C HA P T E R 19

A New Era of Amphibian

Taxonomy
Jennifer B. Pramuk and
Joseph R. Mendelson III

The more than 7000 species of amphibians represent a mono- In the case of amphibians, a stable taxonomy is elusive in
phyletic group (i.e., sharing a common ancestor) that is both part because knowledge of amphibian diversity is still grow-
highly successful and geographically widespread. The earliest ing at an astonishing rate. Compared with the largely stable
amphibians, of which the fossils of Ichthyostega and Elginer- taxonomies for mammals or birds, new major lineages (e.g.,
peton are representatives, date from the Late Devonian, more genera or even families) of amphibians are still being discov-
than 360 million years ago. These fossil forms represent the ered regularly. To put this into perspective, approximately one
earliest known ancestors to all extant tetrapods, which include quarter of all known amphibian diversity has been described
amphibians, birds, reptiles, and mammals. Amphibians were in the past 20 years, and the rate of species discovery has not
the first vertebrates to hunt, vocalize, and breed on land, yet plateaued. To confound matters, the last century had been
and their basic body plan has changed little since the Juras- spent tolerating a flawed traditional taxonomy that used use-
sic period. Because of their highly permeable skin, amphib- less names for artificial and large groups of animals, such as
ians are usually intolerant of saltwater; therefore, their wide “Leptodactylidae,” a catchall group of neotropical frogs that
present-day distribution is largely the result of deeper lineages was far from phylogenetically accurate. The new, molecular
predating the split of continental landmasses—today, they are data–based phylogenies have revealed that the vast age of
present on all continents except Antarctica. amphibians and the phenomenon in many groups in which
Because closely related organisms are expected to function larval characteristics are retained in adults (i.e., neoteny)
similarly, understanding taxonomy and phylogeny might yield have led to repeated appearances and reversals of morpho-
clues to their physiology and natural history. In most cases, all logic traits that were formerly considered to be reliable indi-
but the most commonly kept amphibian species have little to cators of shared ancestry. These results have confounded or
nothing about them in the literature regarding their medical completely revised earlier morphologic-based phylogenies
treatment or husbandry. A fundamental understanding of the and classifications. Presented herein are resources for under-
evolutionary relationships or phylogeny of the amphibians is standing not only current amphibian taxonomy but also the
therefore integral to an understanding of their biology, captiv- challenges of relying on a stable taxonomy that is undergoing
ity, and medical care. For example, if one is treating a member constant flux.
of a poorly known taxon, such as a leaf frog in the genus From activities ranging from teaching to veterinary prac-
Phasmahyla, clues to its captive management (e.g., sensitivities tice, a taxonomy should accurately communicate the state of
to a particular medication or susceptibility to diseases such the art in knowledge related to both actual biodiversity and
as Batrachochytrium dendrobatidis [amphibian chytrid fun- the phylogenetic relationships among species and groups (e.g.,
gus]) might be revealed by researching literature on its most genera). A recent pulse of activity in amphibian systematics,
widely studied sister taxon, in this case, the Red-eyed Tree frog spurred largely by technologic improvements in collecting and
(Agalychnis callidryas). analyzing massive molecular data sets, has greatly increased
It is ironic that the notion of stability is so entrenched in the the understanding of the relationships among and within the
concept of taxonomy that the very system of names designed major groups; this new information has appropriately spurred
to allow communication across cultural and language barriers efforts to revise the antiquated, misleading taxonomy of the
cannot itself be changed without reactionary outbursts from last century or so. Given that novel taxonomic arrangements
the greater community of biologists, land managers, veterinar- for long-familiar groups can be confusing and frustrating, this
ians, and even the taxonomists themselves. It is as if some feel chapter endeavors to summarize in a user-friendly fashion
that taxonomic stability (tradition) is more important than the the most recent taxonomic changes, especially as they relate
development of knowledge of biodiversity (progress). Is one to groups likely to be encountered by readers of this chapter.
not supposed to learn and move forward in understanding of Ideally a taxonomy should reflect the phylogeny—the actual
biodiversity? To witness the reaction some have to proposed evolutionary history—of the creatures under consideration.
taxonomic changes, one would infer not! Taxonomic stabil- Thus, this summary will hopefully be useful for keepers, cura-
ity is a good thing to have but only if the taxonomy in use tors, and veterinarians working with species that are poorly
adequately communicates both the diversity and evolutionary known or with which they are simply unfamiliar. Importantly,
history of the group of organisms in question. knowledge of the phylogenetic position of a species may allow

230
 CHAPTER 19   •  A New Era of Amphibian Taxonomy 231

one to extrapolate information relevant to other closely related accordingly. The International Species Information System
species, genera, or from another family. (ISIS) record-keeping system officially lists Frost23 plus the
Because taxonomic monographs and treatments include addendum by Duellman24 as their taxonomic standard. How-
exhaustive citations of all relevant literature and complete ever, because both of those sources are quite outdated, it can
reviews of taxonomic changes and synonymies, those efforts be assumed that they now refer to the continuation of that
will not be duplicated here and readers are referred to the same project, which now exists online in the form of Amphib-
original monographs. The current renaissance in amphibian ian Species of the World.22 An additional useful Web site is
taxonomy was spurred by a series of three monographs pub- the International Union for Conservation of Nature (IUCN)
lished by the American Museum of Natural History.1-3 The Red List (http://www.iucnredlist.org/initiatives/amphibians),
Amphibian Tree of Life3 was especially influential because it which provides detailed information for every known species
represented the first modern attempt to consider amphibian of amphibian regarding distribution, conservation status, and
phylogeny in its entirety. Never claiming to be the last word more, although this particular database is not intended to be
on knowledge of amphibian phylogeny or taxonomy, it did a source of the most current taxonomic status of any species
proceed to frame and encourage a flood of subsequent articles or group.
that, considered together, have greatly increased knowledge
of the history of Amphibia and represent major steps toward
producing a taxonomy that is consistent with their history. PHYLOGENY AND TAXONOMY
Some important recent articles include treatments of the OF AMPHIBIANS
phyllomedusine frogs,4 hemiphractid frogs (formerly part Amphibians are a distinct group of tetrapod vertebrates,
of Hylidae),5 microhylid frogs,6 glass frogs,7 ranid frogs,8 and despite centuries of communications to the opposite,
bufonid frogs,9-12 and the terraranan frogs (the so-called they are in no way “transitional” between fishes and other
eleutherodactylines, formerly included in Leptodactylidae).13 groups of tetrapods (e.g., reptiles). For the most part, refer-
Additional recent efforts have informed the understanding ring a specimen in hand to one of the three major groups
of the broader groups such as caecilians14-16 and salaman- of amphibians (i.e., caecilians, frogs, and salamanders) has
ders.17-19 Recently, Pyron and Wiens20 assembled virtually never been problematic. However, reconciling the relation-
all of the molecular data produced by the mentioned series ships among these three groups has never been intuitive.
of articles (plus others not here listed) into a single massive This is because frogs and caecilians both have highly derived
analysis. Their study largely confirmed these previous more body forms while salamanders remain with what is frequent-
focused efforts and added some refinements to the ever-prob- ly considered to be the generic tetrapod body plan. Some
lematic assemblage of South American frogs formerly referred debate has occurred regarding the relationships among these
to “Leptodactylidae.” However, relationships within the large three groups,25 but the arrangement shown in Figure 19-1 is
superfamily Hyloidea (including many families widely kept now generally accepted.
in captivity, such as the hylid tree frogs) were mostly poorly Taxonomically, or in terms of nomenclature, this arrange-
supported. Moreover, their phylogeny inexplicably retained ment
   can be summarized as follows:
an untenable nonmonophyletic taxonomy for Bufonidae and Class Amphibia: amphibians
Ranidae. Table 19-1 lists a taxonomic summary of generally Order Gymnophiona: caecilians
recognized families of amphibians, including approximate Infraclass Batrachia: frogs and salamanders
information on species-level diversity within each and geo- Order Caudata: salamanders
graphic distribution. Order Anura: frogs
  
Current generic-level taxonomy has not been emphasized Currently, 7170 species of amphibians are known, and
nor summarized in this chapter because generic limits and spe- the majority of those are anurans (note: the homepage for
cies allocations are in constant flux, and, in the case of amphib- amphibiaweb.org includes a convenient daily update of the
ians, considerable changes at this level can be anticipated in number of recognized species). Frost et al3 characterize
the upcoming years. For current information regarding generic amphibians as having, among many other unique features, loss
allocations, the reader is referred to these frequently updated of several bones in the typical tetrapod skull (e.g., supratempo-
online references: The American Museum of Natural History’s ral, jugal, postorbital bones, Figure 19-2, A), biscuspid, pedi-
Amphibian Species of the World at http://research.amnh.org/ cellate teeth (Figure 19-2, B), a unique papilla amphibiorum
herpetology/amphibia/index.php and The University of Califor­ in the ear associated with the and opercular element (Figure
nia’s Amphibiaweb at http://www.amphibiaweb.org/. 19-2, C), glandular skin (Figure 19-2, D), gonadal fatbodies,
For species occurring in North America, the standard and a lack of epidermal scales.
English and scientific names list cosponsored by all four of
the major herpetological societies based in the United States
is recommended.21 The most recent edition is available for PHYLOGENY AND TAXONOMY OF CAECILIANS
free download by the Society for the Study of Amphibians The caecilians, or Gymnophiona, are the most geographically
and Reptiles at http://www.ssarherps.org/pages/comm_names/ restricted (limited in distribution to tropical South America,
Index.php. Asia, and Africa but absent from Madagascar) and secretive
Readers working in institutions accredited by the Asso- of all amphibian orders. As with all aspects of caecilian biol-
ciation of Zoos and Aquariums (AZA) should note that AZA ogy, their true species-level diversity is poorly known. None-
follows the taxonomy of Amphibian Species of the World Web theless, known taxa are commonly referred to 10 families
site22; however, this Web site is updated frequently, and it has (Figure 19-3), and approximately 192 species are currently
not been standard for AZA institutions to update their records recognized. Future taxonomic changes are to be expected
232 SECTION III   •  ADVANCES IN AMPHIBIAN MEDICINE

TA B LE 1 9-1

A Taxonomic Summary of Generally Recognized Families of Amphibians*

Approximate Number
Taxon English Name of Species General Distribution
GYMNOPHIONA CAECILIANS 192 PANTROPICAL
Caeciliidae N/A 42 South and Central America
Chikilidae N/A 1 India
Dermophiidae N/A 14 Central and South America and Africa
Herpelidae N/A 9 Africa
Ichthyophiidae N/A 53 South and Southeast Asia
Indotyphlidae Including yellow-banded caecilians 20 India, Africa and the Seychelles
Rhinatrematidae N/A 11 South America
Scolecomorphidae N/A 6 Africa
Siphonopidae N/A 23 South America
Typhlonectidae Aquatic caecilians 13 northern South America
CAUDATA SALAMANDERS 648 EURASIA AND AMERICAS
Ambystomatidae Mole salamanders 32 North America
Amphiumidae Amphiumas 3 North America
Cryptobranchidae Giant salamanders and hellbenders 3 Asia and North America
Dicamptontidae Pacific giant salamanders 4 North America
Hynobiidae Asian salamanders 59 Asia
Plethodontidae Lungless salamanders 435 Asia, Europe, and Americas
Proteidae Muduppies and waterdogs 6 Europe and North America
Rhyacotritonidae Torrent salamanders 4 North America
Salamandridae “True” salamanders 98 Eurasia and North America
Sirenidae Sirens 4 North America
ANURA FROGS 6289 COSMOPOLITAN, EXCEPT ANTARCTICA
Allophrynidae N/A, but including Tukeit-hill frog 2 South America
Alsodidae N/A 30 South America
Alytidae Midwife toads 5 Europe and North Africa
Arthroleptidae N/A, but including Hairy frogs 147 Sub-Saharan Africa
Ascaphidae Tailed frogs 2 Northwestern United States and Canada
Batrachylidae N/A 15 Argentina and Chile
Bombinatoridae Fire-bellied toads 10 Eurasia and Southeast Asia
Brachycephalidae N/A 52 South America
Brevicipitidae N/A, but including rain frogs 34 Sub-Saharan Africa
Bufonidae “True” toads 585 Cosmopolitan, except Australo-Papua,
Oceania and Madagascar
Calyptocephalellidae N/A, but including the Helmeted 4 South America
Water toad
Centrolenidae Glass frogs 152 Mesoamerica and South America
Ceratobatrachidae N/A 86 Southeast Asia
Ceratophryidae Including Budgett’s and Horned frogs 12 South America
Ceuthomantidae N/A 4 South America
Conrauidae N/A, but including the Goliath frog 6 Equatorial Africa
Craugastoridae Stream frogs 115 Mesoamerica and South America
Cycloramphidae N/A 34 South America
Dendrobatidae Poison frogs 293 Central and South America
Dicroglossidae N/A 185 Africa and Asia
Discoglossidae Painted frogs 6 Europe and northern Africa
Eleutherodactylidae Rain frogs 206 Americas, including Caribbean
Heleophrynidae Ghost frogs 6 South Africa
Hemiphractidae Marsupial frogs 101 Central and South America
Hemisotidae Shovel-nosed frogs 9 Sub-Saharan Africa
Hylidae Tree frogs 929 Eurasia, Australo-Papua, and Americas
Hylodidae N/A 42 South America
Hyperoliidae Reed frogs 223 Africa and Madagascar
Leiopelmatidae New Zealand frogs 4 North America and New Zealand
Leptodactylidae Southern frogs 200 Mesoamerica and South America
Mantellidae Mantellas 205 Madagascar
Megophryidae N/A but including Horned frogs and 174 Southeast Asia
Toothed toads
Micrixalidae N/A 11 India
Continued
 CHAPTER 19   •  A New Era of Amphibian Taxonomy 233

TA B LE 1 9-1

A Taxonomic Summary of Generally Recognized Families of Amphibians—cont’d

Approximate Number
Taxon English Name of Species General Distribution
Microhylidae Narrow-mouthed toads 525 Americas, Africa, Southeast Asia, and
Australo-Papua
Myobatrachidae Australian ground and water frogs 131 Australo-Papua
Nasikabatrachidae Purple frog 1 India
Nycibatrachidae N/A 28 India
Odontophrynidae Includes Horned frogs 41 South America
Pelobatidae Spadefoot toads 4 Eurasia and North Africa
Pelodytidae Parsley frogs 3 Europe
Petropedetidae N/A 12 Sub-Saharan Africa
Phrynobatrachidae N/A 86 Sub-Saharan Africa
Pipidae Clawed frogs and Surinam toads 33 Central and South America and sub-
Saharan Africa
Ptychadenidae Rocket frogs 51 Sub-Saharan Africa
Pyxicephalidae N/A, including African Bullfrogs 73 Sub-Saharan Africa
Ranidae “True” frogs 362 Cosmopolitan
Ranixalidae N/A 10 India
Rhacophoridae N/A, but including foam-nesting and 361 Sub-Saharan Africa and Asia
flying frogs
Rhinodermatidae Darwin’s frogs 3 Chile
Rhinophrynidae Mexican Burrowing toad 1 Mesoamerica
Scaphiopodidae American Spadefoot toads 7 North America, including Mexico
Sooglossidae Seychelles frogs 4 Seychelles
Strabomantidae Rain frogs 603 Central and South America
Telmatobiidae Includes water frogs 61 South America

*Based on Frost DR, Grant T, Faivovich J, et al. The amphibian tree of life. Bull Am Museum Nat Hist 2006;297:1-370; Pyron RA, Wiens JJ. A large-
scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Mol Phylogenet Evol
2011;61:543-583; and Frost DR. Amphibian species of the world: an online reference. Version 5.5. Electronic database available at http://research.amnh
.org/vz/herpetology/amphibia/. Accessed January 31, 2011. Approximate numbers of species in each group and a very general description of geographic
distribution are also presented (species counts are from Amphibiaweb.org).

including a heavily ossified skull that makes them highly

Gymnophiona

adapted to a fossorial or burrowing lifestyle. Frost et al3 char-

Caudata

acterize the gymnophionans as being legless amphibians that

Anura

lack both limbs and girdles (but one fossil taxon is known to
have had four limbs) and having a unique dual jaw-closing
mechanism, an eversible male intromittent organ (the phal-
lodeum), dermal annuli that encircle the body, and unique
Batrachia paired sensory tentacles on the snout. With the exception of
one species (Atretochoana eiselti), all caecilians have lungs, but
Amphibia often the left lung is reduced in size, which is an adaptation for
an elongate body shape, much as in snakes. Some species also
have dermal calcite scales and are the only living amphibians
FIGURE 19-1 Relationships among the three major groups known to possess them.
of amphibians: Gymnophiona (caecilians), Anura (frogs), and
Caudata (salamanders). (Adapted from Frost DR, Grant T,
Faivovich J, et al. The amphibian tree of life. Bull Am Museum PHYLOGENY AND TAXONOMY
Nat Hist 2006;297:1-370.) OF SALAMANDERS
The salamanders represent a moderately speciose group
(approximately 648 species) distributed primarily in North
in this group as additional material for study, discoveries America, Central America, northern South America, and Eur-
of new species, and studies are forthcoming. For example, asia. Recent efforts in the systematics of salamanders did not
Kamei et al26 recently discovered a new major lineage (fam- greatly modify the understanding of the relationships among
ily Chikilidae) and suggest that it contains numerous yet the major groups (and Figure 19-4) nor their family-level
unnamed species. taxonomy. Readers should be aware that some continuing
Caecilians, superficially resembling earthworms or snakes controversy exists regarding subfamilial classification of sala-
in general appearance, share a suite of morphologic features, manders and should consult the Web sites of Amphibiaweb
234 SECTION III   •  ADVANCES IN AMPHIBIAN MEDICINE

PM N

C
SP FP

TS
PT
P

QJ
PR SQ
A E B RC

U OC GC
PA

MO T
PC

C O
FIGURE 19-2 Amphibians are characterized as having several synapomorphies (i.e.,
unique, shared, derived characteristics) some of which are illustrated here. A, Amphibian
evolution includes the loss of several bones found in the typical tetrapod skull (e.g., supra-
temporal, jugal, postorbital bones) through the fusion of elements such as the frontal and
parietal (frontoparietal). Dorsal view of the skull of the North American bullfrog Lithobates
catesbeianus. E, Exoccipital; FP, frontoparietal; M, maxilla; N, nasal; PR, prootic; PM, premax-
illa, PT, pterygoid; QJ, quadratojugal; SP, sphenethmoid; SQ, squamosal.  (Drawing made
from the DigiMorph online catalog of CT scans: http://www.digimorph.org/specimens/
Rana_catesbeiana//.)  B, Lingual view of the biscuspid, pedicellate teeth of the anuran
Calyptocephalella caudiverbera. C, Crown; P, pedicel; RC, replacement crown; TS, trans-
verse suture.  (Drawing adapted from Parsons TS, Williams EE. The teeth of Amphibia and
their relation to amphibian phylogeny. J Morphol 1962;110:375-389.)  C, The inner ear of
Lithobates catesbeianus illustrating the papilla amphibiorum (PA), as well as the opercular
element (O) and the columella (C). MO, Medulla oblongata; OC, otic capsule; PC, perilym-
phatic cistern; T, tympanum; U, utriculus.  (Drawing adapted from Duellman WE, Trueb L.
Biology of Amphibians. Baltimore: The John Hopkins University Press, 1994.)  D, Cross
section of the skin of the poisonous toad Melanophryniscus montevidensis illustrating
granular glands. (Drawing adapted from Mebs D, Pogoda W, Maneyro R, et al. Studies on
the poisonous skin secretion of individual red bellied toads, Melanophryniscus montevi-
densis [Anura, Bufonidae], from Uruguay. Toxicon 2005;46:641-650.)

and Amphibian Species of the World to consider current have simply retained the primitive ancestral tetrapod body
perspectives; this controversy does not, however, greatly plan. This is an overstatement in many senses, and Frost et al3
affect familial or generic level classifications. Salamanders characterize the caudatans as being unique among amphibians
have always been difficult to characterize in terms of unique by having both forelimbs and a tail and an incomplete maxil-
morphologic aspects because most persons assume that they lary arcade (upper jaw). Additionally, salamanders are unique
 CHAPTER 19   •  A New Era of Amphibian Taxonomy 235

Scolecomorphidae
PHYLOGENY AND TAXONOMY OF FROGS

Typhlonectidae

Rhinatremidae
Ichthyophiidae
Dermophiidae
Siphonopidae
Indotyphlidae
Frogs, represented by more than 6289 species, have always rep-

Caeciliidae

Herpelidae
Chikilidae
resented a great challenge to systematists, as their evolutionary
history evidently includes explosive radiations, along with high
levels of both convergence and extreme morphologic derivation.
Because attempts to recover the phylogeny of frogs with morpho-
logic data alone have generally failed, molecular data have been
crucial in shaping our understanding of this complex phylogeny.
Building on the foundation of the efforts by Frost et al,3 numer-
ous studies have been produced11,20 and are greatly increasing the
knowledge about this vast and taxonomically difficult group. As
soon as an agreement on a phylogeny of frogs is reached, numer-
ous taxonomic changes will be forthcoming. Some stakeholders
have expressed frustration about changes already proposed, such
as when long-familiar species such as Rana catesbeiana and Bufo
americanus suddenly appeared as Lithobates catesbeianus and
Anaxyrus americanus. Simply put, such changes are necessary
and important so that a taxonomy that is both informative and
internally consistent can be used when communicating about
amphibians. Figure 19-5 summarizes the agreement among
Gymnophiona
recent studies on frog phylogeny. Polytomies, or unresolved
branches, represent areas of the tree where recent analyses have
FIGURE 19-3 Relationships among the major clades (fami-
lies) of caecilians. (Adapted from Kamei RG, Mauro DS,
produced conflicting results or poor statistical support; in this
Gower DJ, et al. Discovery of a new family of amphibians sense, the phylogeny shown here is conservative.
from northeast India with ancient links to Africa. Proc Biol Sci Anurans are perhaps the most familiar amphibians to most
2012;279:2396-2401.) people, and indeed, their unique body plan, adapted largely for
saltatorial locomotion (e.g., elongate hindlimbs), makes all but
the most bizarre species instantly recognizable to layperson as
Dicamptodontidae
Cryptobranchidae

frogs. The many unique features of this group that Frost et al3
Rhyacotritonidae
Ambystomatidae
Salamandridae

Plethodontidae

listed as characteristic include the following: reduction of ver-

Amphiumidae

tebral number to nine or fewer; fusion of the caudal vertebrae

Hynobiidae

Proteidae
Sirenidae

to form the bony urostyle; hindlimbs significantly longer than

forelimbs (with a few exceptions), produced, in part, by the
elongation of the ankle bones; fusion of the radius and ulna of
the forelimb and fusion of the tibia and fibula of the hindlimb;
presence of keratinous jaw sheaths and keratodonts on larval
mouthparts (with some exceptions); skin with large subcu-
taneous lymph spaces; and during mating, a unique clasping
behavior (amplexus) common to most species.

CONCLUSION
Tremendous growth in knowledge of the phylogeny of
amphibians has occurred in the last few years. This knowledge
Caudata is crucial for informing the study of all aspects of their biol-
ogy, as well as their husbandry and veterinary care, and for
FIGURE 19-4  Consensus of phylogenetic relationships among pressing conservation concerns. However, more work remains
the major clades (families) of salamanders. (Adapted from to be done, and changes in the trees presented here are antici-
Zhang P, Wake DB. Higher-level salamander relationships pated. Certainly, still more taxonomic changes will accompany
and divergence dates inferred from complete mitochondrial increased knowledge of the evolution of amphibians. The
genomes. Mol Phylogenet Evol 2009;53:492-508 and Pyron RA, regularly updated Amphibian Species of the World22 is recom-
Wiens JJ. A large-scale phylogeny of Amphibia including over
mended as a standard reference for amphibian taxonomy, but
2800 species, and a revised classification of extant frogs, sala-
manders, and caecilians. Mol Phylogenet Evol 2011;61:543-583.) the primary literature should be the source for the most recent
information on amphibian phylogeny.

in that many species, or entire groups, retain various aspects

of the larval morphology into sexual maturity (a phenomenon ACKNOWLEDGMENTS
known as neoteny). Salamanders usually do not vocalize as The authors would like to thank Darrel Frost for providing in-
most anurans do; thus they rely on pheromones and courtship formation on the latest version of the Amphibian Species of the
displays to locate and attract a mate. World Taxonomy.
236 SECTION III   •  ADVANCES IN AMPHIBIAN MEDICINE

Calyptocephalellidae

Eleutherodactylidae

Phrynobatrachidae
Nasikabatrachidae

Ceratobatrachidae
Brachycephalidae

Odontophrynidae

Rhinodermatidae

Nyctibatrachidae
Ceuthomantidae

Cycloramphidae
Scaphiopodidae

Myobatrachidae
Rhinophrynidae

Heleophrynidae
Bombinatoridae

Leptodactylidae
Hemiphractidae

Ceratophryidae
Craugastoridae

Rhacophoridae
Leiopelmatidae

Petropedetidae
Discoglossidae

Pyxicephalidae
Ptychadenidae
Dendrobatidae

Dicroglossidae
Megophryidae

Arthroleptidae
Centrolenidae

Telmatobiidae
Allophrynidae

Breviceptidae
Batrachylidae
Sooglossidae

Microhylidae
Hemisotidae
Hyperoliidae

Micrixalidae
Pelobatidae
Ascaphidae

Ranixalidae
Pelodytidae

Conrauidae

Mantellidae
Bufonidae

Hylodidae
Alsodidae

Ranidae
Alytidae

Pipidae

Hylidae
Hyloidea
Ranoidea

Neobatrachia

Anura
FIGURE 19-5 Consensus of phylogenetic relationships among the major clades (fami-
lies) of anurans or frogs. (Adapted from Pyron RA, Wiens JJ. A large-scale phylogeny of
Amphibia including over 2800 species, and a revised classification of extant frogs, sala-
manders, and caecilians. Mol Phylogenet Evol 2011;61:543-583.)

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nization: molecular phylogenetics and biogeography of the nearctic
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frog family Hylidae, with special reference to Hylinae: phylogenetic anal- 10. Pramuk JB, Robertson T, Sites JW Jr, et al. Around the world in 10 mil-
ysis and taxonomic revision. Bull Am Museum Nat Hist 2005;294:1-240. lion years: biogeography of the nearly cosmopolitan true toads (Anura:
2. Grant T, Frost DR, Caldwell JP, et al. Phylogenetic systematics of the Bufonidae). Global Ecol Biogeogr 2008;17:72-83.
dart-poison frogs and their relatives (Amphibia: Athesphatanura: Den- 11. 
Van Bocxlaer I, Loader SP, Roelants K, et al. Gradual adaptation
drobatidae). Bull Am Museum Nat Hist 2006;299:1-262. toward a range-expansion phenotype initiated the global radiation of
3. Frost DR, Grant T, Faivovich J, et al. The amphibian tree of life. Bull toads. Science 2010;327:679-682.
Am Museum Nat Hist 2006;297:1-370. 12. Mendelson JRIII, Mulcahy DG, Williams TS, et al. A phylogeny and
4. Faivovich JCF, Haddad B, Baeta D, et al. The phylogenetic relation- evolutionary natural history of mesoamerican toads (Anura: Bufoni-
ships of the charismatic poster frogs, Phyllomedusinae (Anura, Hyli- dae: Incilius) based on morphology, life history, and molecular data.
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