Class Amphibia Gray, 182. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classi­fication and survey of taxonomic richness

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ZOOTAXA
Animal biodiversity:
An outline of higher-level classification and survey of taxonomic richness
ZHI-QIANG ZHANG (ED.)
New Zealand Arthropod Collection, Landcare Research, Private Bag 92170, Auckland, New Zealand;
zhangz@landcareresearch.co.nz
Magnolia Press
Auckland, New Zealand
3148

ZHANG (ED.)
2 · Zootaxa 3148 © 2011 Magnolia Press
ZHI-QIANG ZHANG (ED.)
Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness
(Zootaxa 3148)
237 pp.; 30 cm.
23 Dec. 2011
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ISBN 978-1-86977-850-7 (Online edition)
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Zootaxa 3148 © 2011 Magnolia Press · 39
ANIMAL BIODIVERSITY
Class Amphibia Gray, 18251,2,3
Amphibia incertae sedis
†Albanerpetontidae Fox & Naylor, 1982 (†4)4,5
Order Anura Fischer von Waldheim, 1813 (frogs and toads) (410–466 genera; 6090 species; †~84)6
Family Allophrynidae Goin, Goin, & Zug, 1978 (1 genus; 1 species)7
Family Alsodidae Mivart, 1869 (3 genera; 32 species)8
1. By D.C. Blackburn & D.B. Wake (for full contact details, see the list after References). The title of this contribution should be cited as “Class
Amphibia Gray, 182. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness”.
2. The Amphibian Species of the World website (Frost, 2011), a continuation of Frost (1985), has guided our work; this site provides detailed
information on the taxonomic history of the names we discuss here. Our decisions on which taxa to recognize and at what level are based on current
literature and our assessment of current usage. Numbers of species used in our classification are based on the AmphibiaWeb website
(www.amphibiaweb.org; Accessed 13 October 2011).
3. Linnaeus (1758) used Amphibia for a different assemblage of taxa than currently recognized, and authorship for the currently understood taxon is
either de Blainville, 1816 (Dubois, 2004) or Gray, 1825. Frost et al. (2006) detail reasons for rejecting de Blainville as the authority. Phylogenetic
analyses based on molecular data for extant taxa support the monophyly of a clade containing Anura, Caudata, and Gymnophiona (e.g., Zardoya &
Meyer, 2001; San Mauro et al., 2005). Phylogenetic analyses of morphological characters, including from a wide-range of extinct taxa, support this
clade, with the extinct family Albanerpetontidae included in the crown group (e.g., Sigurdsen & Green, 2011; Ruta et al., 2003; Laurin & Reisz,
1997; Trueb & Cloutier, 1991b). However, some morphology-based analyses support non-monophyly of extant amphibians by placing the
Gymnophiona more closely to extant amniotes (e.g., Anderson, 2008; Anderson et al., 2008; but see Marjanovi & Laurin, 2008, 2009) than to
Anura and Caudata. Often the clade comprising extant amphibians has been referred to as Lissamphibia (Parsons & Williams, 1963; Romer, 1966;
Duellman & Trueb, 1986; Bolt, 1991; Trueb & Cloutier, 1991a,b; see also Frost et al., 2006), but we note that this view is not universally held
(Dubois, 1983, 2004). We refer to the most exclusive clade containing crown-group amphibians as Class Amphibia, although we recognize that
Amphibia is also often applied to extinct tetrapod taxa that are included neither within extant amniotes nor extant amphibians.
4. Throughout the manuscript the numbers of genera inclusive of extinct taxa, are listed, followed by the number of extinct (†) genera.
5. Historically, the †Albanerpetontidae has been allied to the extant orders of amphibians (Fox & Naylor, 1982; Milner, 2000; McGowan, 2002).
Phylogenetic analyses have reaffirmed a close relationship between †Albanerpetontidae, comprising four extinct genera (for recent summary see
Sweetman & Gardner, in press), and the extant orders Anura and Caudata, although the precise relationships remain uncertain (Anderson et al.,
2008; Ruta et al., 2003).
6. Many extinct anuran generic-level taxa exist, but most cannot be assigned with confidence to the families recognized here. A few are demonstrably
outside of crown-group Anura (e.g., †Czatkobatrachus, †Mesophryne, †Notobatrachus, †Prosalirus, †Triadobatrachus, †Vieraella,
†Yizhoubatrachus; Báez & Basso, 1996; Gao & Wang, 1998; Gao & Chen, 2004). When phylogenetic analyses or other evidence allow placement
of an extinct genus with some confidence within the crown-group of a family of living species, we have opted to include that extinct taxon within
that family. However, because of either a lack of thorough analyses or changing concepts of families, we cannot place many of these extinct taxa
within currently recognized families. These extinct taxa include †Altanulia, †Aralobatrachus, †Arariphrynus, †Avitabatrachus, †Aygroua,
†Comobatrachus, †Cordicephalus, †Cratia, †Elkobatrachus, †Eobatrachus, †Eopelobates, †Eophractus, †Eorubeta, †Estesiella, †Estesina,
†Estesius, †Eurycephalella, †Hatzegobatrachus, †Itemirella, †Kizylkuma, †Latonia, †Liaobatrachus, †Lithobatrachus, †Liventsovkia,
†Lutetiobatrachus, †Macropelobates, †Messelobatrachus, †Neoprocoela, †Neusibatrachus, †Nezpercius, †Opisthocoelellus, †Palaeophrynos,
†Paralatonia, †Pelophilus, †Pliobatrachus, †Proceratobatrachus, †Ranomorphus, †Ranavus, †Soevesoederberghia, †Sunnybatrachus,
†Thaumastosaurus, †Theatonius, †Thoraciliacus, and †Uldzinia; see Sanchíz (1998) for a review of most extinct anuran taxa.
7. Frost et al. (2006), avoiding families with only one genus, treated Allophryninae and Centroleninae as subfamilies of Centrolenidae. Phylogenetic
analyses support a sister relationship between these clades (Austin et al., 2002; Faivovich et al., 2005; Wiens et al., 2005; Frost et al., 2006;
Guayasamin et al., 2008), but the revision by Guayasamin et al. (2009) maintained Allophrynidae and Centrolenidae as separate families. We see
either as acceptable and viable taxonomies.
8. Pyron & Wiens (2011) were unable to obtain a robust topology of several genera once considered leptodactylids (see also Correa et al., 2006; Frost
et al., 2006; Grant et al., 2006) and recognized eight small families: Alsodidae, Batrachylidae, Ceratophryidae, Cycloramphidae, Hylodidae,
Odontophrynidae, Rhinodermatidae, and Telmatobiidae. Previously, Grant et al. (2006) refined the taxonomy of Frost et al. (2006) by recognizing
Hylodidae as distinct from the Cycloramphidae. Correa et al. (2006) resolved different relationships among these taxa, but used less comprehensive
sampling. Both Nuin and do Val (2005) and Heinicke et al. (2009), with limited taxon sampling, showed that Cycloramphidae was likely not
monophyletic. Frost et al. (2006) showed that Rhinoderma is nested within their Cycloramphidae; this was also suggested by Correa et al. (2006)
who found Rhinoderma to be the sister taxon of Insuetophrynus. Relationships among some of these genera based on morphological data were
discussed by Diaz & Valencia (1985), who included Caudiverbera (now Calpytocephalella) in this lineage, and Diaz (1989), with further details on
potentially useful diagnostic features within this group provided by Alcalde & Blotto (2006), Cárdenas-Rojas et al. (2007), and Rabanal & Formas
(2009). Grant et al. (2006) extended the taxonomy of Frost et al. (2006) by recognizing three subfamilies of Ceratophryidae. However, while
several phylogenetic studies have suggested that these subfamilies form a clade (Faivovich et al., 2005; Frost et al., 2006; Grant et al., 2006), others
have not (Darst & Cannatella, 2004; Wiens, 2005; Correa et al., 2006; Roelants et al., 2007; Heinicke et al., 2009; Pyron & Wiens, 2011; see also
Ruane et al., 2011). Other studies are consistent with monophyly but did not include taxa from all three subfamilies (Wiens et al., 2005). While
reporting monophyly, Frost et al. (2006) and Grant et al. (2006) differed in assessments of relationships among the subfamilies: Grant et al. (2006)
found a sister relationship between Telmatobiinae and Ceratophryinae whereas Frost et al. (2006) reported a sister relationship between
Batrachylinae and Ceratophryinae. Roelants et al. (2007) did not resolve Ceratophryidae as monophyletic, but reported a sister relationship between
taxa in the Batrachylinae and Telmatobiinae. Heinicke et al. (2009) also did not resolve Ceratophryidae as monophyletic, but instead found a sister
relationship between taxa in the Telamatobiinae and Ceratophryinae. Bossuyt & Roelants (2009) recognized two families, Telmatobiidae and
Ceratophyridae, yet did not specify the content of these families, which is crucial given the uncertainty in relationships among the subfamilies.
Given that Pyron & Wiens (2011) is the analysis to date with the most complete taxonomic sampling of taxa previously placed in the
Ceratophryidae, Cycloramphidae, and Hylodidae, we follow their elevation to family level of the three subfamilies of Ceratophryidae of Grant et al.
(2006). Córdova & Descailleaux (2005) provide a cladistic analysis of karyotypic data suggesting that Telmatobius is paraphyletic with respect to
Batrachophyrnus. For the families Alsodidae and Batrachylidae, we follow the genus-level revision of Pyron & Wiens (2011) that resulted in
placing Hylorhina and several Batrachyla species in Eupsophus.

BLACKBURN & WAKE: CLASS AMPHIBIA GRAY, 1825
40 · Zootaxa 3148 © 2011 Magnolia Press
Family Alytidae Fitzinger, 1843 (2 genera; 12 species; †6)9
Subfamily Alytinae Fitzinger, 1843 (1 genus; 5 species)
Subfamily Discoglossinae Günther, 1858 (1 genus; 7 species)
Family Aromobatidae Grant, Frost, Caldwell, Gagliardo, Haddad, Kok, Means, Noonan, Schargel & Wheeler,
2006 (5 genera; 103 species)10
Family Arthroleptidae Mivart, 1869 (8 genera; 144 species)11
Family Ascaphidae Fejérváry, 1923 (1 genus; 2 species)12
Family Batrachylidae Gallardo, 1965 (2 genera; 11 species)13
Family Bombinatoridae Gray, 1825 (2 genera; 10 species; †1)14
Family Brachycephalidae Günther, 1858 (2 genera; 49 species)15
Family Brevicipitidae Bonaparte, 1850 (5 genera; 32 species)16
Family Bufonidae Gray, 1825 (~10– ~48 genera; 568 species)17
Family Calyptocephalellidae Reig, 1960 (2 genera; 4 species)18
Family Centrolenidae Taylor, 1951 (12 genera; 150 species)19
Subfamily Centroleninae Taylor, 1951 (10 genera; 118 species)
Subfamily Hyalinobatrachinae Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, & Vilà, 2009 (2
genera; 32 species)
9. Analyses of molecular data support a clade containing Alytes, Barbourula, Bombina, and Discoglossus (Hay et al., 1995; Hoegg et al., 2004; San
Mauro et al., 2004a, 2005; Roelants & Bossuyt, 2005; Frost et al., 2006; Gissi et al., 2006; Roelants et al., 2007; Wiens, 2007; Blackburn et al.,
2010). Alytes and Discoglossus have long been recognized as closely related from morphology-based phylogenetic analyses (e.g., Cannatella, 1985;
Gao & Wang, 2001) and molecular phylogenetic analyses agree. Historically, the taxon containing these genera has been referred to as
Discoglossidae. Sanchíz (1998) and Dubois (2005) noted the nomenclatural priority of Alytidae. Bossuyt & Roelants (2009) recognize Alytidae and
Discoglossidae as separate families based on the extent of “evolutionary time” separating these clades, and this remains an appropriate alternative
taxonomy, as would an alternative more inclusive family containing Barbourula and Bombina (and thus the Bombinatoridae, see below). We
tentatively include six extinct taxa in the Alytidae (†Callobatrachus, †Enneabatrachus, †Eodiscoglossus, †Prodiscoglossus, †Scotiophryne,
†Wealdenbatrachus, †Zaphrissa), although their phylogenetic relationships remain unclear (Sanchíz, 1998; Gao & Wang, 2001; Gao & Chen,
2004; Marjanoviæ & Laurin, 2007).
10. Grant et al. (2006) separated a family Aromobatidae (with three subfamilies: Allobatinae, Anomaloglossinae, Aromobatinae) from the
Dendrobatidae. While some have rejected this partitioning (Santos et al., 2009; Santos & Cannatella, 2011; Pyron & Wiens, 2011), many in the
research community (e.g., Verdade & Rodrigues, 2007; Manzanilla et al., 2009; Brown & Twomey, 2009) have found this partitioning useful and
follow the taxonomy of Grant et al. (2006). We view either family-level taxonomy as a viable taxonomy. A supplementary document associated
with Santos et al. (2009) disputes the higher-level taxonomy of Grant et al. (2006), but monophyly of the Dendrobatidae and Aromobatidae and the
proposed subfamilies of Dendrobatidae receive strong support. The basic topology of generic relationships within Aromobatidae recovered by
Santos et al. (2009), as well as by Pyron & Wiens (2011), differs little from that of Grant et al. (2006). Santos et al. (2009) find no support for the
Anomaloglossinae. One composite taxon, which combined molecular data for Allobates alagoanus with morphological data from A. olfersioides,
was resolved as sister to the remaining species of Allobates by Grant et al. (2006). Verdade & Rodrigues (2007) synonymized these two species,
with A. olfersioides having priority. Santos et al. (2009) found this species (referred to in Supplementary Materials as Colosthethus alagoanus) to
be the sister-taxon of all other species of Aromobatidae. This result renders Allobates, and thus Allobatinae, paraphyletic. Pending further analysis
of intrafamilial relationships, we list no subfamilies for the Aromobatidae. Grant et al. (2006) and Santos et al. (2009) provide conflicting views on
the number of genera recognized and allocated to the Aromobatidae and Dendrobatidae (see also Brown et al., 2011).
11.The Arthroleptidae (sensu Frost et al., 2006) has been recognized as a morphologically distinctive lineage for decades (Laurent 1941, 1942, 1951;
see also Dubois, 1981). This includes recognition that Leptopelis may be more closely related to genera in the Arthroleptidae than to those in the
Hyperoliidae, a hypothesis supported by a variety of molecular phylogenetic studies (Emerson et al., 2000; Biju & Bossuyt, 2003; Vences et al.,
2003b; Bossuyt et al., 2006; Frost et al., 2006; Roelants et al., 2007). Frost et al. (2006) recognized two subfamilies: Leptopelinae (Leptopelis) and
Arthroleptinae (Arthroleptis, Astylosternus, Cardioglossa, Leptodactylodon, Nyctibates, Scotobleps, and Trichobatrachus). However, uncertainty
remains in the placement of Leptopelis, including the possibility that the Arthroleptinae sensu Frost et al. (2006) is paraphyletic with respect to
Leptopelis (Vences et al., 2003b; Scott, 2005; Frost et al., 2006; Blackburn, 2008), and thus we do not recognize subfamilies of Arthroleptidae.
12.Frost et al. (2006) recognized Ascaphidae and Leiopelmatidae as subfamilies of Leiopelmatidae. Morphology-based hypotheses of phylogeny have
either separated these two families as successively branching lineages at the base of anuran phylogeny (e.g., Cannatella, 1985) or resolved them as
sister taxa (e.g., Báez & Basso, 1996; Wang et al., 2001). Green et al. (1989) found these taxa to be genetically divergent and suggested that
Ascaphus could be more closely related to other clades of extant anurans than to Leiopelma, though recent molecular phylogenetic analyses have
recognized these as sister taxa (e.g., Frost et al., 2006; Roelants et al., 2007; Irisarri et al., 2010; Pyron & Wiens, 2011). Bossuyt & Roelants (2009)
maintained two families because of the degree of genetic divergence, as did Pyron & Wiens (2011).
13.See footnote 8.
14.Monophyly of Bombinatoridae is supported by both molecular (Blackburn et al., 2010) and morphology-based (Cannatella, 1985; Gao & Wang,
2001) phylogenetic studies. We include †Paradiscoglossus in the Bombinatoridae (Estes & Sanchíz, 1982). See also footnote 9.
15. Hedges et al. (2008) restricted Brachycephalidae to Brachycephalus and Ischnocnema. Brachycephalidae is one of four recognized families in the
unranked taxon Terrarana (Hedges et al., 2008; Heinicke et al., 2009). A viable alternative taxonomy that avoids reliance on unranked taxa would
be to recognize the families of Terrarana as subfamilies of Brachycephalidae.
16.Brevicipitidae (sensu Frost et al., 2006), long included in Microhylidae, is now recognized as a member of a larger clade endemic to sub-Saharan
Africa (e.g., van der Meijden et al., 2004; Frost et al., 2006; Roelants et al., 2007; Wiens, 2007; Kurabayashi et al., 2011). Dubois (2005) enlarged
the Brevicipitidae to also contain the taxa recognized here as Arthroleptidae, Hemisotidae, and Hyperoliidae, which is a viable alternative
taxonomic arrangement that would produce a taxon with nearly 400 species.

Zootaxa 3148 © 2011 Magnolia Press · 41
ANIMAL BIODIVERSITY
Family Ceratobatrachidae Boulenger, 1884 (5 genera; ~86 species)20
Family Ceratophryidae Tschudi, 1838 (3 genera; 12 species; †3)21
Family Ceuthomantidae Heinicke, Duellman, Trueb, Means, MacCulloch & Hedges, 2009 (1 genus; 4 species)22
Family Conrauidae Dubois, 1992 (1 genus; 6 species)23
Family Craugastoridae Hedges, Duellman, & Heinicke, 2008 (2 genera; 115 species)24
Family Cycloramphidae Bonaparte, 1850 (2 genera; 33 species)25
Family Dendrobatidae Cope, 1865 (13 genera; 182 species)26
Subfamily Colostethinae Cope, 1867 (4 genera; 64 species)
Subfamily Dendrobatinae Cope, 1865 (8 genera; 59 species)
17.Generic-level taxonomy within Bufonidae is in a state of flux (Frost et al., 2006, 2009; Pauly et al., 2009). Controversy arises because the many
species historically referred to Bufo do not form an exclusive clade with respect to morphologically distinct satellite taxa (e.g., Ansonia,
Capensibufo, Nectophrynoides, Pedostibes, Schismaderma, Stephopaedes). In the interests of taxonomic stability, some workers prefer to maintain
Bufo for readily recognized “toad”-like taxa, even to the point of reducing morphologically divergent taxa long recognized as genera to subgenera.
This would result in a very large genus Bufo, with more than 450 species. Alternatively, approximately 50 genera would be recognized (Frost et al.,
2006; Frost, 2011), with many species long placed in Bufo and having extensive literature references being placed in newly (or recently) created
genera. Many of the generic-level taxonomic changes have been embraced already by much of the community of amphibian taxonomists (Pramuk
et al., 2007; Van Bocxlaer et al., 2009, 2010; Maciel et al., 2010). Pyron & Wiens (2011) recognize 35 genera.
18. Calyptocephalellidae is recovered as monophyletic and a sister taxon of our Myobatrachidae (San Mauro et al., 2005; Wiens et al., 2005; Correa et
al., 2006; Frost et al., 2006; Pyron & Wiens, 2011). Frost et al. (2006) referred to the clade containing Calyptocephalella and Telmatobufo as the
Batrachophrynidae because it was not yet clear that Batrachophrynus is likely embedded within the genus Telmatobius (Aguilar & Pacheco, 2005;
Córdova & Descailleaux, 2005; Aguilar & Valencia, 2009).
19. Guayasamin et al. (2009) recognized two subfamilies within the Centrolenidae. We follow Pyron & Wiens (2011) in placing the monotypic Ikakogi
in the Centroleninae. See also footnote 7.
20.Multiple phylogenetic studies have revealed complicated relationships among clades variously assigned to the Ranidae (see Dubois, 1981, 1983,
1992, 2005; Duellman & Trueb, 1986), including clades recognized here as the Arthroleptidae, Mantellidae, and Rhacophoridae (Emerson et al.,
2000; Vences et al., 2003bc; Roelants et al., 2004, 2007; Scott, 2005; van der Meijden et al., 2005; Bossuyt et al., 2006; Frost et al., 2006; Wiens,
2007; Wiens et al., 2009; Pyron & Wiens, 2011; see also Ford, 1990; Ruvinsky & Maxson, 1996; Biju & Bossuyt, 2003; Haas, 2003). Dubois
(2005) suggested recognizing fourteen subfamilies of Ranidae (Ceratobatrachinae, Conrauinae, Dicroglossinae, Lankanectinae, Mantellinae,
Micrixalinae, Nyctibatrachinae, Petropedetinae, Phrynobatrachinae, Ptychadeninae, Pyxicephalinae, Raninae, Ranixalinae, and Rhacophorinae);
for a summary of the taxonomic history of “ranid” frogs, see Frost et al. (2006). The subfamilial taxa of Dubois (2005) were elevated to the family
level by Frost et al. (2006), although several were combined into single families; Lankanectinae and Nyctibatrachinae were combined into the
Nyctibatrachidae, and Conrauinae, Petropedetinae, and Ranixalinae were combined into the Petropedetidae (although Ranixalidae is recognized as
a separate family by subsequent authors; Van Bocxlaer et al., 2006; Bossuyt & Roelants, 2009; Wiens et al., 2009). Some authors (Bossuyt et al.,
2006; Wiens et al., 2009) follow the concept of Ranidae advocated by Dubois (2005), but there is growing use of the family-level taxonomy
advocated by Frost et al. (2006). As pointed out by Frost et al. (2006), their unranked taxon Natatanura roughly corresponds to what previous
workers have referred to as “ranids”. There is high support for the Natatanura of Frost et al. (2006) from a variety of phylogenetic studies, although
the relationships among these families remain unresolved (Vences et al., 2003bc; Roelants et al., 2004, 2007; Scott, 2005; Bossuyt et al., 2006;
Wiens, 2007; Wiens et al., 2009; Pyron & Wiens, 2011). Ceratobatrachidae is supported as monophyletic (Bossuyt et al., 2006; Frost et al., 2006;
Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens et al., 2009; Ruane et al., 2011; Pyron & Wiens, 2011; see also Brown, 2004).
21.Based on recent phylogenetic studies (Evans et al., 2008; Ruane et al., 2011), we include three extinct genera in the Ceratophryidae
(†Baurubatrachus, †Beelzebufo, †Wawelia). See also footnote 8.
22.Heinicke et al. (2009) described the Ceuthomantidae, as well as its sole genus Ceuthomantis, and showed that it is sister to a clade containing other
families placed in the unranked taxon Terrarana.
23.The monophyly of a clade containing Conraua and Petropedetes (including taxa formerly placed in Arthroleptides) is supported in several
phylogenetic analyses (Bossuyt et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens et al., 2009; Ruane et al., 2011), but not others
(van der Meijden et al., 2005; Zimkus et al., 2010; Pyron & Wiens, 2011). Frost et al. (2006) resolved this clade to also contain the taxon here
recognized as Ranixalidae, but this is not supported by subsequent analyses. Dubois (1992) considered the Conrauini to be a tribe within his
subfamily Dicroglossinae (Ranidae of Dubois, 1992) and, by implication, included Petropedetes within the Phrynobatrachidae; Dubois (2005) later
treated both Conrauinae and Petropedetinae as subfamilies of the Ranidae (sensu Dubois, 2005). In light of the uncertain sister relationship between
these two clades and their likely deep divergence (Roelants et al., 2007), we recognize these as two distinct families, Conrauidae and
Petropedetidae. For further details on taxonomy of taxa placed in the Ranidae of Dubois (2005), see footnote 20.
24. Hedges et al. (2008) proposed the Craugastoridae for the diverse Craugastor and its sister taxon, a new genus Haddadus. The Craugastoridae is one
of four families in the unranked taxon Terrarana (Hedges et al., 2008; Heinicke et al., 2009; Padial et al., 2009). Pyron & Wiens (2011) found
Craugastoridae sensu Hedges et al. (2009) to be embedded within the Strabomantidae sensu Hedges et al. (2009), and expanded the Craugastoridae
to include all taxa previously assigned to Strabomantidae. However, because of low support values among basal nodes in this larger clade, the
analysis of Pyron & Wiens (2011) does not reject the hypothesis that Craugastoridae is sister to the Strabomantidae. Higher-level relationships
among these clades require further study.
25.See footnote 8.
26. Grant et al. (2006) recognize three subfamilies of Dendrobatidae (for discussion regarding taxa in the Aromobatidae, see footnote 10). The
phylogenetic relationships resolved in other studies (Vences et al., 2000, 2003a; Roberts et al., 2006; Santos et al., 2009; Santos & Cannatella,
2011) support the tree topology on which the subfamily taxonomy of Grant et al. (2006) is based. Santos et al. (2009; see also Santos & Cannatella,
2011) argued that the partitioning of Dendrobates into six genera was unnecessary (Adelphobates, Dendrobates, Excidobates, Minyobates,
Oophaga, Ranitomeya). Brown et al. (2011) discuss this matter at length and elect to recognize all six genera and describe a seventh genus,
Andinobates.

BLACKBURN & WAKE: CLASS AMPHIBIA GRAY, 1825
42 · Zootaxa 3148 © 2011 Magnolia Press
Subfamily Hyloxalinae Grant, Frost, Caldwell, Gagliardo, Haddad, Kok, Means, Noonan, Schargel &
Wheeler, 2006 (1 genus; 59 species)
Family Dicroglossidae Anderson, 1871 (12–14 genera; 177 species)27
Subfamily Dicroglossinae Anderson, 1871 (10–12 genera; 155 species)
Subfamily Occidozyginae Fei, Ye, & Huang, 1990 (2 genera; 22 species)
Family Eleutherodactylidae Lutz, 1954 (4 genera; 202 species)28
Subfamily Eleutherodactylinae Lutz, 1954 (2 genera; 195 species)
Subfamily Phyzelaphryninae Hedges, Duellman, & Heinicke, 2008 (2 genera; 7 species)
Family †Gobiatidae Ro ek & Nesov, 1993 (†3)29
Family Heleophrynidae Noble, 1931 (2 genera; 6–7 species)30
Family Hemiphractidae Peters, 1862 (6 genera; 95 species)31
Family Hemisotidae Cope, 1867 (1 genus; 9 species)32
Family Hylidae Rafinesque, 1815 (42–46 genera; 904 species; †1)33
Subfamily Hylinae Rafinesque, 1815 (36–39 genera 647 species;)
Subfamily Pelodryadinae Günther, 1858 (1–2 genera; 198 species)
Subfamily Phyllomedusinae Günther, 1858 (5 genera; 59 species)
Family Hylodidae Günther, 1858 (3 genera; 42 species)34
Family Hyperoliidae Laurent, 1943 (18 genera; 214 species)35
Family Leiopelmatidae Mivart, 1869 (1 genus; 4 species)36
Family Leiuperidae Bonaparte, 1850 (7 genera; 85 species)37
Family Leptodactylidae Werner, 1896 (6 genera; 105 species)38
27. Phylogenetic analyses of molecular data support monophyly of the Dicroglossidae (Kosuch et al., 2001; Roelants et al., 2004, 2007; Kurabayashi et
al., 2005; van der Meijden et al., 2005; Bossuyt et al., 2006; Frost et al., 2006; Van Bocxlaer et al., 2006; Che et al., 2007; Wiens, 2007; Wiens et
al., 2009; Ruane et al., 2011). Two clades, the subfamilies Dicroglossinae and Occidozyginae, also receive strong support (Kosuch et al., 2001;
Roelants et al., 2004; Bossuyt et al., 2006; Frost et al., 2006; Che et al., 2007; Wiens et al., 2009; Ruane et al., 2011; Pyron & Wiens, 2011).
Generic-level taxonomy and phylogenetic relationships within the Dicroglossinae remain unresolved (Dubois et al., 2001; Jiang et al., 2005; Che et
al., 2007, 2009, 2010; Pyron & Wiens, 2011). For further details on taxonomy of taxa placed in the Ranidae of Dubois (2005), see footnote 20.
28. Hedges et al. (2008) recognized two well-supported clades within the Eleutherodactylidae, one including Diasporus and the mega-diverse genus
Eleutherodactylus and the other including the species-poor Adelophryne and the monotypic Phyzelaphryne.
29.The †Gobiatidae is an extinct family of three genera (†Cretasalia, †Gobiates, and †Gobiatoides) with obscure relationships to other families of
Anura (Ro ek, 2008). Sanchíz (1998) questioned the validity of †Gobiatoides.
30.Molecular phylogenies have resolved the Heleophrynidae as the sister taxon of all other Neobatrachia (e.g., Hoegg et al., 2004; Frost et al., 2006;
Roelants et al., 2007; Wiens, 2007). Van Dijk (2008) erected Hadromophryne, the validity of which is further supported by Pyron & Wiens (2011).
31. Monophyly of the Hemiphractidae has been controversial. Based on molecular phylogenetic analyses, some authors have found the Hemiphractidae
to be paraphyletic (Darst & Cannatella, 2004), or polyphyletic (Faivovich et al., 2005; Frost et al., 2006), whereas others resolved it to be
monophyletic (Wiens, 2007; Guayasamin et al., 2008; Heinicke et al., 2009; Pyron & Wiens, 2011). Frost et al. (2006) recognized three families
(Amphignathodontidae, Cryptobatrachidae, Hemiphractidae). Because more recent studies have resolved this family as monophyletic, we maintain
the Hemiphractidae for Cryptobatrachus, Flectonotus, Gastrotheca, Hemiphractus, and Stefania, as well as the resurrected Fritziana (Duellman et
al., 2011).
32. The family Hemisotidae, containing only the genus Hemisus, is assumed to be monophyletic, though we know of no explicit test. Hemisotidae is the
sister taxon of the Breviciptidae (Frost et al., 2006; Roelants et al., 2007; van der Meijden et al., 2007b; Wiens, 2007), a relationship previously
suggested by morphological phylogenetic studies (Blommers-Schlösser, 1993; Channing, 1995). One viable alternative taxonomic arrangement
would be to include Hemisotidae as a subfamily of the Brevicipitidae.
33.The content of the Hylidae has been extensively modified in recent years. Those frogs recognized above as the Hemiphractidae were previously
considered part of the Hylidae (Duellman, 1970), but they are not necessarily closely related (Darst & Cannatella, 2004; Frost et al., 2006; Wiens,
2007; Heinicke et al., 2009). Most analyses agree in resolving a monophyletic lineage containing three subfamilies (Darst & Cannatella, 2004;
Faivovich et al., 2005; Wiens et al., 2005; Frost et al., 2006; Pyron & Wiens, 2011), two of which (Pelodryadinae and Phyllomedusinae) form a
clade sister to the third (Hylinae). However, there is some disagreement on the monophyly of Hylidae (Roelants et al., 2007), which, combined with
deep genetic divergences, has led some authors to recognize each subfamily as a distinct family (Bossuyt & Roelants, 2009). We recognize a single
family with three subfamilies. Faivovich et al. (2005) extensively revised the generic-level taxonomy of Hylinae. Generic-level taxonomy of the
Pelodryadinae remains in flux with authors recognizing one or two genera (Faivovich et al., 2005; Frost et al., 2006; Rosauer et al., 2009; Tyler et
al., 2009; Pyron & Wiens, 2011). For the Phyllomedusinae, Faivovich et al. (2010) synonymized Hylomantis and Pachymedusa with Agalychnis.
The relationships of †Australobatrachus to currently recognized clades remains unclear.
34.The three genera of Hylodidae form a clade (Nuin & do Val, 2005; Frost et al., 2006; Grant et al. 2006; see also Pyron & Wiens, 2011) that Grant
et al. (2006) resolved as the sister taxon of the Dendrobatidae + Aromobatidae (also suggested by morphological and karyological data; e.g., Lynch,
1971; Bogart, 1991; Augiar et al., 2004) and thus removed it from the Cyclorhamphidae, where it had been placed by Frost et al. (2006). Pyron &
Wiens (2011) resolved Hylodidae as the sister taxon of Alsodidae, and not near the Dendrobatidae + Aromobatidae.
35. The Hyperoliidae is a diverse clade of eighteen genera, many of which have long been recognized as sharing morphological features (Laurent 1986;
Drewes, 1984) and both morphological (Drewes, 1984) and molecular phylogenetic analyses support monophyly (Vences et al., 2003b; Frost et al.,
2006; Veith et al., 2009). As detailed in footnote 11, the genus Leptopelis is now considered part of the Arthroleptidae, which is the sister taxon to
Hyperoliidae.
36.See footnote 12.

Zootaxa 3148 © 2011 Magnolia Press · 43
ANIMAL BIODIVERSITY
Family Mantellidae Laurent, 1946 (12 genera; 198 species)39
Subfamily Boophinae Vences & Glaw, 2001 (1 genus; 72 species)
Subfamily Laliostominae Vences & Glaw, 2001 (2 genera; 4 species)
Subfamily Mantellinae Laurent, 1946 (9 genera; 122 species)
Family Megophryidae Bonaparte, 1850 (10 genera; 160 species)40
Family Micrixilidae Dubois, Ohler, & Biju, 2001 (1 genus; 11 species)41
Family Microhylidae Günther, 1858 (68 genera; 495 species)42
Subfamily Asterophryinae Günther, 1858 (22 genera; 252 species)
Subfamily Cophylinae Cope, 1889 (7 genera; 58 species)
Subfamily Dyscophinae Boulenger, 1882 (1 genus; 3 species)
Subfamily Gastrophryninae Fitzinger, 1843 (13 genera; 59 species)
Subfamily Hoplophryninae Noble, 1931 (2 genera; 3 species)
Subfamily Kalophryninae Mivart, 1869 (1 genus; 17 species)
Subfamily Melanobatrachinae Noble, 1931 (1 genus; 1 species)
Subfamily Microhylinae Günther, 1858 (9 genera; 71 species)
Subfamily Otophryninae Wassersug & Pyburn, 1987 (2 genera; 6 species)
Subfamily Phrynomerinae Noble, 1931 (1 genus; 5 species)
Subfamily Scaphiophryninae Laurent, 1946 (2 genera; 12 species)
Family Myobatrachidae Schlegel, 1850 (20 genera; 127 species; †1)43
Subfamily Limnodynastinae Lynch, 1969 (8 genera; 44 species)
Subfamily Myobatrachinae Schlegel, 1850 (12 genera; 83 species; †1)
Family Nasikabatrachidae Biju & Bossuyt, 2003 (1 genus; 1 species)44
Family Nyctibatrachidae Blommers-Schlösser, 1993 (2 genera; 29 species)45
Family Odontophrynidae Lynch, 1969 (3 genera; 36 species)46
Family †Palaeobatrachidae Špinar, 1972 (†4)47
37. Grant et al. (2006) recognized Leiuperidae for a clade of seven genera that Frost et al. (2006) included within the Leptodactylidae. Frost et al.
(2006) did not resolve the Leiuperidae as monophyletic but both Correa et al. (2006) and Grant et al. (2006) did. Leiuperidae is treated as a
subfamily of Leptodactylidae by Pyron & Wiens (2011).
38. Ruvinsky & Maxson (1996) showed that Leptodactylidae sensu Lynch (1971, 1973) is polyphyletic (see also Darst & Cannatella (2004), Faivovich
et al. (2005), Wiens et al. (2005), and Carrera et al. (2006). Frost et al. (2006) partitioned Leptodactylidae into multiple families; Grant et al. (2006)
modified this scheme by further partitioning (see also Pyron & Wiens, 2011). Based in part on previous work of Heyer (1998) and Kokobum &
Giaretta (2005) and finding that Adenomera and Lithodytes form a clade sister to Leptodactylus, Frost et al. (2006) considered these genera to be
synonyms of Leptodactylus. However, subsequent authors have provided evidence that these genera are morphologically distinct (Ponssa & Heyer,
2007), and both molecular (Pyron & Wiens, 2011) and morphology-based (de Sá et al., 2005; Ponssa et al., 2010) phylogenetic analyses suggest
that Adenomera and Lithodytes are not nested within Leptodactylus.
39.Phylogenetic analyses provide strong support for the monophyly of the Mantellidae (Emerson et al., 2000; Vences et al., 2003bc; Roelants et al.,
2004, 2007; Bossuyt et al., 2006; Frost et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens, 2007; Vieites et al., 2009; Wiens et al.,
2009; Ruane et al., 2011). The subfamilies (Boophinae, Laliostominae, and Mantellinae) are also strongly supported (Vences et al., 2003c; Roelants
et al., 2004; Bossuyt et al., 2006; Frost et al., 2006; Glaw et al., 2006; Kurabayashi et al., 2008; Vieites et al., 2009; Wiens et al., 2009; Pyron &
Wiens, 2011). Frost et al. (2006) included the Laliostominae in the Mantellinae, but the taxonomic community has not accepted this proposal (e.g.,
Glaw & Vences, 2006; Glaw et al., 2006; Hiobiarilanto et al., 2010) and these clades may not be sister to one another (Pyron & Wiens, 2011). The
relationship of the monotypic Tsingymantis to the recognized subfamilies remains a point for future research (Glaw et al., 2006; Kurabayashi et al.,
2008). For further details on taxonomy of taxa placed in the Ranidae of Dubois (2005), see footnote 20.
40.The monophyly of the Megophryidae is widely accepted (e.g., Ford & Cannatella, 1993; Frost et al., 2006). The most significant recent taxonomic
change is the recognition that Leptobrachium is paraphyletic with respect to Vibrissaphora (Rao & Wilkinson, 2008; Brown et al., 2009). We
refrain from recognizing subfamilies within Megophryidae (Leptobrachiinae, Leptolalaginae, Megophryinae, as advocated by Delorme et al., 2006)
until phylogenetic analyses with more inclusive taxon sampling are available. However, the molecular phylogenetic analysis by Pyron & Wiens
(2011) generally supports the subfamilies of Delorme et al. (2006), but lacks sampling for Leptobatrachella; their analysis also suggests that
Xenophrys may be paraphyletic with respect to both Megophrys and Brachytarsophrys.
41.The monophyly of the Micrixalidae, containing only the genus Micrixalus, is supported by phylogenetic analyses (Bossuyt et al., 2006; Van
Bocxlaer et al., 2006; Frost et al., 2006; Wiens et al., 2009; see also Dubois et al., 2001). For further details on taxonomy of taxa placed in the
Ranidae of Dubois (2005), see footnote 20.
42.Following recognition of the Brevicipitidae (see footnote 16), there is strong support for the monophyly of the Microhylidae (Ford & Cannatella,
1993; Haas, 2003; van der Meijden et al., 2004, 2007b; Frost et al., 2006; Roelants et al., 2007; Kurabayashi et al., 2011). Based in part on the
substantial divergences between subfamilies observed in DNA sequence data, Bossuyt & Roelants (2009) elevated each subfamily to family rank.
However, Bossuyt & Roelants (2009) did not mention the Otophryninae, leaving its status uncertain, and failed to provide any details on the many
taxa not assigned to subfamilies within the classification scheme of Frost et al. (2006). At this point, adopting a strategy in which each subfamily of
Microhylidae is elevated to family would lead to many genera of unclear affinities being orphaned. Thus, we advocate recognizing the family
Microhylidae with multiple subfamilies and leaving certain genera without subfamily designation. Based on Greenbaum (2006), we suggest that
Altigius, Melanophryne, Myersiella, and Syncope should be included in the Gastrophryninae. Following Pyron & Wiens (2011) and Trueb et al.
(2011), we tentatively include Synapturanus in the Otophryninae.

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44 · Zootaxa 3148 © 2011 Magnolia Press
Family Pelobatidae Bonaparte, 1850 (1 genus; 4 species)48
Family Pelodytidae Bonaparte, 1850 (1 genus; 3 species; †2)49
Family Petropedetidae Noble, 1931 (1 genus; 12 species)50
Family Phrynobatrachidae Laurent, 1941 (1 genus; 84 species)51
Family Pipidae Gray, 1825 (4–5 genera; 33 species; †7)52
Family Ptychadenidae Dubois, 1987 (3 genera; 51 species)53
Family Pyxicephalidae Bonaparte, 1850 (13 genera; 67 species)54
Subfamily Cacosterninae Noble, 1931 (11 genera; 61 species)
Subfamily Pyxicephalinae Bonaparte, 1850 (2 genera; 6 species)
Family Ranidae Rafinesque, 1814 (10–16 genera; 369 species)55
Family Ranixalidae Dubois, 1987 (1 genus; 10 species)56
Family Rhacophoridae Hoffman, 1932 (15 genera; 333 species)57
Subfamily Buergeriinae Channing, 1989 (1 genus; 5 species)
43.From one to three families of myobatrachids (Myobatrachidae, Limnodynastidae, Rheobatrachidae) have been recognized. Ford & Cannatella
(1993) questioned the monophyly of a single family (e.g., Heyer & Liem, 1976) and proposed that †Rheobatrachus may be most closely related to
the Myobatrachinae (sensu Heyer & Liem, 1976). Several molecular phylogenetic analyses did not test the monophyly of each family (e.g., Read et
al., 2001; Morgan et al., 2007). Frost et al. (2006) found that twenty genera variously assigned to the three families form a clade, yet chose to
recognize two families (Limnodynastidae and Myobatrachidae, the latter containing Mixophyes and †Rheobatrachus). Much of the uncertainty of
higher-level taxonomy in this clade relates to Mixophyes and †Rheobatrachus, the relationships of which remain uncertain (Heyer & Liem, 1976;
Farris et al., 1982; Frost et al., 2006; Roelants et al., 2007; Ruane et al., 2011; Pyron & Wiens, 2011). In several analyses, Mixophyes and
†Rheobatrachus are not sister taxa and the relationships of these two genera to the other taxa in the Limnodynastidae and Myobatrachidae (sensu
Frost et al., 2006) remain unclear (Roelants et al., 2007; Pyron & Wiens, 2011; Ruane et al., 2011). Bossuyt & Roelants (2009) recognized
Rheobatrachidae, Limnodynastidae, and Myobatrachidae but mentioned neither †Rheobatrachus nor Mixophyes explicitly. To date, the most
complete relevant analyses are those of Frost et al. (2006) and Pyron & Wiens (2011), which largely agree with one another, yet disagree on
whether Rheobatrachus and Mixophyes form a clade. Because of remaining uncertainties in the placement of †Rheobatrachus and Mixophyes, we
follow Pyron & Wiens (2011) by using two subfamilies for the single family Myobatrachidae. The genus †Rheobatrachus became extinct in the late
20th century.
44.The monotypic family Nasikabatrachidae (Biju & Bossuyt, 2003) is nearly universally accepted, although Frost et al. (2006) included the sole
genus in the Sooglossidae. We follow the generally accepted practice of maintaining these as two distinct families (e.g., Frost, 2011; Pyron &
Wiens, 2011).
45.Frost et al. (2006) subsumed the Lankanectinae and Nyctibatrachinae of Dubois (2005) into a single family, Nyctibatrachidae. The monophyly of
this family, containing the two genera Lankanectes and Nyctibatrachus, receives high support from phylogenetic analyses (van der Meijden et al.,
2005; Bossuyt et al., 2006; Frost et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens, 2007; Wiens et al., 2009; Ruane et al., 2011;
Pyron & Wiens, 2011; see also Dubois & Ohler, 2001). For further details on taxonomy of taxa placed in the Ranidae of Dubois (2005), see
footnote 20.
46.Pyron & Wiens (2011) found strong support for the monophyly of a clade of three genera (Macrogenioglottus, Odontophrynus, and
Proceratophrys), which they recognized as the Odontophrynidae. See also footnote 8.
47. †Palaeobatrachidae contains four recognized extinct genera (†Albionbatrachus, †Messelobatrachus, †Palaeobatrachus, and †Pliobatrachus) and is
generally allied to the extant Pipidae although the phylogenetic relationships of this family remain obscure (Sanchíz, 1998). The phylogenetic
analyses of Gao & Wang (2001) and Gao & Chen (2004), which included †Palaeobatrachus, suggest that †Palaeobatrachidae may nest within the
Pipidae, but a phylogenetic analysis with greater taxonomic sampling of extinct pipoid taxa supports the monophyly of Pipidae to the exclusion of
†Palaeobatrachus
48.The well-supported Pelobatidae contains a single genus (Pelobates) with four species (Cannatella, 1985; Ford & Cannatella, 1993; Lathrop, 1997;
García-París et al., 2003; Roelants & Bossuyt, 2005; Frost et al., 2006; Veith et al., 2006). Based in part on the morphological phylogenetic analysis
of Cannatella (1985), Ford & Cannatella (1993) defined Pelobatidae to include Pelobates as well as Scaphiopus and Spea (see also, e.g., Noble,
1925). Analyses of morphological data by Lathrop (1997), Henrici & Haynes (2006), and Henrici (2009) further support this result as well as
including the extinct taxa †Elkobatrachus, †Macropelobates, and †Eopelobates in the Pelobatidae. However, subsequent molecular phylogenetic
analyses, as well as a combined analysis of larval and adult morphology (Pugener et al., 2003) indicate that Pelobates, Scaphiopus, and Spea do not
form a clade exclusive of Pelodytidae and Megophryidae (see also footnote 39). We follow the results of these recent studies by recognizing
Pelobatidae and Scaphiopodidae to be distinct families, although the relationships of the extinct taxa now remain uncertain and we consider these
Anura incertae sedis.
49.The monophyly of the Pelodytidae, which contains a single genus (Pelodytes) with three species, is supported by phylogenetic analysis of
mitochondrial DNA sequence data (García-París et al., 2003). Analyses of both morphological and molecular data support the Pelodytidae as a
member of a larger clade containing the Pelobatidae, Scaphiopodidae, and Megophryidae (Cannatella, 1985; Ford & Cannatella, 1993; Hay et al.,
1995; García-París et al., 2003; Pugener et al., 2003; Roelants & Bossuyt, 2005; San Mauro et al., 2005; Frost et al., 2006; Veith et al., 2006;
Roelants et al., 2007; Wiens, 2007; Ruane et al., 2011). †Miopelodytes and †Tephrodytes have been included in the Pelodytidae based on the fusion
of the tibiale and fibulare (unique among archaeobatrachians; Cannatella, 1985), but the precise relationships of these two genera to Pelodytes
remains unclear (Taylor, 1941; Henrici, 1994).
50.See footnote 23.
51. The monophyly of the Phrynobatrachidae, containing the single diverse genus Phrynobatrachus, is well supported by phylogenetic analyses (Scott,
2005; Bossuyt et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens et al., 2009; Ruane et al., 2011; Zimkus et al., 2010). While
combined molecular and morphological data of Scott (2005) suggested paraphyly of Phrynobatrachus with respect to Natalobatrachus, the recent
phylogenetic analysis of van der Meijden et al. (2011) demonstrate Natalobatrachus to be within the Pyxicephalidae. For further details on
taxonomy of taxa placed in the Ranidae of Dubois (2005), see footnote 20.

Zootaxa 3148 © 2011 Magnolia Press · 45
ANIMAL BIODIVERSITY
Subfamily Rhacophorinae Hoffman, 1932 (14 genera; 327 species)
Family Rhinodermatidae Günther, 1858 (2 genera; 3 species)58
Family Rhinophrynidae Günther, 1859 (1 genus; 1 species; †2–3)59
Family Scaphiopodidae Cope, 1865 (2 genera; 7 species)60
Family Sooglossidae Noble, 1931 (2 genera; 4 species)61
Family Strabomantidae Hedges, Duellman, & Heinicke, 2008 (17–19 genera; 572 species)62
Subfamily Holoadeninae Hedges, Duellman, & Heinicke, 2008 (6 genera; 47 species)
Subfamily Strabomantinae Hedges, Duellman, & Heinicke, 2008 (11–13 genera; 525 species)
Family Telmatobiidae Fitzinger, 1843 (2 genera; 60 species)63
Order Caudata Fischer von Waldheim, 1813 (salamanders) (67–68 genera; 614 species; †~66)64
52. Phylogenetic analyses of morphological and molecular data support the monophyly of the Pipidae (Cannatella, 1985; Cannatella & Trueb, 1988ab;
Ford & Cannatella, 1993; Haas, 2003; Pugener et al., 2003; Evans et al., 2004; Roelants & Bossuyt, 2005; San Mauro et al., 2005; Frost et al.,
et al., 2007; Wiens, 2007; Irisarri et al., 2011; Ruane et al., 2011). The
relationships of Hymenochirus and Pseudhymenochirus are unclear; some studies find these genera (typically represented only by Hymenochirus)
to be more closely related to extant Xenopus (including Silurana; Roelants & Bossuyt, 2005; San Mauro et al., 2005; Roelants et al., 2007; Wiens,
2007; Irisarri et al., 2011; Ruane et al., 2011), or to Pipa (Cannatella, 1985; Cannatella & Trueb, 1988ab; Evans et al., 2004; Pugener et al., 2003;
Báez & Harrison, 2005; Trueb et al., 2005; Marjanovi & Laurin, 2007), or possibly sister to a clade containing both Xenopus and Pipa (Frost et al.,
2006); see also de Sá & Hillis (1990). Because of these uncertainties, we refrain from recognizing subfamilies within the Pipidae. Cannatella &
Trueb (1988a) recognized the genus Silurana as distinct from Xenopus, although one of these authors states that this was an unsound decision
(Pauly et al., 2009). Diverse extinct taxa of the Pipidae include †Eoxenopoides, †Llankibatrachus, †Pachycentrata (Pipinae; Báez & Harrison,
2005; Trueb & Báez, 2006), †Saltenia, †Shelania, †Singidella (Pipinae; Báez & Harrison, 2005), and †Vulcanobatrachus. Other extinct taxa that
may be more closely related to Pipidae than to Rhinophrynidae include †Avitabatrachus, †Cordicephalus, and †Thoraciliacus, and possibly the
Palaeobatrachidae (Báez et al., 2000; Trueb et al., 2005; Trueb & Báez, 2006), but we do not include these taxa within the Pipidae.
53.The monophyly of the Ptychadenidae receives strong support from phylogenetic analyses (Scott, 2005; Bossuyt et al., 2006; van Bocxlaer et al.,
2006; Wiens et al., 2009). For further details on taxonomy of taxa placed in the Ranidae of Dubois (2005), see footnote 20.
54.Phylogenetic analyses of molecular data provide strong support for the monophly of the African endemic family Pyxicephalidae (van der Meijden
et al., 2005, 2011; Bossuyt et al., 2006; Frost et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens, 2007; Wiens et al., 2009; Zimkus
et al., 2010; Pyron & Wiens, 2011) and its component subfamilies Cacosterninae and Pyxicephalinae (Scott, 2005; van der Meijden et al., 2005,
2011; Bossuyt et al., 2006; Frost et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens et al., 2009; Zimkus et al., 2010; Pyron &
Wiens, 2011). For further details on taxonomy of taxa placed in the Ranidae of Dubois (2005), see footnote 20.
55. Phylogenetic analysies of molecular data provide strong support for the monophly of the African endemic family Pyxicephalidae (van der Meijden
et al., 2005, 2011; Bossuyt et al., 2006; Frost et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens, 2007; Wiens et al., 2009; Zimkus
et al., 2010; Pyron & Wiens, 2011) and its component subfamilies Cacosterninae and Pyxicephalinae (Scott, 2005; van der Meijden et al., 2005,
2011; Bossuyt et al., 2006; Frost et al., 2006; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens et al., 2009; Zimkus et al., 2010; Pyron &
Wiens, 2011). For further details on taxonomy of taxa placed in the Ranidae of Dubois (2005), see footnote 20.
56. The Ranixalidae, containing only the genus Indirana, a well supported clade (Roelants et al., 2004, 2007; Bossuyt et al., 2006; Van Bocxlaer et al.,
2006; Wiens et al., 2009; Ruane et al., 2011; Pyron & Wiens, 2011). For further details on taxonomy of taxa placed in the Ranidae of Dubois
(2005), see footnote 20.
57.Rhacophoridae and its two subfamilies are clades (Emerson et al., 2000; Haas, 2003; Kurabayashi et al., 2005; Scott, 2005; van der Meijden et al.,
2005; Bossuyt et al., 2006; Frost et al., 2006; Vences et al., 2003bc; Van Bocxlaer et al., 2006; Roelants et al., 2007; Wiens, 2007; Li et al., 2009;
Wiens et al., 2009; Yu et al., 2009; Ruane et al., 2011; Pyron & Wiens, 2011). Phylogenetic analyses led to the recognition of several additional
rhacophorine genera (Biju et al., 2010; Meegaskumbura et al., 2010). For further details on taxa placed in the Ranidae by Dubois (2005), see
footnote 20.
58.See footnote 8.
59. The Rhinophrynidae, represented today only by the monotypic Rhinophrynus, is the sister taxon of the extant Pipidae (Cannatella, 1985; Hay et al.,
1995; Haas, 2003; Pugener et al., 2003; Roelants & Bossuyt, 2005; Frost et al., 2006; Roelants et al., 2007; Wiens, 2007; Ruane et al., 2011). The
phylogenetic analyses of Gao & Wang (2001) and Gao & Chen (2004) support a sister relationship of Rhinophrynus with †Palaeobatrachus, but
analyses with greater sampling of extinct pipoid taxa do not support this relationship (Trueb et al., 2005; Trueb & Báez, 2006; Marjanovi &
Laurin, 2007). We follow Henrici (1998) by including the fossil taxa †Chelomophrynus and †Rhadinosteus, and possibly †Eorhinophrynus, in the
Rhinophrynidae.
60.The genera Scaphiopus and Spea form the monophyletic Scaphiopodidae (Cannatella, 1985; Lathrop, 1997; García-París et al., 2003; Roelants &
Bossuyt, 2005; Frost et al., 2006; Roelants et al., 2007; Wiens, 2007; Ruane et al., 2011), which Ford & Cannatella (1993) included in the
Pelobatidae. While the phylogenetic analyses of morphological data by Henrici & Haynes (2006) and Henrici (2009) did not resolve Scaphiopus
and Spea as sister taxa to the exclusion of Pelobates, other phylogenetic analyses present evidence that these two genera form a clade. The
phylogenetic relationships and family-level taxonomy of fossil taxa previously allied to Pelobates, Scaphiopus, and Spea remain unclear (Zweifel,
1956; Henrici & Haynes, 2006; Henrici, 2009) and we thus leave these unassigned to family. See also footnote 48.
61. Sooglossidae is a well supported clade (Nussbaum, 1982; Ford & Cannatella, 1993; Nussbaum & Wu, 2007; Frost et al., 2006; van der Meijden et
al., 2007a; Pyron & Wiens, 2011). Partitioning of the diversity of this family into genera is supported by morphological, molecular, and acoustic
data (Nussbaum & Wu, 2007; van der Meijden et al., 2007a; see also Nussbaum et al., 1982).
62. Hedges et al. (2008) proposed the Strabomantidae and its two subfamilies (Holoadeninae and Strabomantinae) for a diverse clade of Central and
South American genera. Strabomantidae is one of four families in the unranked taxon Terrarana (Hedges et al., 2008; Heinicke et al., 2009).
Subsequent molecular phylogenetic analyses with less taxon sampling have not resolved the two subfamilies as monophyletic (Heinicke et al.,
2007, 2009), or even as forming a clade (Padial et al., 2009), but because these are based on substantially less sampling of taxa and/or genetic loci,
we follow the subfamily taxonomy proposed by Hedges et al. (2008).See also footnote 24.
63.See footnote 8.

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46 · Zootaxa 3148 © 2011 Magnolia Press
Family Ambystomatidae Gray, 1850 (1 genus; 32 species; †2)65
Family Amphiumidae Gray, 1825 (1 genus; 3 species; †1)66
Family †Batrachosauroididae Auffenberg, 1958 (†7)67
Family Cryptobranchidae Fitzinger, 1826 (2 genera; 3 species; †6)68
Family Dicamptodontidae Tihen, 1958 (1 genus; 4 species; †5)69
Family Hynobiidae Cope, 1859 (9 genera; 53 species; †1)70
Family †Karauridae Ivachnenko, 1978 (†2–3)71
Family Plethodontidae Gray, 1850 (27 genera; 418 species)72
Subfamily Hemidactyliinae Hallowell, 1856 (20 genera; 322 species)
Subfamily Plethodontidinae Gray, 1850 (7 genera; 96 species)
Family Proteidae Gray, 1825 (2 genera; 6 species; †2)73
Subfamily Proteinae Gray, 1825 (1 genus; 1 species; †2)
64.We consider Caudata and Urodela to be synonyms and follow the argumentation of Frost et al. (2006) by recognizing Caudata as the appropriate
name (for a contrasting view, see Dubois, 2004). Family-level assignment is currently not possible for ~25 extinct genera. Of these, †Karaurus and
†Kokartus may be outside crown-group Caudata. Unassigned taxa, some of which are based on very limited material (e.g., †Galverpeton), are
†Apricosiren, †Batrachosauroides, †Bishara, †Chrysotriton, †Chunerpeton, †Comonecturoides, †Galverpeton, †Hemitrypus, †Hylaeobatrachus,
†Iridotriton, †Jeholotriton, †Kiyatriton, †Laccotriton, †Liaoxitriton, †Marmorerpeton, †Mynbulakia, †Nesovtriton, †Paleoamphiuma,
†Pangerpeton, † Prosiren, †Ramonellus, †Regalerpeton, †Seminobatrachus, †Sinerpeton, †Triassurus (possibly not within Caudata; Estes, 1981;
Milner, 2000), and †Valdotriton.
65.The monophyly is well established for Ambystomatidae and Dicamptodontidae, and the clades are sister taxa (Larson, 1991; Larson & Dimmick,
1993; Frost et al., 2006; Roelants et al., 2007; Vieites et al., 2007, 2009; Wiens, 2007; Zhang & Wake, 2009a; Pyron & Wiens, 2011). Frost et al.
(2006) reduced Dicamptodontidae to a subfamily of Ambystomatidae to avoid two taxa each with only one genus. Significantly, the decision by
Frost et al. (2006) to place the Dicamptodontidae in the Ambystomatidae because “each contain[s] a single genus” (p. 118) disregards the long
recognition of extinct genera as belonging to these families (e.g., Estes, 1981). In light of the deep separation of the taxa (probably in excess of 100
my: Roelants et al., 2007; Zhang & Wake, 2009a; Vieites et al., 2009), the long fossil record of dicamptodontids, documentation of their former
occurrence in Europe (Venczel, 2008), and substantial biological differences between the two clades, we recognize these as two distinct families.
We follow Estes (1981) by including five extinct genera (†Ambystomichnus [an ichnotaxon; Peabody, 1954], †Bargmannia, †Chrystotriton,
†Geyeriella, and †Wolterstorffiella) in the Dicamptodontidae. While we agree with Milner (2000) that the inclusion of these in Dicamptodontidae is
based on very limited data (i.e., vertebral morphology; Estes, 1981), it is the only evidence at hand and serves as a hypothesis to be tested with
additional data. Rogers (1976) described †Amphitriton as an extinct genus of Ambystomatidae; Estes (1981) and Milner (2000) have shown that the
morphological features of this extinct taxon fall within the diversity observed in Ambystoma.
66.The monophyletic Amphiumidae contains three extant species of Amphiuma. Amphiumidae is the sister-taxon of the Plethodontidae (Larson &
Dimmick, 1993; Frost et al., 2006; Roelants et al., 2007; Vieites et al., 2007, 2009; Wiens, 2007; Zhang & Wake, 2009a; Pyron & Wiens, 2011).
We include †Proamphiuma, and refrain from including †Paleoamphiuma because of disagreements as to its affinities (Rieppel & Grande, 1998;
Gardner, 2003).
67.The †Batrachosauroididae is an extinct, enigmatic family of uncertain phylogenetic affinity with seven currently recognized genera
(†Batrachosauroides, †Mynbulakia, †Opisthotriton, †Palaeoproteus, †Parrisia, †Peratosauroides, and †Prodesmodon; Estes, 1969; Denton &
O’Neill, 1998; Milner 2000). Estes (1981) considered batrachosauroidids to be closely related to extant proteids. Other enigmatic genera such as
†Hylaeobatrachus and †Prosiren may also belong to this lineage (Milner, 2000)
68.The monophyletic Cryptobranchidae (e.g., Wiens, 2007; Zhang & Wake, 2009a; Pyron & Wiens, 2011) contains two extant genera (Andrias and
Cryptobranchus) and two extinct genera (†Aviturus and †Ulanurus). Gubin (1991) placed †Aviturus and †Ulanurus in the subfamily †Aviturinae
within the Cryptobranchidae, but we refrain from recognizing this subfamily taxonomy pending cladistic analyses. Four other extinct genera
(†Chunerpeton, †Jeholotriton, †Pangerpeton, and †Regalerpeton) form a clade with extant cryptobranchids exclusive of other salamanders but
these have not been formally included within the Cryptobranchidae (Zhang et al. 2009; Skutschas & Gubin, in press; see also Wang & Evans,
2006). Estes (1981) provides details on other taxa considered junior synonyms of Andrias. Skutschas (2009) also includes both †Eoscapherpeton
and †Horezmia in the Cryptobranchidae.
69.See footnote 65.
70.Hynobiidae is monophyletic (e.g., Frost et al., 2006; Zhang & Wake, 2009a; Zheng et al., 2011; Pyron & Wiens, 2011) and sister to the
Cryptobranchidae (Larson, 1991; Larson & Dimmick, 1993; Frost et al., 2006; Roelants et al., 2007; Vieites et al., 2007, 2009; Wiens, 2007; Zhang
& Wake, 2009a; San Mauro, 2010; Pyron & Wiens, 2011). A close relationship between the Hynobiidae and Cryptobranchidae is supported by
morphological data (Noble, 1925; Larsen, 1963; Larson & Dimmick, 1993). Molecular phylogenetic analysis of the recently rediscovered
Protohynobius puxiongensis reveals that it is closely related to Pseudohynobius and nested well within Hynobiidae (Peng et al., 2010), thus
invalidating the former subfamily Protohynobinae (Fei & Ye, 2000). We follow Venczel (1999) by including the extinct †Parahynobius in the
Hynobiidae.
71.†Karauridae is an extinct family comprising two genera (†Karaurus and †Kokartus) from the Jurassic of middle Asia (Ivachnenko, 1978; Nessov,
1988). The †Karauridae is believed to be the sister taxon of extant salamanders (Evans & Milner, 1996; Evans et al., 2005; Skutschas & Martin,
2011; Skutschas & Gubin, in press). Another extinct genus, †Marmorerpeton, may be allied to the †Karauridae (Milner, 2000).
72.Evidence for two major clades within the Plethodontidae is strong (Vieites et al., 2007, 2011; Camp et al., 2009) and two subfamilies,
Hemidactyliinae and Plethodontinae, are recognized (contra Pyron & Wiens, 2011). Vieites et al. (2011) recognized four tribes in the
Hemidactyliinae and five in the Plethodontinae; their Bolitoglossini and Spelerpini were treated as subfamilies by Chippindale et al. (2004) and
Pyron & Wiens, (2011). Each subfamily and each tribe is well resolved as monophyletic in molecular phylogenetic analyses (e.g., Chippindale et
al., 2004; Mueller et al., 2004; Macey, 2005; Min et al., 2005; Frost et al., 2006; Wiens, 2007; Kozak et al., 2009; Vieites et al., 2011, Pyron and
Wiens, 2011). However, inferred relationships among the tribes have changed over time with increases in phylogenetic data (Hedges & Maxson,
1993; Hay et al., 1995; Chippindale et al., 2004; Mueller et al., 2004; Macey, 2005; Min et al., 2005; Frost et al., 2006; Roelants et al., 2007;
Wiens, 2007; Kozak et al., 2009; Vieites et al., 2011).

Zootaxa 3148 © 2011 Magnolia Press · 47
ANIMAL BIODIVERSITY
Subfamily Necturinae Fitzinger, 1843 (1 genus; 5 species)
Family Rhyacotritonidae Tihen, 1958 (1 genus; 4 species)74
Family Salamandridae Goldfuss, 1820 (21–22 genera; 86 species; †9)75
Subfamily Pleurodelinae Tschudi, 1838 (16–17 genera; 69 species; †7)
Subfamily Salamandrinae Goldfuss, 1820 (4 genera; 15 species; †1)
Subfamily Salamandrininae Fitzinger, 1843 (1 genus; 2 species)
Family †Scapherpetontidae Auffenberg & Goin, 1959 (†3)76
Family Sirenidae Gray, 1825 (2 genera; 4 species; †3)77
Order Gymnophiona Müller, 1832 (caecilians) (31 genera; 188; †3)78
Family Caeciliidae Rafinesque, 1814 (2 genera; 42 species)79
Family Dermophiidae Taylor, 1969 (4 genera; 14 species; †1)80
Family Herpelidae Laurent, 1984 (2 genera; 9 species)81
Family Ichthyophiidae Taylor, 1968 (3 genera; 50 species)82
Family Indotyphlidae Lescure, Renous & Gasc, 1986 (7 genera; 21 species)83
Family Rhinatrematidae Nussbaum, 1977 (2 genera; 11 species)84
Family Scolecomorphidae Taylor, 1969 (2 genera; 6 species)85
73. Monophyly of the Proteidae has long been debated (e.g., Larsen & Guthrie, 1974; Hecht & Edwards, 1976). Molecular phylogenetic studies recover
a monophyletic Proteidae but with very long internal branches and a split likely in excess of 120 my (Frost et al., 2006; Roelants et al., 2007;
Wiens, 2007; Zhang & Wake, 2009a). While we recognize a single family, there are extinct taxa assigned to Proteinae and a reasonable alternative
would be to raise the subfamilies (Proteinae and Necturinae) to family level (Zhang & Wake, 2009a). The extinct taxa †Mioproteus and
†Orthophyia are morphologically similar to Proteus (Estes, 1981; Milner, 2000) and we place these genera in the Proteinae.
74.The Rhyacotritonidae, containing only the genus Rhyacotriton, is sister to the clade comprising the Amphiumidae and Plethodontidae (Mueller et
al., 2004; Min et al., 2005; Frost et al., 2006; Wiens et al., 2005; Roelants et al., 2007; Vieites et al., 2007, 2011; Wiens, 2007; Zhang & Wake,
2009a; Pyron & Wiens, 2011; Zheng et al., 2011).
75. Monophyly of the Salamandridae is well established based on morphological and molecular evidence (Wake & Özeti, 1969; Titus & Larson, 1995;
Frost et al., 2006; Wiens, 2007; Roelants et al., 2007; Zhang et al., 2008; Zhang & Wake, 2009a; Pyron & Wiens, 2011). Molecular phylogenetic
analyses reveal three well resolved clades of salamandrids (Weisrock et al., 2006; Zhang et al., 2008; Pyron & Wiens, 2011) that are each now
recognized as a subfamily (Dubois & Raffaëlli, 2009). Following Estes (1981) and Dubois & Raffaëlli (2009), we recognize nine extinct genera in
the Salamandridae, with seven in the Pleurodelinae (†Brachycormus, †Carpathotriton, †Chelotriton, †Koallelia, †Oligosemia [considered a
possible synonym of Triturus by Estes, 1981], †Palaeopleurodeles, and †Procynops) and one in the Salamandrinae (†Megalotriton). Unlike Dubois
& Raffaëlli (2009), we consider the extinct †Archaeotriton as Salamandridae incertae sedis because its affinities remain unclear (Estes, 1981;
Venczel, 2008).
76.The extinct family †Scapherpetontidae includes three genera: †Lisserpeton, †Piceoerpeton, and †Scapherpeton (Estes, 1969; Naylor & Krause,
1981). Following Estes (1981), we recognize this as a distinct taxon instead of subsuming these genera within the Dicamptodontidae based on
vertebral morphology (i.e., Edwards, 1976). The relationship of the †Scapherpetontidae to other salamander families remains unclear.
77.The phylogenetic relationship of the two genera of the Sirenidae (Pseudobranchus and Siren) to other salamanders has long been enigmatic (e.g.,
Boyden & Noble, 1933; Larsen, 1963; Estes, 1965). Its monophyly is well established, with most studies resolving it as the sister to a large clade
(Salamandroidea) containing Ambystomatidae, Dicamptodontidae, Plethodontidae, Proteidae, Rhyacotritonidae, and Salamandridae (Wiens et al.,
2005; Roelants et al., 2007; Wiens, 2007; Pyron & Wiens, 2011), although Zhang & Wake (2009a), using complete mitochondrial genomes, found
Sirenidae to be sister to all other salamanders. We follow Evans et al. (1996) by including three extinct genera in the Sirenidae (†Habrosaurus,
†Kababisha, and †Noterpeton), though we note that the affinities of these taxa remain enigmatic (Rage et al., 1993; Milner, 2000).
78.Doubts about monophyly led Frost et al. (2006) to recognize only three families: Caeciliidae, Typhlonectidae, and Rhinatrematidae. In order to
resolve apparent paraphyly, Wilkinson et al. (2011) recognized nine familes. Their classification is compatible with the results of the most
comprehensive molecular phylogenetic analyses (Roelants et al., 2007; Zhang & Wake, 2009b; Pyron & Wiens, 2011), yet many taxa remain
unsampled. We adopt the classification of Wilkinson et al. (2011), which identifies major clades and presents detailed justifications for recognizing
these as families (rather than subfamilies as in Pyron & Wiens, 2011); because of this recent presentation, we do not go into detailed summaries for
each family of caecilians. The Caeciliidae (sensu Nussbaum & Wilkinson, 1989; see also Wilkinson & Nussbaum, 2006) is the family-level taxon
most affected by changes proposed by Wilkinson et al. (2011); it is divided into five families (Caeciliidae, Dermophiidae, Herpelidae,
Indotyphlidae, and Siphonopidae). †Apodops, described by Estes & Wake (1972) as a caeciliid based on stated similarities to Dermophis,
Gymnophis, and Geotrypetes, is tentatively placed in the Dermophiidae (M. Wake, pers. comm.). Of the three extinct genera of Gymnophiona, only
†Apodops is within crown-group Gymnophiona; †Eocaecilia and Rubricaecilia are likely sister to extant caecilians (Jenkins & Walsh, 1993; Evans
& Sigogneau-Russell, 2001; Jenkins et al., 2007).
79.See footnote 78.
80.See footnote 78.
81.See footnote 78.
82.Most phylogenetic analyses agree in finding a close relationship between the Icthyophiidae and the former Uraeotyphlidae (Wilkinson &
Nussbaum, 1996; Gower et al., 2002; Wilkinson et al., 2003; San Mauro et al., 2004b, 2009; Frost et al., 2006; Loader et al., 2007; Roelants et al.,
2007; Gower et al., 2008; Gower & Wilkinson, 2009; Zhang & Wake, 2009b), although most of these studies are based on a single representative of
each taxon. Frost et al. (2006) subsumed the Uraeotyphlidae within the Ichthyophiidae based on data for three specimens, one of which was
unidentified to species. In studies with richer taxon sampling, Gower et al. (2002), Zhang & Wake (2009b), and Pyron & Wiens (2011) found
Ichthyophis to be paraphyletic with respect to Uraeotyphlus, and Ichthyophis is paraphyletic with respect to Caudicaecilia (Roelants et al., 2007;
Zhang & Wake, 2009b; Pyron & Wiens, 2011). The validity of genera in the Ichthyophiidae requires further research (i.e., Wilkinson et al., 2011).
83.See footnote 78.

BLACKBURN & WAKE: CLASS AMPHIBIA GRAY, 1825
48 · Zootaxa 3148 © 2011 Magnolia Press
Family Siphonopidae Bonaparte, 1850 (7 genera; 22 species)86
Family Typhlonectidae Taylor, 1968 (4 genera; 13 species)87
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Author Addresses
David C. Blackburn, Department of Vertebrate Zoology & Anthropology, California Academy of Sciences, 55 Music Concourse
Drive, San Francisco, California 94118 USA; dblackburn@calacademy.org
David B. Wake, Museum of Vertebrate Zoology and Department of Integrative Biology, University of California, Berkeley, California
94708-3160 USA; wakelab@berkeley.edu