla

tym

anti

sL

upa

colu

sL

upa

colu

ssp

elae

us

Pla

tym

anti

sta

ylor

iB

row

n,A

lcal

a&

Die

mso

s,19

99P

laty

man

tis

Pla

tym

anti

sL

upa

colu

sL

upa

colu

sta

ylor

i



Nomenclature designated Halophila vitiensis Girard,1853, as the type species of Cornufer. As H. vitiensishas been placed consistently in either Platymantis orCornufer within the Ceratobatrachidae, Cornufer andCornuferinae would not be considered part ofEleutherodactylidae.

ALCALINAE SUBFAM. NOV.Type genusAlcalus (see account below).

DiagnosisThe diagnosis for Alcalinae is the same as for the genusAlcalus, below.

Phylogenetic definitionWe have not defined Alcalinae as a phylogenetic namebecause it would be redundant with Alcalus; it addsno new information about phylogenetic relationships.However, we name the ranked subfamily Alcalinae, eventhough it is also redundant in content with Alcalus,to provide a coordinate name for its sister-taxonCeratobatrachinae.

ContentOne genus, Alcalus, which includes the speciesAl. mariae, Al. baluensis, and Al. rajae. We antici-pate that Ingerana sariba eventually will be trans-ferred to the new genus as well.

EtymologySee Etymology section for the genus Alcalus below.

ALCALUS GEN. NOV.Type speciesMicrixalus mariae Inger 1954.

DiagnosisMembers of the genus Alcalus can be distinguishedfrom many members of the clade Anurajen (species ofthe genera Cornufer and Platymantis) by having (1)an intermediate body size (Al. baluensis: males 20–25 mm Snout-Vent Length (SVL), females 26–31;Al. mariae: males 32–37 mm SVL, females 35–43); (2)a broad head (vs. slender to moderately broad); (3) acoarsely textured, shagreened, or ‘wrinkled’ skin ap-pearance in all species (vs. smooth, tuberculate, or withlongitudinal dorsolateral dermal ridges); (4) widelyexpanded, terminally squared, spatulate toe discs (vs.non- or minimally expanded, terminally rounded);(5) semi-aquatic microhabitat preferences (vs. prefer-ences for terrestrial or arboreal microhabitats in mostspecies); and by the (6) presence of nuptial pads in males(vs. absence); (7) absence of vocal sacs (vs. presence

of median subgular vocal sacs); (8) absence of super-numerary tubercles on hand (vs. presence in mostspecies); (9) presence of elongate subarticular tuber-cles (vs. presence, round); (10) absence of outer meta-tarsal tubercles on plantar surface of feet (vs. presencein most species); and (11) presence of extensive, usuallyfull, interdigital webbing of the feet (vs. absence).

Phylogenetic definitionAlcalus (NCN) is a maximum crown-clade name thatrefers to the crown clade (C) originating in the lastcommon ancestor of Al. mariae and all extant speciesthat share a more recent common ancestor withAl. mariae than with Cornufer vitiensis or Platymantiscorrugatus. It can also be conceived of as the largestcrown clade containing Al. mariae, but not Co. vitiensisor Pl. corrugatus.

ContentSoutheast Asian (Sunda Shelf and Palawan Island)species formerly placed in Ingerana (Table 3): Al. mariae,Al. baluensis, Al. rajae, and presumably Al. sariba(Shelford, 1905), which was not sampled (Table 3).

CommentIt is not surprising that the montane, semi-aquatic,Southeast Asian island archipelago species formerlyreferred to Ingerana comprise a monophyletic group,unrelated to the ecologically dissimilar andbiogeographically disjunct mainland species of Ingerana(as presently understood, from Andaman Islands,Bhutan, China, north eastern India, Myanmar, andNepal). Erection of a new genus to accommodate thesetaxa is undertaken here with reference to thephylogenetic placement of the type species of ‘true’Ingerana (I. tenasserimensis), which in our phylog-eny is more closely related (but with weak support)to the Dicroglossidae than to the Ceratobatrachidae(Fig. 2). The placement of Alcalus as the sister groupof the clade Anurajen (containing genera Platymantisand Cornufer) has been confirmed elsewhere (Bossuytet al., 2006; Frost et al., 2006; Wiens et al., 2009; Pyron& Wiens, 2011), although taxon sampling was not asextensive. The phylogenetic relationships and pos-sible additional generic subdivision of the non-ceratobatrachid (perhaps dicroglossid) species referredto Ingerana remain unstudied.

EtymologyA masculine noun honouring our long-term collabora-tor, friend, and mentor Angel C. Alcala for his numer-ous contributions to the systematics, ecology,conservation, and developmental biology of South-east Asian amphibians. Suggested common name:Alcala’s dwarf mountain frogs.



CERATOBATRACHINAE BOULENGER, 1884

DiagnosisOther species of Ceratobatrachinae differ from Alcalusby having (1) a relatively narrow head (vs. wide); and(2) smooth skin, with or without dermal tubercles and/or dermal ridges (vs. coarsely textured, shagreened,or ‘wrinkled’ in appearance); (3) rounded terminal toediscs (vs. spatulate toe discs with squarish terminalshape); (4) absence of nuptial pads in males (vs. pres-ence); (5) presence of a medial subgular vocal sacin most species (vs. absence); (6) presence of palmarsupernumerary tubercles (vs. absence); and (7)presence of rounded subarticular tubercles and outermetatarsal tubercles on plantar surface (vs. pres-ence, elongate). Finally, most species ofCeratobatrachinae (except Cornufer guppyi, species ofthe subgenus Potamorana, and Platymantis levigatus)differ from Alcalus by the absence of interdigital webbingon the feet (vs. presence); and by having terrestrialor arboreal microhabitat preferences (vs. semi-aquatic). Morphological synapomorphies forCeratobatrachinae have not been identified.

ContentThe genera Cornufer and Platymantis (see below).

CommentWe have not converted Ceratobatrachinae (Clade D)to a clade name, but have instead coined a newunranked clade name, Anurajen. If Ceratobatrachinaewere converted to a clade name, and if the familyCeratobatrachidae (Node B) were re-ranked asCeratobatrachinae in the future, then the phylogeneticname Ceratobatrachinae and the ranked nameCeratobatrachinae would refer to different clades, whichwould cause confusion. Therefore, we define a new cladename below denoting the same clade as the subfam-ily Ceratobatrachinae.

ANURAJEN NEW CLADE NAME

Phylogenetic definitionA maximum crown-clade name referring to the crownclade (D) originating with the most recent common an-cestor of Co. vitiensis and all extant species that sharea more recent common ancestor with Co. vitiensis thanwith Alcalus mariae.

ContentSpecies of Platymantis and Cornufer (as forCeratobatrachinae).

CommentThis clade is supported by high bootstrap propor-tions and posterior probabilities (Fig. 2), and consistsof two large subclades (Platymantis and Cornufer) situ-

ated on either side of Wallace’s Line (Fig. 1).Ceratobatrachinae is the ranked name equivalent incontent to Anurajen (Table 4).

EtymologyWe are pleased to name the new clade after JenniferAnne Weghorst in appreciation of the many times thatshe has arduously proofread our manuscripts and forthe devoted support and encouragement that she hasprovided to R. M. B. for many years. Anurajen is derivedfrom the Latin noun Anura and the abbreviated ap-pellation Jen.

GENUS PLATYMANTIS GÜNTHER, 1858

Type speciesPlatymantis pliciferus Günther, 1858, currently con-sidered a junior subjective synonym of Pl. corrugatus(Duméril, 1853); subsequent designation by Zweifel(1967).

DiagnosisMembers of the exclusively Philippine genus Platymantiscan be distinguished from the three or four knownspecies of Alcalus (with the exception of Al. mariae,all Alcalus occur outside the Philippines) by the (1)absence of interdigital webbing or the presence of highlyreduced webbing (vs. presence); (2) presence of median

Table 4. Comparison of ranked and phylogenetic taxono-mies for Ceratobatrachidae

Ranked taxonomy Phylogenetic taxonomy

Ceratobatrachidae Ceratobatrachia NCNAlcalinae subfam. nov. No name needed*

Alcalus new genus Alcalus NCNCeratobatrachinae Anurajen NCN

Cornufer Cornufer CCNCornufer (subgenus) Yanuboto NCN†Potamorana new subgenus Potamorana NCNCeratobatrachus (subgenus) Ceratobatrachus CCNDiscodeles (subgenus) Discodeles CCNPalmatorappia (subgenus) Palmatorappia CCNBatrachylodes (subgenus) Batrachylodes CCNAenigmanura new subgenus Aenigmanura NCN

Platymantis Platymantis CCNPlatymantis (subgenus) Tagomukhus NCN†Lahatnanguri new subgenus Lahatnanguri NCNTirahanulap new subgenus Tirahanulap NCNTahananpuno new subgenus Tahananpuno NCNLupacolus new subgenus Lupacolus NCN

CCN, converted clade name; NCN, new clade name.*A new phylogenetic name equivalent to the Linnean nameCeratobatrachinae is not necessary as this name would have thesame content as the clade (genus) Alcalus.†A NCN is provided to avoid having the same name apply to twodifferent clades, as is the case with the genus and subgenus rank.



subgular vocal sacs (vs. absence); (3) absence of nuptialpads (vs. presence); (4) presence of supernumerary tu-bercles on the hands (vs. absence); and (5) presenceof metatarsal tubercles on the foot (vs. absence).

Although all Philippine Platymantis are readily di-agnosed from members of the genus Alcalus, and se-lected species of the genus Cornufer, subgenera Cornufer(= Yanuboto), Potamorana, Discodeles, Ceratobatrachus,Palmatorappia, and Batrachylodes (see diagnoses ofthose clades), characters universally distinguishing Phil-ippine Platymantis from all members of the genusCornufer (in particular, the subgenus Aenigmanura andspecies formerly referred to ‘Platymantis’ from theSolomon−Bismarck−Admiralty Archipelago, Palau, NewGuinea, and eastern Indonesia, Table 3; see species notassigned to subgenus) have not been identified. We areunaware of any morphological synapomorphies forPlatymantis, although our phylogenetic analysis pro-vides very strong support (PP = 1.0) for this clade (Fig. 2,Clade M).

Phylogenetic definitionPlatymantis (CCN) is a maximum crown-clade namereferring to the crown clade (M) originating with themost recent common ancestor of Pl. corrugatus (synonymPl. plificerus, the type species of Platymantis) and allextant species that share a more recent common an-cestor with Pl. corrugatus than with Al. mariae orCo. vitiensis.

ContentPhilippine taxa (currently 31 species) of the subgen-era Platymantis (Tagomukhus, NCN), Lahatnanguri,Tirahanulap, Tahananpuno, and Lupacolus (Table 3).Numerous Philippine species await description, sug-gesting that the content of this genus will expandrapidly in the near future (Siler et al., 2007, 2009, 2010,2011, 2012; Brown et al., 2008, 2012, 2013a, 2013b;Brown & Stuart, 2012).

CommentThe content of the genus Platymantis Günther, 1858,is hereby restricted to the primary Philippine clade(M) and we apply Cornufer Tschudi 1838 to its sistergroup (Clade E), which includes the type species of thegenus Cornufer, Halophila vitiensis Girard, 1853. Giventhat the relationships amongst clades O, N, P, Q, andR show some degree of uncertainty, we have used onespecifier from each clade to assure that the phylogeneticdefinition of the name of Clade M will remain stable.

EtymologyFrom the Greek adjective ‘platy’, meaning flat and‘mantis’. The meaning of ‘mantis’ here is confusing; oftenit is stated that generic names ending in ‘mantis’ arederived from the Greek noun ‘mantis’, a term com-

monly meaning prophet or soothsayer (Liddell & Scott,1996). However, Günther (1858) specifically stated inhis etymology of Platymantis that the Greek noun‘mantis’ referred to ‘tree-frog’ rather than soothsayer.‘Mantis’ was applied by ancient Greeks to the speciesHyla arborea (a species perceived to be akin to proph-ets because it produces advertisement calls prior tothe arrival of rain; Liddell & Scott, 1996). Kraus &Allison (2007) resolved previous confusion concerningthe gender of Platymantis, stemming from Günther’s(1858) mistaken use of both masculine and feminineepithets for the two species included in the originaldefinition of the genus, and R. Günther’s (1999) as-sertion that Platymantis should be treated as a femi-nine noun. Günther (1999) stated that ‘According toGünther (1858) . . . mantis is Greek, of feminine gender,and means tree frog.’ (pp. 327–328), but did not explainhis opinion. We follow Kraus & Allison (2007) in con-sidering the gender of Platymantis as masculine.

SUBGENUS PLATYMANTIS GÜNTHER, 1858

DiagnosisThe subgenus Platymantis (currently a single recog-nized species, Pl. corrugatus) differs from other speciesof Platymantis by having (1) elongate longitudinaldermal ridges along the dorsal body surfaces (vs. dorsumsmooth or tuberculate); (2) distinctive ‘quaaack’ ad-vertisement calls (vs. frequency sweeps, pure tones,or complex calls); and (3) distinctive dark lateral headcoloration (of varying shades; vs. lateral head pigmentundifferentiated from surrounding coloration). The di-agnostic dark lateral head coloration forms a dark ‘face-mask’ that we consider a synapomorphy of this clade.Additionally, members of this subgenus can be diag-nosed from species of arboreal variable Philippine forestfrogs of the genera Lahatnanguri (Platymantis banahao,Pl. cornutus, and Pl. insulatus), and all members ofthe subgenera Tahananpuno and Tirahanulap, by theabsence of expanded digital tips of fingers and toes (vs.presence of some degree of terminal digital expan-sion of fingers and toes), and by having a terrestrialmicrohabitat preference (normally calling beneath leaflitter) and a crepuscular (vs. nocturnal) calling activ-ity pattern (Table 3).

ContentThe allopatric populations of (1) the Luzon and WestVisayan faunal regions, (2) the Camiguin Norte lineage,(3) the populations from the Mindanao faunal regionislands, and (4) the Mindoro Island populations, allcurrently referred to Pl. corrugatus (Table 3). The sub-genus Platymantis is equivalent in content to theunranked taxon Tagomukhus.

Conversion of the name Platymantis wouldresult in two different clades, ranked as genus and



subgenus, with the same name, Platymantis. There-fore we define a NCN, denoting the same clade as sub-genus Platymantis.

TAGOMUKHUS NEW CLADE NAME

Phylogenetic definitionTagomukhus (NCN) is a maximum crown-clade namereferring to the crown clade (Clade N) originating withthe most recent common ancestor of Pl. corrugatus andall extant species and populations that share a morerecent common ancestor with Pl. corrugatus than withPl. levigatus, Pl. hazelae, Pl. guentheri, or Pl. dorsalis.

ContentPlatymantis (Tagomukhus) corrugatus (syn. P. plicifera,type species of the genus Platymantis).

CommentAt a minimum, we anticipate that the Luzon (+ W.Visayan) Pleistocene Aggregate Island Complex (PAIC;Brown & Diesmos, 2009; Brown et al., 2013a), theMindanao PAIC, the Mindoro PAIC, and CamiguinNorte Island populations will all eventually be recog-nized as distinct species (K. Cobb, R.M.B., A.C.D., C.D.S.,& A.C. Alcala, unpubl. data). Platymantis pliciferus,the type species of the genus, is an available namethat applies to the Mindanao PAIC population (Günther,1859; Peters, 1873), should it be demonstrated to bea diagnosable evolutionary lineage worthy of taxo-nomic recognition.

EtymologyFrom the Tagalog adjective tago, meaning ‘concealed’or ‘unseen’ and the Tagalog noun mukha, meaning ‘coun-tenance’, in reference to the darkly pigmented face-mask present in varying degrees of distinctiveness inmost populations. The name is masculine in gender.Suggested common name: Philippine masked frogs.

LAHATNANGURI SUBGEN. NOV.Type speciesPlatymantis levigatus Brown & Alcala, 1974.

DiagnosisIndividual species of the subgenus Lahatnanguri differfrom other members of Platymantis by characters relatedto their general classification as arboreal tree frogs(Pl. banahao, Pl. cornutus, readily distinguished fromall Philippine Platymantis except members of the sub-genus Tahananpuno), distinctive mottled-coloured lime-stone specialist species (Pl. insulatus, readily diagnosedfrom all Philippine Platymantis except Platymantisbayani, Platymantis biak, and Platymantis speleaus),miniaturized species (Platymantis pygmaeus, SVL 13–

15 mm, readily distinguished from all PhilippinePlatymantis except possibly Platymantis naomiae), anda unique Romblon Province semi-aquatic speciesPl. levigatus (vs. all remaining Philippine species ter-restrial, scansorial, or arboreal). The wide range of mor-phological and ecological variation in this clade rendersan unambiguously exclusive diagnosis of Lahatnanguriimpossible. We are unaware of morphologicalsynapomorphies for this group, although ourphylogenetic analysis provides very strong support forthis clade (Clade O, Fig. 2).

Phylogenetic definitionLahatnanguri (NCN) is a maximum crown-clade namereferring to the crown clade (O) originating with themost recent common ancestor of Platymantis(Lahatnanguri) levigatus and all extant species thatshare a more recent common ancestor with Pl. levigatusthan with Pl. corrugatus, Pl. hazelae, Pl. guentheri, orPl. dorsalis.

ContentPlatymantis banahao, Pl. biak, Pl. cornutus,Pl. insulatus, Pl. levigatus, and Pl. pygmaeus (Table 3).

CommentSeveral unrecognized terrestrial species eventually willbe assigned to the subgenus Lahatnanguri, includingat least three from Mindanao Island (species 20, 21,and 40), a miniature ground frog from the RomblonProvince islands of Sibuyan and Tablas (R. M. Brown,A. C. Diesmos & C. D. Siler, unpubl. data), and at leastone arboreal species from Luzon Island (species 10)(Fig. 2). Although some species (Pl. banahao,Pl. insulatus) of the subgenus Lahatnanguri (Clade O)are phenotypically very similar to some species(Platymantis diesmosi, Pl. bayani, Pl. guentheri, Pl.rabori, Pl. negrosensis) of the subgenus Tahananpuno(Fig. 2, Clade Q) and were, in fact, grouped in anonphylogenetic taxonomy as the Pl. guentheri group(Brown et al., 1997a, b; Alcala & Brown, 1999), thisphenotypic similarity appears to be a case ofecomorphological convergence.

EtymologyFrom the Tagalog (Filipino) phrase lahat ng uri,meaning ‘all kinds’ or ‘every type’ in reference to thefull range of morphological and ecological variationwithin this clade, including miniature semifossorialspecies, large terrestrial ground frogs, semiaquaticspecies, limestone cave specialists, and high-elevationtree canopy frogs. The name is masculine in gender.Suggested common name: variable Philippine forestfrogs.



TIRAHANULAP SUBGEN. NOV.Type speciesPhilautus hazelae (Taylor, 1920).

DiagnosisThe morphologically, ecologically, and acoustically similarspecies of Tirahanulap differ from all other subgen-era of Platymantis by having: (1) widely expanded ter-minal discs of fingers and toes (vs. non- or minimallyexpanded): (2) subdigital surfaces relatively flat withlow subarticular tubercles (vs. subarticular tuberclesprominently rounded to pointed); (3) greatly reducedFinger I (vs. Finger I as long or nearly as long as FingerII); (4) tonal advertisement calls of constant frequen-cy (vs. possession of frequency sweep calls or calls withmultiple syllables of different frequencies); (5) smallclutch sizes (four to eight eggs vs. clutches typicallyof 20 or more eggs); and (6) a mid- to upper montaneshrub-layer vegetation microhabitat preference (vs. ter-restrial, semiaquatic, forest canopy, limestone, orsemifossorial). We consider the reduced length of FingerI, and the low, flat subarticular tubercles to be uniquesynapomorphies for the clade, which is strongly sup-ported in phylogenetic analyses (Fig. 2, Clade P).

Phylogenetic definitionTirahanulap (NCN) is a maximum crown-clade namereferring to the crown clade (Fig. 2, Clade P) origi-nating with the most recent common ancestor ofPlatymantis (Tirahanulap) hazelae and all extant speciesthat share a more recent common ancestor withPl. hazelae than with Pl. corrugatus, Pl. levigatus,Pl. guentheri, or Pl. dorsalis.

ContentPlatymantis hazelae, Pl. isarog, Pl. lawtoni, Pl.montanus, Pl. panayensis, Pl. polillensis, Pl.sierramadrensis, and Pl. subterrestris (Table 3).

CommentSpecies of Tirahanulap form a morphologically and eco-logically cohesive group that corresponds to thePl. hazelae group of Brown et al. (1997a; 1999a) andAlcala & Brown (1999). The members of this clade areecologically and phenotypically most similar to Cornufer(Palmatorappia) heffernani (formerly Palmatorappiasolomonis) and the high-elevation shrub frogs of NewBritain (Cornufer macrosceles, Cornufer citrinospilus,and Cornufer mamusiorum) and Manus Island (Cornufercustos). We are aware of at least four currently un-recognized species in this clade (species 2, 3, 5, and42; Fig. 2).

EtymologyFrom the Tagalog verb tumira, meaning, when con-jugated (‘tirahan’), to ‘inhabit’ or ‘reside within’, and

the Tagalog noun ulap, meaning cloud; together meaning‘cloud-dwellers’ or ‘they come from the clouds’. The nameis masculine in gender. Suggested common name: Phil-ippine cloud frogs.

TAHANANPUNO SUBGEN. NOV.Type speciesCornufer guentheri Boulenger, 1882.

DiagnosisMembers of this tree canopy specialist clade or Phil-ippine rain frogs, subgenus Tahananpuno, differ fromother all species of Platymantis (except Pl. banahao,Pl. cornutus, and Pl. insulatus, see below) by having(1) widely expanded terminal discs of fingers and toes(vs. non- or minimally expanded in terrestrial speciesof Tagomukus and Lupacolus); (2) prominent, roundedto pointed subarticular tubercles (vs. flattened on ventralsurfaces in cloud frog species of the subgenusTirahanulap); (3) pulsed advertisement calls (vs. tonal,constant frequency calls of cloud frogs, subgenusTirahanulap; vibrational, stridulated, or complexmultisyllable calls of species of terrestrial frogs of thesubgenera Tagomukus and Lupacolus); and (4) under-story (Pl. guentheri), limestone (Pl. bayani), cliff-edge(Pl. diesmosi), or canopy vegetation microhabitat pref-erences (all other Tahananpuno species). Althoughwidely expanded terminal discs of fingers and toesappear to be a synapomorphy for this clade, they haveevolved independently and diagnose a small subcladeof variable Philippine forest frogs only, subgenusLahatnanguri (Pl. banahao, Pl. cornutus, andPl. insulatus). We are unaware of any unique charac-ters that distinguish species of this new subgenus fromother species of Platymantis. Nevertheless, ourphylogenetic analyses provide strong support for thisclade (Fig. 2, Clade Q).

Phylogenetic definitionTahananpuno (NCN) is a maximum crown-clade namereferring to the crown clade (Fig. 2, Clade Q) origi-nating with the most recent common ancestor ofPlatymantis (Tahananpuno) guentheri and all extantspecies that share a more recent common ancestor withPl. guentheri than with Pl. corrugatus, Pl. dorsalis,Pl. hazelae, or Pl. levigatus.

ContentPlatymantis bayani, Pl. diesmosi, Pl. guentheri, Pl.luzonensis, Pl. negrosensis, and Pl. rabori (Table 3).

CommentThe subgenus Tahananpuno corresponds to thereadily distinguished Pl. guentheri group as defined byBrown et al. (1997a, b) and Alcala & Brown (1999).



Interestingly, and in contrast to expectations based onmorphology and understory/canopy microhabitat pref-erences (Brown et al., 1997a), Pl. banahao, Pl. cornutus,and Pl. insulatus are not part of this clade (or of theformer Pl. guentheri group; Brown et al., 1997a, b; Alcala& Brown, 1999), but rather fall in Clade O (Lahatnanguri).We are aware of at least four additional unrecognizedspecies in this clade (species 6, 7, 8, and 9; Fig. 2).

EtymologyTahananpuno is a masculine noun, derived from theTagalog verb tahanan meaning ‘to dwell upon’, or ‘tooccupy’ and noun puno, ‘tree’, in reference to the pre-vailing microhabitat preference of species in this clade:understory and canopy treefrogs. The name is mas-culine in gender. Suggested common name: Philip-pine rain frogs.

LUPACOLUS SUBGEN. NOV.Type speciesCornufer dorsalis Duméril, 1853.

DiagnosisPhilippine forest ground frogs of the subgenus Lupacolusare distinguished from other species of Platymantis (excepta few species of subgenus Lahatnanguri, see below) byhaving (1) non- to minimally expanded terminal discsof fingers and toes [vs. finger and toe discs expandedin cloud frogs of the subgenus Tirahanulap, rain frogsof the genus Tahananpuno, and three species of vari-able forest frogs of the subgenus Lahatnanguri(Pl. banahao, Pl. cornutus, and Pl. insulatus)]; (2) promi-nently rounded to pointed subarticular tubercles (vs.ventrally flattened in cloud frogs of the subgenusTirahanulap); (3) highly variable and often complexmultisyllable advertisement calls [vs. tonal, constantfrequency calls in cloud frogs of the subgenus Tirahanulap,or repeatedly pulsed calls in rain frogs of the subgenusTahananpuno and a few species of variable Philippineforest frogs, subgenus Lahatnanguri (Pl. banahao,Pl. cornutus, and Pl. insulatus)]; and (4) a predomi-nantly terrestrial, forest-floor microhabitat prefer-ence, with a tendency to call from slightly elevated perchesof 0.2–0.5 m [vs. perching in shrub and understory veg-etation in cloud frogs, subgenus Tirahanulap, and rainfrogs, subgenus Tahananpuno, and a few species of vari-able Philippine forest frogs, subgenus Lahatnanguri(Pl. banahao, Pl. cornutus, and Pl. insulatus)]. We areunaware of any morphological synapomorphies for thisgroup, but our phylogenetic analysis provides very strongsupport for this clade (Fig. 2, Clade R).

Phylogenetic definitionLupacolus (NCN) is a maximum crown-clade name re-ferring to the crown clade (Fig. 2, Clade R) originat-

ing with the most recent common ancestor of Pl. dorsalisand all extant species that share a more recent commonancestor with Pl. dorsalis than with Pl. corrugatus,Pl. hazelae, Pl. guentheri, or Pl. levigatus.

ContentPlatymantis cagayanensis, Pl. dorsalis, Pl. indeprensus,Pl. mimulus, Pl. naomiae, Pl. paengi, Pl. pseudodorsalis,Pl. spelaeus, and Pl. taylori (Table 3).

CommentFor the most part, Lupacolus corresponds to thePl. dorsalis group of W. Brown et al. (1997a, 1999a, b)and Alcala & Brown (1999). This clade of generalizedterrestrial ground frogs contains a large percentage ofcurrently unrecognized species (at least 15 small- tomedium-sized ground frogs from the northern andcentral islands of the archipelago, including species 12,14, 15, 18, 19, 22–25, 27, 29, 30, 31, 33, and 43; Fig. 2),but contrary to predictions from taxonomy (Brown et al.,1997a; Alcala & Brown, 1999), does not include thePl. corrugatus clade (Tagomukhus) or the morphologi-cally similar terrestrial, species Pl. pygmaeus andPl. levigatus (Lahatnanguri) (Brown, Brown & Alcala,1997a; Brown et al., 1997b, 1999a; Alcala & Brown,1999).

EtymologyLupacolus is derived from the combination of theTagalog noun Lupa, meaning ‘ground’ or ‘terrestrial’and the Greek colos, meaning ‘inhabitants’ or ‘dwell-ers’ in reference to the largely terrestrial microhabitatof the included species. It is masculine in gender. Sug-gested common name: Philippine forest ground frogs.

GENUS CORNUFER TSCHUDI, 1838

Type speciesHalophila vitiensis Girard, 1853 by subsequent des-ignation, following Opinion 1104 of the Commission(Anonymous, 1978).

DiagnosisMembers of the genus Cornufer can be distinguishedfrom species of the genus Alcalus by the presence of(1) median subgular vocal sacs (vs. absence); (2) absenceof nuptial pads (vs. presence); (3) presence of super-numerary tubercles on hands (vs. absence); (4) pres-ence of metatarsal tubercles beneath feet (vs. absence);and (4) absence or presence but highly reduced ofinterdigital webbing (vs. presence in species of Alcalusand members of Cornufer, subgenera Discodeles andPotamorana).

Although species of Cornufer, subgenera Cornufer,Potamorana, Ceratobatrachus, Discodeles, Palmatorappia,and Batrachylodes are phenotypically diagnosable



from species of the genera Alcalus and Platymantis(see diagnoses of those clades), species of Cornufer(subgenus Aenigmanura) and former members ofSolomon−Bismarck−Admiralty, Palau, Papuan, andeastern Indonesian ‘Platymantis’ (Table 3; see speciesnot assigned to subgenus) cannot be readily distin-guished from species of the genus Platymantis on thebasis of any one morphological character. We are unawareof any morphological synapomorphies for this clade,although it is strongly supported (Fig. 2, Clade E).

Phylogenetic definitionCornufer (CCN) is a maximum crown-clade name re-ferring to the crown clade (Fig. 2, Clade E) originat-ing with the most recent common ancestor ofCo. vitiensis and all extant species that share a morerecent common ancestor with Co. vitiensis than withAl. mariae or Pl. corrugatus.

ContentSpecies of the subgenera (clades) Potamorana, Cornufer,Ceratobatrachus, Palmatorappia, Discodeles,Batrachylodes, Aenigmanura, and species of the Pacific(non-Philippine) clade, genus Cornufer, formerly re-ferred to ‘Platymantis’ and not assigned to subgenusor subclade within Cornufer (Fig. 2; Table 3).

CommentUpon discovering that the overlooked type of Cornufer(Cornufer unicolor Tschudi, 1838) was in fact a speciesof the Neotropical taxon Eleutherodactylus, Zweifel(1966) petitioned the Commission to suppress the namesCornufer and its type species Cornufer unicolor Zweifel,1967, to avoid synonymy of Eleutherodactylus withinCornufer. His argument was that this discovery wouldrequire the assignment of the > 200 species ofEleutherodactylus to Cornufer. Suppression of Cornuferwould mean that the next available name for that groupof ranoids would, at the time, have been PlatymantisGünther, 1858.

Darlington et al. (1967) countered that Cornufer shouldnot be suppressed and that both names, Cornufer andPlatymantis, should be retained as available becauseCornufer had been widely used for some ranoid species.Additionally, the non-overlapping geographical distri-butions of Cornufer (east of Wallace’s Line) andPlatymantis (west of Wallace’s Line) strengthened theargument that both genera should be retained as valid(Darlington et al., 1967).

Prior to Zweifel (1966, 1967), Cornufer and Platymantiswere commonly used (Boulenger, 1918b; Taylor, 1920;Noble, 1931; Gorham, 1965; but see Inger, 1954). Al-though Zweifel (1967: 117) stated that ‘the name Cornuferis unavailable’ (and he was largely followed by workingtaxonomists), the Commission had not yet ruled on hisrequest (Zweifel, 1966) to suppress this name. A decade

later, the Committee ruled against his proposal(Anonymous, 1978) and eventually held that Halophilavitiensis Girard, 1853, be designated as the type speciesof Cornufer and that this genus should be considereda junior subjective synonym of Platymantis, which ‘. . .isto be given precedence over Cornufer Tschudi, 1838, byany zoologist who considers the type-species of thosenominal genera to belong to the same taxonomic genus(Anonymous, 1978; italics added).’ The Committee alsosuppressed all previous designations of the type speciesof Cornufer. Importantly, Cornufer was not sup-pressed; both names remain available and may be usedeither as genera or subgenera.

Given our choice not to place these two type species(Co. vitiensis and Pl. pliciferus, the latter currently asynonym of Pl. corrugatus) in the same genus, and thatthe name Cornufer Tschudi, 1838, remains available(Anonymous, 1978), we recognize both Platymantis (westof Wallace’s Line, i.e. Philippine species, excludingAl. mariae) and Cornufer (all species east of Wal-lace’s Line, i.e. taxa from eastern Indonesia, NewGuinea, Palau, the Solomon Islands, theBismarck−Admiralty archipelagos, and Fiji). Thesenames correspond to our newly defined clades (Fig. 2,Clades M and E, respectively).

Because relationships amongst some species of thegenus Cornufer have low support (Fig. 2), we have usedthe type species of subgenera as specifiers to ensurethat the content of Cornufer will remain stable.

EtymologyAlthough Tschudi (1838) provided no etymology forCornufer, we assume that the name is derived fromthe Latin ‘cornu’ meaning horn, and the Latin verb‘ferre’ (present infinitive), meaning to carry or bear,in reference to the presence of supraocular dermal tu-bercles in Co. vitiensis (the type species). Suggestedcommon names: Fijian ground frog (Cornufer vitianus),Fijian tree frog (Co. vitiensis).

SUBGENUS CORNUFER TSCHUDI, 1838

DiagnosisMembers of the subgenus Cornufer differ from othermembers of the genus Cornufer by having (1) a largemale body size (65–150+ mm SVL, vs. male body sizeusually ∼25–40 mm); (2) terminal discs of fingers andtoes non- to minimally expanded in Co. vitianus (vs.widely expanded in some arboreal riddle frogs of sub-genus Aenigmanura, palm frogs of subgenusPalmatorappia, giant water frogs of subgenus Discodeles,and a few sticky-toed frogs of subgenus Batrachylodes),or widely expanded in Co. vitiensis (vs. non- to mini-mally expanded in some terrestrial riddle frogs of sub-genus Aenigmanura, horned frogs of subgenusCeratobatrachus, river frogs of subgenus Potamorana,



and a few sticky-toed frogs of subgenus Batrachylodes).Additionally, both species are restricted to the islandsof Fiji, where they are the only native ranoid frogs;they do not possess overlapping distributions with anyother known ceratobatrachids. We are unaware of anymorphological synapomorphies for this group, al-though our molecular data clearly provides strongsupport for Fijian frogs as a monophyletic group.

Conversion of the name Cornufer (referring to thesubgenus) to a phylogenetic name would result in twodifferent clades bearing the name Cornufer. There-fore, we define a new clade name denoting the sameclade (Fig. 2, Clade E) as the subgenus Cornufer.

ContentCornufer vitiensis, Co. vitianus, and (provisionally) theextinct taxon Cornufer megabotoniviti (Worthy, 2001;Table 3). The subgenus Cornufer is equivalent in contentto the unranked taxon Yanuboto.

YANUBOTO NEW CLADE NAME

Phylogenetic definitionYanuboto (NCN) is a node-based name referring to theclade (Fig. 2, Clade T) originating with the most recentcommon ancestor of Co. vitiensis and Co. vitianus (bothspecies formerly in Platymantis).

CommentThe two living species of Fijian ceratobatrachids(Yanuboto) possess nearly a full complement of theecomorphological variation in the genus Cornufer(Brown, 2004). Cornufer vitiensis is a fully arborealtree frog characterized by widely expanded terminaldiscs of the fingers and toes and an arborealmicrohabitat preference and Co. vitianus is a large-bodied, fully terrestrial ground frog (with narrowly tonon-expanded terminal finger and toe discs). Surpris-ingly, these morphologically and ecologically dispar-ate forms (Gorham, 1965, 1968; Morrison, 2003; Zug,2013) are sister species (Fig. 2).

The fossil Co. megabotoniviti is known only from FijianQuaternary deposits. Worthy (2001) allied it toCo. vitianus and Co. vitiensis. Because of the lack ofsynapomorphies that ally it to the other Fijian species,we place it tentatively in Yanuboto (subgenus Cornufer)because of its provenance, but it would not be unrea-sonable to consider it unassigned to subgenus.

EtymologyYanuboto is derived from the Fijian terms yanuyanu,meaning ‘island’, and boto meaning ‘frog’, in refer-ence to the status of the included species status as theonly native anurans of Fiji. The name is masculine ingender. Suggested common names: Fijian ground frog(Co. vitianus), Fijian tree frog (Co. vitiensis).

POTAMORANA SUBGEN. NOV.Type speciesRana bufoniformis Boulenger, 1884.

DiagnosisRiver frog species of the subgenus Potamorana differfrom other subgenera of Cornufer, except giant waterfrogs of the subgenus Discodeles, and Fijian frogs sub-genus Cornufer (= Tanuboto) by having (1) a large bodysize (males 50–75 mm SVL; females 65–140; vs. mostspecies male SVL ∼25–40 mm; (2) moderately exten-sive, but reduced compared with Co. (Discodeles) guppyi,interdigital webbing of feet (vs. highly reduced to ves-tiges (Cornufer nexipus) or absent (all other species);(3) extensive rugosity of dorsal body skin (vs. smooth,weakly rugose, or slightly shagreened body skin); (4)non-expanded terminal discs of fingers and toes (vs.widely expanded in some arboreal riddle frogs of sub-genus Aenigmanura, palm frogs of subgenusPalmatorappia, giant water frogs of subgenus Discodeles,and a few sticky-toed frogs of subgenus Batrachylodes);and (5) semiaquatic microhabitat preferences (vs. ter-restrial). Based on the phylogeny, we consider theirlarge body size, interdigital webbing of the feet, andsemiaquatic microhabitat preferences to be shared,derived characters that unambiguously distinguish thespecies of Potamorana from all other species of Cornuferexcept Discodeles guppyi, in which these characters mostlikely have independently evolved (Fig. 2).

Phylogenetic definitionPotamorana (NCN) is an apomorphy-based name forthe clade (Fig. 2, Clade F) originating in the ancestorof Cornufer bufoniformis and Cornufer malukuna inwhich the following apomorphy, synapomorphic withthat in the various populations of Co. bufoniformis, origi-nated: moderately extensive webbing between the digitsof the feet.

ContentCornufer malukuna and Co. bufoniformis (Table 3). Wedid not sample Cornufer (Discodeles) opisthodon orCornufer (Discodeles) vogti but we tentatively place themin Potamorana because these species share thesynapomorphy (moderately extensive webbing betweenthe digits of the feet) of the clade Potamorana.

CommentThe newly discovered relationships of the former‘Discodeles’ malukuna and ‘Discodeles’ bufoniformisreveal that Discodeles was polyphyletic in its formersense. These species are unrelated to the clade (Fig. 2,Clade H) containing the type species D. guppyi. In retro-spect, it is not surprising that these four morphologi-cally similar (moderate body size, moderate interdigital



webbing, terminal digital discs non-expanded) speciesare now recognized as distinct from Co. (Discodeles)guppyi, a much larger species with full interdigitalwebbing between the toes and moderately expandedterminal digital discs.

EtymologyDerived from the Greek term potamo, meaning ‘riveror stream’, and the Latin rana, meaning ‘frog’, in ref-erence to the semiaquatic habitat preferences of thenew clade. The name is feminine in gender. Suggest-ed common name: Solomon−Bismarck river frogs.

SUBGENUS CERATOBATRACHUS BOULENGER, 1884

Type speciesCeratobatrachus guentheri Boulenger, 1884, by monotypy.

DiagnosisThe sole species of the subgenus Ceratobatrachus,Cornufer (Ceratobatrachus) guentheri, is one of the mostcharismatic and distinctive species in theCeratobatrachidae and is readily diagnosed from allmembers of the genus Cornufer by having (1) elabo-rated ossification of the squamosal (vs. absence ofornamental ossification); and by the (2) presence of man-dible odontoids (vs. absence); and (3) presence of ornatedermal protuberances above the eyes (‘horns’), at thetip of the snout, and along the posterior edges of fore-and hindlimbs (vs. absence). These characters areuniquely derived in this lineage (Fig. 2, Clade S).

Phylogenetic definitionCeratobatrachus is an apomorphy-based name for theclade (Fig. 2, Clade S) originating in the ancestor inwhich the following apomorphy, synapomorphic withthose in the known populations of Co. (Ce.) guentheri,originated: ornate dermal protuberances above the eyes(‘horns’), at the tip of the snout, and in the form ofserrated flaps along the outer edges of the limbs.

Content: Cornufer (Ce.) guentheri (Table 3).

CommentCornufer (Ce.) guentheri is most closely related to theextremely phenotypically dissimilar miniaturized speciesCornufer acrochordus (Fig. 2). This bizarre and com-pletely unexpected relationship stands as a testa-ment to the highly variable and at times bewilderingpatterns of morphological variability and phylogeneticrelationships in the family Ceratobatrachidae.

EtymologyAlthough Boulenger (1884) provided no etymology forCeratobatrachus, the name is probably derived fromthe Greek ‘kerato’, meaning ‘horned’ and the Greek

‘batrachos’, meaning ‘frog’. Suggested common name:Solomon Islands horned frogs.

SUBGENUS DISCODELES BOULENGER, 1918

Type speciesRana guppyi Boulenger, 1884.

DiagnosisThe sole species of the subgenus Discodeles is easilydiagnosed from species of the genus Cornufer by having(1) an extremely large body size [females up to 250 mmSVL (mass of up to 1 kg) vs. most species with femaleSVL ≤ 65 mm]; (2) moderately expanded terminal discsof fingers and toes (vs. widely or non-expanded); (3)fully webbed feet (vs. interdigital webbing absent, limitedto basal vestige, or present but with one or two ter-minal phalanges free of web); and (4) aquaticmicrohabitat preference (vs. terrestrial or arboreal). Weconsider its body size and full interdigital foot webbingto be synapomorphies of this distinct lineage (Fig. 2,Clade H).

Phylogenetic definitionDiscodeles is the apomorphy-based name for the clade(Fig. 2, Clade H) originating in the ancestor in whichthe following apomorphies, synapomorphic with thatin the known populations of D. guppyi, originated: ex-tremely large body size and fully webbed feet.

ContentComposed of highly divergent isolated allopatric andinsular lineages of the nominal species, Co. (D.) guppyiis most likely a complex of evolutionary lineages (species)from New Britain, Bougainville, and various SolomonIslands populations (Table 3).

CommentTwo species of Ceratobatrachidae have the specificepithet guppyi: Rana guppyi Boulenger, 1884 (the typespecies of the aquatic genus Discodeles) and Cornuferguppyi Boulenger, 1884 (a tree frog native to theSolomon Islands). Our inclusion of the two species inthe resurrected genus Cornufer creates homonymybetween the names. Under the principle of priority(ICZN, 1999) we normally would retain the seniorhomonym, the older available name. However, bothspecies were named in the same year, in the same workand on the same page (Boulenger, 1884: 211), an ex-tremely unusual situation.

Under the Code, the preferred and most conserva-tive action would be the substitution of a valid juniorsynonym of one of these species. Rana guppyi Boulenger,1884, purportedly has a junior synonym; Zweifel (1960)treated Rana bufoniformis cognata Hediger, 1933(NHMB 4605, holotype; Forcart, 1946) as a synonym



of R. guppyi Boulenger, 1884. However, we reject cognataas a junior synonym of R. guppyi because its type lo-cality, ‘Iriu’, Admiralty Islands, falls within the knowngeographical range of ‘D.’ vogti and not within that ofD. guppyi. Hediger’s (1933) description additionally listsmorphological character states (narrowly expanded toediscs, relatively small body size) that lead us to believethat this species is not referable to Co. (D.) guppyi. Wetherefore consider R. bufoniformis cognata Hediger, 1933,as a junior subjective synonym of D. vogti (Hediger,1934). Thus, there is no junior synonym that can besubstituted for R. guppyi Boulenger, 1884.

The second species in this conundrum is Co. guppyiBoulenger, 1884, which also lacks any junior syno-nyms. However, the Code provides for a resolution insuch cases. Article 24.2 of the Code states that the prin-ciple of first reviser (ICZN, 1999:30) is to be used insituations in which the precedence between namescannot be determined and an available junior synonymdoes not exist. Acting as first reviser, we fix prec-edence of R. guppyi Boulenger, 1884, over Co. guppyiBoulenger, 1884. This action maintains the name ofthe well-known species D. guppyi (= Rana guppyiBoulenger, 1884), which is also the type species ofDiscodeles. We provide a new replacement name forCo. guppyi Boulenger, 1884, below (see under subge-nus Aenigmanura).

EtymologyAlthough Boulenger (1918b) provided no etymology forDiscodeles, he distinguished it from other Papuan andMelanesian forms on the basis of the ‘horseshoe-shaped groove’ (Boulenger, 1918b:238) evident on thetips of fingers and toes. Thus, we assume that the nameis derived from the Latin ‘discus’, meaning a flat andround shape, and the Greek ‘delos’, meaning visibleor evident, in reference to presence of the digital discs.Suggested common name: giant Pacific water frogs.

SUBGENUS PALMATORAPPIA AHL, 1927

Type speciesHylella solomonis Sternfeld, 1920.

DiagnosisThe single species Cornufer (Palmatorappia) heffernani(formerly Palmatorappia solomonis; see below) can bereadily diagnosed from other members of the genusCornufer by having (1) a small, delicate, slender bodyand limbs (vs. more robust body form and limbs); (2)widely expanded terminal discs of fingers and toes (vs.non- to minimally expanded in some terrestrial riddlefrogs of subgenus Aenigmanura, horned frogs of sub-genus Ceratobatrachus, river frogs of subgenusPotamorana, and a few sticky-toed frogs of subgenusBatrachylodes); (3) flattened subarticular tubercles of

hands and feet (vs. subarticular tubercles rounded topointed); (4) moderate interdigital webbing of fingers(unique amongst species of the genus Cornufer) andtoes (present as vestiges in Co. (Aenigmanura) nexipus(vs. absent or much more extensive); and (5) interdigitalwebbing extensive in Co. (D.) guppyi but moderate (oneor two terminal phalanges free) in species of the sub-genus Potamorana. We consider this suite of charac-ters to be uniquely derived within Cornufer. Based onour phylogeny (Fig. 2), moderate interdigital webbingof the manus appears to be a unique apomorphy dis-tinguishing Palmatorappia from all otherceratobatrachids.

Phylogenetic definitionPalmatorappia is an apomorphy-based name for theclade (Fig. 2, Clade U) originating in the ancestor inwhich the following apomorphy, synapomorphic withthat in the various populations of Palmatorappiaheffernani, originated: moderate interdigital webbingof the fingers.

ContentCornufer (Pa.) heffernani (Kinghorn, 1928); formerlya junior synonym of Pa. solomonis (Sternfeld, 1920);here designated a nomen substitutum; see below(Table 3).

CommentIn general phenotypic characteristics and microhabitatpreferences, the sole species of the subgenusPalmatorappia is unlike any other Solomon memberof the genus Cornufer and, in fact, phenotypically andecologically much more closely resembles the unrelat-ed members of the clade Platymantis (Tirahanulap) ofthe Philippines (formerly referred to as the Platymantishazelae Group, sensu Brown et al., 1997a) and speciesof Cornufer (Aenigmanura) from the mountains of NewBritain Island (Co. macrosceles, Co. citrinospilus,Co. mamusiorum) and Manus Island (Co. custos).

The allocation of the Solomon Islands palm frog,Pa. solomonis, originally Hylella solomonis, andPlatymantis solomonis (Boulenger, 1884) (a wide-spread Solomon Islands ground frog), originally Cornufersolomonis, to the genus Cornufer creates homonymy.That the identical species names belong to differentsubgenera within Cornufer is not relevant to the issueof homonymy (Article 57.4). Following the principle ofpriority we retain the senior homonym Co. solomonisBoulenger, 1884; in our classification the new combi-nation is Cornufer (Aenigmanura) solomonis.

Hylella solomonis Sternfeld, 1920, is the type andonly species of Palmatorappia Ahl, 1927; the princi-ple of homonymy requires that this junior homonymbe replaced even though it would be desirable to main-tain the name of the type species in the interest of



stability. The only available junior synonym ofPa. solomonis is Hypsirana heffernani Kinghorn, 1928,which was synonymized under Pa. solomonis by Brown(1952). Therefore, we designate Hyps. heffernaniKinghorn, 1928, as a substitute name for Hyl. solomonisSternfeld, 1920. The species commonly known asPa. solomonis will be Cornufer (Palmatorappia)heffernani comb. nov.

EtymologyMost likely from the Latin ‘palmat-’, meaning the con-dition in which the spaces between the digits are filledin (as by webbing), and Rappia, a patronym for Rapp,who named the genus Hyperolius. Günther (1865) un-justifiably proposed Rappia as a substitute name forHyperolius Rapp, 1842, so Rappia is a patronym andthus Palmatorappia is to be treated as masculine.Common name: Solomon Islands palmate frogs.

SUBGENUS BATRACHYLODES BOULENGER, 1887

Type speciesBatrachylodes vertebralis Boulenger, 1887.

DiagnosisSpecies of the subgenus Batrachylodes form a pheno-typically and ecologically cohesive group, differing fromother members of the genus Cornufer by having (1) asmall body size (males 17–24 mm SVL; vs. ≥ 25 mm);(2) stout, triangular bodies (vs. body shape slender, nottriangular); (3) pointed snouts (vs. rounded); and (4)slightly expanded to widely expanded terminal discsof fingers and toes (vs. terminal discs non-expanded);and by the (5) presence of darkened loreal stripes con-tinuing diagonally across the flank to form a distinct-ly stratified lateral body marking (i.e. clearly demarcateddarker dorsal and lighter ventral colours) in most species(vs. absence); and (6) absence of interdigital webbing(vs. presence in Potamorana and Discodeles). We con-sider body shape (microhylid-like; generally triangu-lar bodies with very small heads and strongly pointedsnouts) and stratified coloration (light above, dark onlateral surfaces) to be synapomorphies for the subge-nus (Boulenger, 1887; Sternfeld, 1920; Brown & Parker,1970), which is strongly supported in our phylogeny(Fig. 2, Clade L).

Phylogenetic definitionBatrachylodes is an apomorphy-based name for the clade(Fig. 2, Clade L) originating in the ancestor of Cornufer(Batrachylodes) vertebralis and Cornufer (Batrachylodes)trossulus, in which the following apomorphy,synapomorphic with that in Batrachylodes vertebralis,originated: very small, triangular bodies with smallheads and strongly pointed snouts (Fig. 2).

ContentSeven species formerly referred to the genusBatrachylodes (i.e. excluding Cornufer minutus; Fig. 2,and below), exclusively from the Solomon Islands (Brown& Parker, 1970: Cornufer elegans, Co. gigas, Co.mediodiscus, Co. montanus, Co. trossulus, Co. vertebralis,and Co. wolfi; Table 3). Brown et al. (2013) discussedan undescribed species from New Britain Island, Bis-marck Archipelago (the first report of a species of thisgenus outside the Solomon Island Archipelago;Foufopoulos & Richards, 2007).

CommentThe species not sampled by us (B. elegans, B. gigas,B. mediodiscus, and B. montanus) from the morpho-logically cohesive and biogeographically circum-scribed Batrachylodes are also placed in Batrachylodesbecause they share the synapomorphy on which thephylogenetic name is based. We exclude Co. minutusfrom this group on the basis of its unstable phylogeneticaffinities (Fig. 2), which, in the combined data setsuggest a closer relationship to Melanesian (Cornufersp. Halmahera, Cornufer batantae, and Cornuferbimaculatus) species than to members of the subge-nus Batrachylodes, with the caveat that support forthis relationship is low (Fig. 2).

EtymologyAlthough Boulenger (1887) provided no etymology forBatrachylodes, the name is most likely derived fromthe Greek ‘batrachus’, meaning frog, and possibly‘hylodes’, in reference to the genus Hylodes. Boulenger’s(1882) concept of Hylodes included 45 species that aretoday allocated to Pristimantis, Eleutherodactylus,Lithodytes, Batrachyla, and other genera. One ofBoulenger’s diagnostic characters for Hylodes was ex-panded digital discs, such as are present in someBatrachylodes species. Hylodes is almost certainlyderived from Hyla- + ‘-odes’ (Greek), meaning like orsimilar to Hyla, implicitly with expanded discs. Commonname: Solomon Islands sticky-toed frogs.

AENIGMANURA SUBGEN. NOV.Type speciesPlatymantis papuensis schmidti Brown & Tyler, 1968.

DiagnosisIndividual species of the subgenus Aenigmanura differfrom other members of Cornufer by characters relatedto their general classification as either generalized ter-restrial species with narrow finger and toe discs or ar-boreal forms with widely expanded finger and toe discs.The arboreal tree frogs of Aenigmanura (Cornufercitrinospilus, Co. custos, Co. hedigeri [formerlyPlatymantis guppyi; see below], Co. macrosceles, Co.



mamusiorum, Co. nakanaiorum, Co. neckeri, Co. nexipus,Co. parilis, Co. sp. B. Manus and Co. sp. C. Manus)can be readily distinguished from ground frogs (se-lected members of subgenus Aenigmanura and allCornufer species not assigned to subgenera, all withnon-expanded discs of fingers and toes), aquatic species(subgenera Potamorana and Discodeles, characterizedby the presence of interdigital webbing), sticky-toedfrogs (subgenus Batrachylodes, small, triangular-shaped bodies with strongly pointed snouts), the Fijianground frog (Co. vitianus, with non-expanded discs offingers and toes), palm frogs (subgenus Palmatorappia,with interdigital webbing present on hands), and hornedfrogs (subgenus Ceratobatrachus, with elaborately casquedskull morphology and dermal horns above the eyelids).The terrestrial species of Aenigmanura (Cornuferadiastolus, Cornufer admiraltiensis, Cornuferakarithymus, Cornufer boulengeri, Cornufer bufonulus,Cornufer desticans, Cornufer gilliardi, Cornufer latro,Cornufer magnus, Cornufer papuensis, Cornuferpelewensis, Cornufer schmidti, Cornufer solomonis,Cornufer sulcatus, and Cornufer weberi) can be dis-tinguished from river frogs and giant frogs (subgeneraPotamorana and Discodeles, characterized by the pres-ence of interdigital webbing), sticky-toed frogs (sub-genus Batrachylodes, small, triangular-shaped bodieswith strongly pointed snouts), the Fijian tree frog(Co. vitiensis, with widely expanded discs of fingers andtoes), palm frogs (subgenus Palmatorappia, an arbor-eal species with expanded finger and toe discs, andinterdigital webbing present on hands and feet), andhorned frogs (subgenus Ceratobatrachus, with elabo-rately casqued skull morphology and dermal horns abovethe eyelids).

As implied by the name, the wide range of morpho-logical and ecological variation in this clade rendersan unambiguously exclusive diagnosis of Aenigmanuraimpossible. We are unaware of morphologicalsynapomorphies for this group, although ourphylogenetic analysis provides very strong support forthis phenotypically and ecologically diverse clade (Fig. 2,Clade J).

Phylogenetic definitionAenigmanura (NCN) is a maximum crown-clade namereferring to the crown clade (Fig. 2, Clade J) originat-ing with the most recent common ancestor ofCo. papuensis and all extant species that share a morerecent common ancestor with Co. papuensis than withany of the other species of the clade Cornufer. Alter-natively it can be conceived of as the largest crownclade containing Co. papuensis, but not any other speciesof the clade Cornufer.

ContentCornufer adiastolus, Co. admiraltiensis, Co. akarithymus,Co. boulengeri, Co. citrinospilus, Co. custos, Co. desticans,

Co. gilliardi, Co. hedigeri (formerly Pl. guppyi;see below), Co. latro, Co. macrosceles, Co. magnus,Co. mamusiorum, Co. nakanaiorum, Co. neckeri,Co. nexipus, Co. parilis, Co. papuensis, Co.pelewensis, Co. schmidti, Co. solomonis, Co. sulcatus,Co. weberi, the newly described Co. custos (Richardset al., 2014), and two undescribed species fromManus Island (sp. B Manus and sp. C Manus; Fig. 2.,Clade J).

CommentThe range of body sizes in this large clade is strik-ing. From miniaturized terrestrial species such asCo. sulcatus and Co. akarithymus (males 17–27 mmSVL), to giant ground species such as Co. magnus(males 75–150 mm SVL), to large canopy frogs suchas Co. neckeri and Co. hedigeri (formerly Pl. guppyi;see below), to delicate, high-elevation, arboreal shrubspecies such as Co. macrosceles and Co. mamusiorum,to widespread terrestrial generalists such asCo. papuensis, Co. weberi, Co. schmidti, andCo. solomonis – the subgenus Aenigmanura exhibitsnearly the full range of ceratobatrachid ecomorphologicaldiversity (Brown, 2004), all within one clade of closelyrelated species.

As noted above, allocation of Pl. guppyi Boulenger,1884 (not to be confused with D. guppyi) to the genusCornufer presented a case of secondary homonymywith respect to R. guppyi Boulenger, 1884. Given thatno available junior synonym exists for the latterand that it is also the type species of Discodeles, weelected not to alter this name, and we have givenR. guppyi precedence over Pl. guppyi following the prin-ciple of first reviser (ICZN, 1999:30). Thus, the estab-lishment of a replacement name for Pl. guppyiBoulenger, 1884, is necessary. Accordingly, we desig-nate Co. hedigeri as a nomen novum for Pl. guppyiBoulenger, 1884. The epithet hedigeri is a patronymfor Heine Hediger (1908–1992) in recognition of hiscontributions (Hediger, 1933, 1934) to the taxonomyof the genus Cornufer sensu lato and the biology ofthe South Pacific.

EtymologyFrom the Latin enigma, meaning something ‘obscureor unknown, a riddle’, and anura, meaning ‘frog’, inreference to the unanticipated and confusing range ofmorphological and ecological variation represented bythe closely related species of the new subgenus. Thename is masculine in gender. Suggested common name:Pacific Island riddle frogs.

DISCUSSIONPHENOTYPIC CHARACTERS AND DIAGNOSES

The phylogenetic framework for our new ceratobatrachidclassification was derived from a new multilocus DNA



sequence data set. Although some phenotypic charac-ters are easily identified as synapomorphies (e.g. thedermal ‘horns’ of Ceratobatrachus), we have not com-prehensively surveyed phenotypic characters to deter-mine their value as possible synapomorphies, and inmany cases it is not possible to provide informationthat will place a species within a clade without usingDNA sequences. As a result, some unsampled speciesand/or species of uncertain phylogenetic affinities(Figs. 2, 3), are not yet referable to subgenera (Table 3).

Our approach takes a top-down perspective in thatwe have begun with a phylogeny and will progres-sively incorporate information about phenotype. Thenext steps in our studies of ceratobatrachid evolutioninclude the description of many new species, with acomprehensive survey of external morphology andphylogenetic analysis of advertisement calls. Our ex-perience with these frogs suggests that we will gleanmany synapomorphies from the phenotype and acous-tic data. Integrating these new data into thisphylogenetic framework will provide a broader viewof ceratobatrachid evolution.

TAXONOMY

Our phylogeny of the Ceratobatrachidae is a major steptowards the development of a stable taxonomy for this

poorly understood clade of frogs from Southeast Asiaand the Pacific islands. Clearly Platymantis as previ-ously defined is not monophyletic (Fig. 3A). Setting asidefor the moment the genus Alcalus (formerly South-east Asian species of Ingerana) as uncontroversial, wecarefully considered the following options for the tax-onomy of Clade D (Fig. 2).

1. Recognizeone genus Platymantis for Clade D, withno subgenera. This would subsume several species-poor genera such as Discodeles, Palmatorappia, etc.as junior synonyms of Platymantis. However, a ‘flat’taxonomy such as this would conceal the phylogenetichierarchy elucidated by this study. Because > 80%of ceratobatrachid species are Platymantis, few genus-species combinations would change.

2. Recognize one genus Platymantis for Clade D, withseveral subgenera, many of them new. This wouldconvey both phylogenetic hierarchy and diversityin morphology, ecology, and biogeography. It wouldalso retain the use of well-known clade names suchas Discodeles, Ceratobatrachus, Batrachylodes, andPalmatorappia, which are associated with widelyappreciated and distinct phenotypes. Similar to (1),few genus-species combinations would change.

3. Split Clade D into several genera. This woulddissolve Platymantis, reduce its content, and so

Figure 3. A schematic representation of ceratobatrachid phylogeny (based on Fig. 2), summarizing (A) the previous tax-onomy of ceratobatrachid frogs and (B) the new classification scheme proposed here.



re-allocate many species to different genera.Numerous genus-species combinations wouldchange.

4. Adopt an intermediate option, which is to recognizetwo genera, Cornufer and Platymantis, for Clade D.Subgenera would be used to reflect hierarchical struc-ture and biotic diversity. The number of changes ingenus-species combinations would be fewer than inoption (3) but more than in (1) and (2).

We have adopted option (4), but we can be criti-cized for not using option (2), and in fact some of theauthors (D. C. C. vs. R. M. B.) disagree on this choice.The latter would maintain the generic name Platymantisfor the large number of species in Clade J(Aenigmanura) and minimize changes in combina-tions. Our use of the Cornufer−Platymantis arrange-ment (Fig. 3B) increases the number of changes ingenus-species combination, although not as much asoption (3), but emphasizes a trenchant biogeographi-cal pattern between Clade M and Clade E, each situ-ated on either side of Wallace’s Line.

Under options (2) and (4) subgeneric ranks could beused. Under option (2), one might use two subgen-era, Platymantis and Cornufer, with smaller taxa withineach. These less inclusive taxa might be ranked as sec-tions or series. According to the Code (Article 10.4; ICZN,1999) such ranks are treated as subgenera. Nested levelsof subgenera are an ideal but underused way to provideadditional hierarchical information that is not evidentin ‘flat’ taxonomies; for an example see Hillis et al.(2001). However, the use of nested subgenera isproblematic under the Code (see Dubois, 2007;Hillis, 2007), which reflects the Code’s non-evolutionaryorigins.

Although subgenera are an excellent means of en-hancing phylogenetic information in taxonomy, they haveproblems that derive from the Code’s emphasis on ranks.For example, although the proper form ofCeratobatrachus guentheri under our taxonomy isCornufer (Ceratobatrachus) guentheri, Ce. guentheri aloneunambiguously refers to that species without explicitmention of the subgenus rank. Unfortunately, the Codeprohibits omission of the genus name when the sub-genus name is used (Article 4.2), but ignoring thisrule has little negative effect if the name is used incontext. Additionally, monotypic subgenera (e.g.Ceratobatrachus in our taxonomy) add no informa-tion about relationships to other taxa, but we retainthese names to connect the species epithets to previ-ous taxonomies.

A second problem is that a subgenus containing thetype species of the genus must be denoted by the samename as the genus (Articles 43.1 and 44.1). For example,from its creation Platymantis exists both as a genusand subgenus name, and simple reference to

‘Platymantis’ is ambiguous as to rank. A simple solu-tion is to define a new, unranked name in place of thesubgeneric name, as we have done (Table 4), so thatthe name Platymantis refers to only one node.

It is possible that a future worker will propose raisingthe subgenera to generic rank. We feel that this actionwould be ill-advised and unwarranted because it wouldresult in changes in a large number of genus–speciescouplet names. The practice of unnecessarily split-ting a genus into several genera destabilizes taxono-my and hides nested phylogenetic information [see forexample the proposal to split Anolis by Nicholson et al.(2012) and responses by Poe (2013) and R. Glor (unpubl.data)]. In many cases of oversplitting, the possibilityof using subgenera is typically not considered or is re-jected without discussion.

In weighing the goals of naming diversity that cor-responds to phenotypic or geographical distinctiveness(and with the goals, some may feel, of optimizing edu-cational and conservation benefits that may be asso-ciated with more atomized classification), vs. avoidingan excess of names of equal rank owing to the splittingof clades (Cannatella & de Queiroz, 1989; Glaw, Vences& Böhme, 1998; Vences et al., 2000; Vences & Glaw,2001; Glaw & Vences, 2006; Glaw, Hoegg & Vences,2006; Pauly, Hillis & Cannatella, 2009; Poe, 2013), wehave adopted a compromise between changing genus–species couplets and retaining the presumed inten-tions of earlier taxonomists (Tschudi, 1838; Günther,1858; Boulenger, 1884, 1887, 1896, 1918a; Ahl, 1927)who apparently recognized, appreciated, and formallynamed the morphological, biogeographical, and eco-logical distinctiveness of the taxa (Noble, 1931; Gorham,1965). We feel that this compromise both recognizesthe marked diversity within the Ceratobatrachidae, andalso imparts a stable hierarchical classification that isconservative in that it requires relatively few changesto existing species names (Fig. 3).

Challenges exist for improved understanding of therelationships of ceratobatrachids. First, additional taxonsampling will provide new information to this initialestimate of phylogeny. With the addition of possibly40–65 undescribed species (R. M. Brown, S. J. Rich-ards, A. C. Diesmos & C. D. Siler, unpubl. data), somerelationships and classification schemes will prob-ably change. Additionally, poor resolution amongst thesubgenera Ceratobatrachus, Discodeles, Potamorana,Batrachylodes, Palmatorappia, Aenigmanura, and thespecies of Cornufer not assigned to subgenera willrequire additional gene sampling and taxonomic re-vision. For the meantime, we consider the classifica-tion of Ceratobatrachidae to be a work in progress (sensuGraybeal & Cannatella, 1995; Linkem, Diesmos &Brown, 2011) and we anxiously await future studiesthat will address the remaining problems identifiedhere.



ACKNOWLEDGEMENTS

We would like to express our gratitude to Angel C.Alcala and the late Walter C. Brown for encouragingus to work collaboratively on frogs of the familyCeratobatrachidae. For loans of specimens (Appendix1), access to genetic samples, use of photographs, and/or assistance in the field, we thank A. Allison, C. Austin,D. Bickford, R. Crombie, S. Donnellan, R. Drewes, B.Evans, R. Fisher, R. Günther, R. Inger, D. Iskandar,F. Kraus, T. LaDuc, A. Leviton, J. McGuire, C. Mor-rison, R. Norris, P. Pikacha, A. Resetar, I. Setiadi, andJ. Vindum. William E. Duellman provided commentson a previous version of the manuscript. We thank thePhilippine Department of Environment and NaturalResources (DENR), the Solomon Islands Forestry andEnvironment departments, the Indonesian Institute ofSciences (LIPI), and the Papua New Guinea Nation-al Research Institute and Department of Environ-ment and Conservation, for research, collecting, andexport permits. We thank K. de Queiroz for consulta-tion regarding phylogenetic taxonomy. Support for thisresearch was provided by South Australian Museumand Conservation International funds provided toS. J. R., a Rufford Foundation award to A. C. D.,Fulbright and Fulbright-Hayes fellowships to C. D. S.,and multiple U.S. National Science Foundation grants:DEB 0804115 to C. D. S.; DEB 9981631 and DEB0206729 to D. C. C.; and EB 073199, DEB 0640737,DEB 0743491, and EF 0334952 to R. M. B.

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12



SUPPORTING INFORMATION

Additional Supporting Information may be found in the online version of this article at the publisher’s web-site:

Figure S1. Partitioned Bayesian analyses (letters denote nodes of interest; see text and Figs. 2 and 3). Strong-ly supported conflict between nuclear [left; four partitions: recombinase activating gene 1 (RAG1), tyrosinase(Tyr), proopiomelanocortin (POMC), third codon positions combined) and mitochondrial (right; two partitions:12S and 16S) gene partitions is most probably an artefact of missing 12S data.



Multilocus phylogeny and a new classification for Southeast … · 2015. 4. 21. · Multilocus...

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