RESPONSE TO THE POINT OF VIEW OF GREGORY B. PAULY, DAVID
M. HILLIS, AND DAVID C. CANNATELLA, BY THE ANURAN
SUBCOMMITTEE OF THE SSAR/HL/ASIH SCIENTIFIC AND
STANDARD ENGLISH NAMES LIST
DARREL R. FROST
1,4
,ROY W. MCDIARMID
2
, AND JOSEPH R. MENDELSON, III
3
1
Division of Vertebrate Zoology (Herpetology), American Museum of Natural History, Central Park West at 79th Street,
New York, NY 10024, USA
2
US Geological Survey, Patuxent Wildlife Research Center, National Museum of Natural History, MRC 111,
P.O. Box 37012, Washington, DC 20013-7012, USA
3
Zoo Atlanta, 800 Cherokee Avenue Southeast, Atlanta, GA 30315, USA
??Stultum facit Fortuna quem vult perdere???Publilius Syrus, Maxim 911
ABSTRACT: The Point of View by Gregory Pauly, David Hillis, and David Cannatella misrepresents the
motives and activities of the anuran subcommittee of the Scientific and Standard English Names Committee,
contains a number of misleading statements, omits evidence and references to critical literature that have
already rejected or superseded their positions, and cloaks the limitations of their nomenclatural approach in
ambiguous language. Their Point of View is not about promoting transparency in the process of constructing
the English Names list, assuring that its taxonomy is adequately reviewed, or promoting nomenclatural
stability in any global sense. Rather, their Point of View focuses in large part on a single publication, The
Amphibian Tree of Life, which is formally unrelated to the Standard English Names List, and promotes an
approach to nomenclature mistakenly asserted by them to be compatible with both the International Code of
Zoological Nomenclature and one of its competitors, the PhyloCode.
Key words: Amphibia; Bufo; Rana; Species names; Taxonomic stability; Zoological nomenclature
In their Point of View (p. 115?128) Gregory
B. Pauly, David B. Hillis, and David C.
Cannatella (subsequently referred to as
PHC) allege that the Society for the Study
of Amphibians and Reptiles (SSAR), Ameri-
can Society of Ichthyologists and Herpetolo-
gists (ASIH), and Herpetologists? League
(HL), which sponsor the Scientific and
Standard English Names of Amphibians and
Reptiles of North America (Crother, 2008),
through their selection of committee mem-
bers and endorsement of review procedures,
deemed inadequate by PHC, have confound-
ed and confused users of such lists, thereby
engendering taxonomic chaos (their term).
While some of PHC?s points deserve careful
consideration, their document is in fact an
attack on the amphibian taxonomy proposed
and the analytical methods used by Frost et al.
(2006)
5
. PHC suggest that their criticisms of
our anuran Scientific and Standard English
Names subcommittee are elements of a
widely-held taxonomic and nomenclatural
viewpoint. However, evidence from publica-
tions, including those of their own research
group, suggests otherwise. As we will show,
the PHC document contains an abundance of
misinformation and misleading statements
and reflects a pattern of omission or mis-
citation of critically relevant literature, as well
as exclusion of evidence where it serves their
purpose.
PHC pose as disinterested scientists at-
tempting to protect non-systematists from
dangerous ideas. However, if their approach
were to be applied generally the central
principles of the International Code of Zoo-
logical Nomenclature (Dubois, 2006b, 2007;
4
CORRESPONDENCE: e-mail, frost@amnh.org
5
Relevant to this discussion is that Hillis and
Cannatella are two of the PI?s on a major NSF grant,
AmphibiaTree, funded in 2003, to produce a not-yet-
published comprehensive tree and taxonomy of all
amphibians. The amphibian biology website, Amphibia-
Web (http://amphibiaweb.org/aw/about/index.html) is an
extension of this same AmphibiaTree consortium and
largely reflects the viewpoints promoted by PHC. The
funded proposal, made available by NSF in redacted
form, can be downloaded from ftp://files8.cyberlynk.net
(username: NSF_grant; password: anonymous).
Herpetologica, 65(2), 2009, 136?153
E
2009 by The Herpetologists? League, Inc.
136
ICZN, 1999), which regulate a nomenclatural
system that has been universally accepted
effectively for 250 years, must be substantially
abandoned or modified.
But, regardless of what we, Pauly, Hillis, or
Cannatella do at this moment in time, the
amphibian systematics community will follow
its own course, as it has for more than two
centuries. The taxonomy it adopts and no-
menclatural codes it follows will depend
largely on the state of revisionary publications
and perceptions of what is useful. No one gets
to have the last word. Lifetimes are short,
discoveries are effectively unlimited, and the
intellectual and social fabric of systematics
changes, not only with each scientific revolu-
tion (Kuhn, 1962), but with each retirement
and subsequent hire.
AGREEMENT WITH PAULY,HILLIS,
AND CANNATELLA
Before we address the specific issues raised
by PHC, let us first state that we agree with
them that professional societies should not
officially adopt scientific names. As discussed
elsewhere (Crother, 2009: 133), the term
??official?? on the title page was used primarily
to promote common usage of non-scientific
names and to distinguish this list from an
online list of English names produced by the
Center for North American Herpetology
(Collins and Taggart, 2009: http://www.
naherpetology.org). Any implication of scien-
tific names being official (whatever that might
mean in practice) would obviate the intent of
the SSAR-HL-ASIH annotated names list,
which, beyond its primary objective of pre-
senting a set of standardized English names
for North American taxa, was to invite
additional work on the species listed by
pointing out lacunae in our knowledge of
these taxa. PHC indicated that we should have
discussed taxonomic levels above species
more fully and commented on every change
in nomenclature since 2001. Moreover, they
wondered why we did not adopt or discuss
conventions that they considered important,
and they suggested why they thought this was
so, including the allegation that we are
legislating taxonomy. Obviously, additional
discussion was warranted and we provide that
here. Regardless, none of this would have
changed the scientific names we chose to
adopt. Had we felt the need to justify the
name changes more fully, we would simply
have cited the literature (none of which was
referenced by PHC) that had already rejected
or superseded the positions advocated by
PHC (e.g., Che et al., 2007; Dubois, 2006b,
2007; Frost et al., 2008a); these citations will
appear in the Standard English Names list
when it goes online.
We also agree with PHC about two goals of
a taxonomy: to provide a reference system that
(1) facilitates communication and (2) reflects
evolutionary history. The first goal may be
guided by the principle of stability and the
second by the notion of monophyly. But, we
think that PHC have missed a third goal of
taxonomy, which is to promote cogent com-
parisons and to invite additional systematic
work by bringing attention to previously
overlooked taxonomic units. The traditional
methods to this end have been the publication
of sound taxonomic work with appropriate
taxonomic partitions and nomenclatural des-
ignations. The history of taxonomy reflects a
balancing act among these three goals, at
different times tipping in different directions
as progress is made. PHC lean strongly in
favor of stability (part of goal 1) but com-
pletely neglect goal 3. In the short term this
has merit, as do all three of the goals, but in
the longer term, as discussed below, this
makes for practical problems.
??TAXONOMIC CHAOS?? - ISITREAL?
The issues addressed in the PHC paper
surround the notion that by adopting a
taxonomy that recognizes genera smaller than
those preferred by PHC, the English Names
anuran subcommittee promoted ??taxonomic
chaos,?? something we have not noticed in our
heavy use of current systematic and conser-
vation literature, and something we do not
find evident in any other field of herpetology,
either. In addition, Frost and McDiarmid are,
in PHC?s opinion, overly influential in pro-
moting confusion, apparently because of their
willingness both to support new, useful
taxonomies and to work on taxonomic catalogs
that serve to limit confusion among non-
systematists (e.g., Amphibian Species of
June 2009] HERPETOLOGICA 137
the World [ASW; http://research.amnh.org/
herpetology/amphibia/index.php]; Integrated
Taxonomic Information System [ITIS; http://
www.itis.gov/]). Neither Frost nor McDiarmid
has any relationship with AmphibiaWeb (http://
amphibiaweb.org/; associated with PHC?s
AmphibiaTree consortium), which did not
adopt any of the Frost et al. (2006) taxonomy
below the level of family-group, or with the
IUCN Red List (http://www.iucnredlist.org/)
which largely did adopt the taxonomy of Frost
et al. (2006). So, the larger issue for PHC seems
to be that for some taxa there is more than one
message being provided among websites and in
the literature. What PHC do not mention is that
taxonomic practice by non-Austin members of
the AmphibiaTree consortium does not meet
PHC?s current standard of taxonomic practice,
either, so we can only conclude that PHC?s
point of view is not widely shared among
workers within that work-group.
PHC apparently support the idea that
nomenclatural changes, unless required to
represent monophyletic taxonomies (but re-
gardless of any other kind of utility, such as
reflecting the age of a taxon or reducing a
nomenclatural unit to a more manageable
size), are bad if they affect species found in
the United States, while identical classes of
changes?such as those supported by their
own AmphibiaTree grant?that affect the
names of severely endangered species outside
of the United States are good. Examples of
??taxonomic chaos?? laid by PHC at the feet of
Frost et al. (2006) are (1) a redelimiting of the
former genus ??Rana?? such that species in the
Americas are now members of the either the
genus Lithobates or a redelimited Rana
(which was further partitioned in the Old
World; see also Che et al., 2007), and (2) a
partitioning of the former genus ??Bufo?? such
that species in the Americas are now members
of the genera Anaxyrus, Incilius, Nanno-
phryne, Rhaebo,andRhinella (and, like the
former Rana, further partitioned in the Old
World). Similarly, Hedges et al. (2008), in a
publication arising from the AmphibiaTree
grant, partitioned the formerly enormous
genus Eleutherodactylus (.800 species) into
four families: Brachycephalidae: Ischnocnema
(part); Craugastoridae: Craugastor and Had-
dadus; Eleutherodactylidae: Diasporus and
Eleutherodactylus (redelimited); and Strabo-
mantidae: Pristimantis, Hypodactylus, Lynch-
ius, and Strabomantis. It may be that PHC do
not approve of the taxonomic changes in
Hedges et al.?s (2008) landmark revision
because many of their changes go beyond
the minimum required to render a monophy-
letic taxonomy. Cannatella, nevertheless,
seems to have adopted these nomenclatural
novelties (e.g., see Elmer and Cannatella,
2008).
Numerous major taxonomic revisions of
recent years (e.g., Faivovich et al., 2005
[revision of Hylinae]; Glaw and Vences, 2006
[revision of Mantellidae]; Grant et al., 2006
[revision of Dendrobatoidea]; Hedges et al.,
2008 [revision of former Brachycephalidae,
now Brachycephalidae, Craugastoridae, Eleu-
therodactylidae, and Strabomantidae]) and
their resulting nomenclatural changes have
been rapidly and widely accepted by investi-
gators who deal with the substantial amphib-
ian biodiversity outside of the United States
(of which at least 32% are threatened or
extinct; Stuart et al., 2008). These revisions
reduced the ambiguity resulting, in part, from
extremely large genera. Hyla, Mantidactylus,
Colostethus, and Eleutherodactylus were all
partitioned beyond the minimal amount
allowed under the PHC dictate, but these
partitions have enhanced clarity, not promot-
ed any substantial confusion. It is also clear
that workers from biodiverse countries (e.g.,
Argentina, Brazil, China, Colombia, India,
Mexico, South Africa, Sri Lanka) as well as
those from newer centers of taxonomic
expertise outside of the United States (e.g.,
Belgium, Germany, Spain) are now increas-
ingly driving amphibian taxonomy, with the
United States becoming less influential. A few
contrary Americans are not going to hold back
this tide.
The taxonomic changes in Bufonidae and
Ranidae with which PHC take particular issue
were published more than three years ago
(Frost et al., 2006) and enjoy a healthy level of
acceptance and professional discussion out-
side of the immediate AmphibiaTree work-
group, by those who publish on the system-
atics of these groups (e.g., Caramaschi and
Pombal, 2006; Chaparro et al., 2007; Che et
al., 2007; Di Candia and Routman, 2007;
138 HERPETOLOGICA [Vol. 65, No. 2
Fouquet et al., 2007; Goebel et al., 2009; Lehr
et al., 2007; Lima et al., 2007; Maciel et al.,
2007; Padial et al., 2006; Pramuk et al., 2008).
PHC provide no evidence to support their
assertion that adopting scientific names in
our English Names list that delimit groups
smaller than they prefer promotes misunder-
standing more than clarity. Moreover, if
evolving taxonomies create such a funda-
mental problem, one wonders why ecologists,
cell biologists, physiologists, government
agencies, and conservation biologists, rather
than the AmphibiaTree competitors of the
Frost et al. (2006) team, are not calling for
resolution.
PHC also cite several authors for support of
taxonomic stability, which like mother and
apple pie is tough to argue against when taken
out of context. The authors cited by PHC
predictably include, among others, the pre-
phylogenetic authors George Gaylord Simp-
son and Ernst Mayr, one of the two authors of
the PhyloCode (Philip D. Cantino), and
Frost?s colleague Eugene S. Gaffney. The
Gaffney citation only tenuously supports the
PHC view and appears to hinge on a single
phrase within Gaffney?s section Classification:
Stability is ignorance (Gaffney, 1979: 103): ??I
hardly advocate change for its own sake ???
Nevertheless, Gaffney supports neither the
methods of PHC nor their nomenclatural
philosophy (e.g., see Gaffney et al., 2006). The
paper by Godfray and Knapp (2004) also does
not support the positions of PHC, even
though PHC cited it as supporting their
statement about taxonomies based on ??poorly
supported branches, or for issues unrelated to
monophyly ??? and their statement, simply
untrue, that Godfray and Knapp (2004)
supported the view that changes have
??brought the ire of many biologists ? and
only promote the increasing disregard of the
field.??
Godfray and Knapp?s (2004) paper warrants
complete and careful reading; it is a great deal
more textured than implied by PHC. A
relevant quotation (pp. 561?562, see below)
clarifies PHC?s misleading citation, provides
enormous irony, and suggests that PHC?s
proposed changes to the traditional practices
of zoological nomenclature are fraught with
new problems:
??There is intense competition for science
funding today and any field seeking new or
continuing monies is obliged both to point to
its past history of success and list the
enthusiastic end-users of its products. Taxon-
omy as currently practiced can do both, yet the
adoption of PhyloCode or similar proposals
risks both. Reinventing nomenclature suggests
that the Linnaean system of the past 250 years
is not the success we claim, but rather a blind
alley. It also risks severely alienating most
people who use taxonomy. You need only look
at how taxonomists are sometimes portrayed
by their colleagues in other fields: as scientists
who do a valuable job yet have the irritating
habit of changing names for no apparent
purpose. The wholesale abandonment of the
Linnaean naming system and its replacement
by a new and untried method would destroy
the support base for the field and imperil its
survival. And it is not as if we cannot have our
cake and eat it too. Especially today with
modern Web technologies we can link Lin-
naean taxonomies with phylogenetic hierar-
chies and have the benefits of both.??
We think that Godfray and Knapp (2004)
are correct (see also Godfray et al., 2007; Sluys
et al., 2004) and that PHC have underesti-
mated the power of the web and online
nomenclators to ameliorate ??the irritating
habit of changing names for no apparent
purpose??, but readers should realize that PHC
are the ones casting aspersions on systema-
tists, not Godfray and Knapp (2004), and that
PHC should be pointing to their past history
of success and to the enthusiastic end-users of
their products, not deriding Frost et al. (2006)
and certainly not criticizing the anuran
subcommittee of the Names list. We concur
with Godfray and Knapp?s (2004) that replac-
ing one system of nomenclature with another,
the PhyloCode (or the related approach
asserted by PHC to represent a compromise),
which has its own problems of instability (see
see Dominguez and Wheeler, 1997, and Frost
et al., 2006: 144, and footnote 26 therein), may
be ill-advised, especially if the change is
promoted through misleading statements and
misrepresentations, and contrary evidence is
ignored, as was pointedly noted by Dubois
(2006b, 2007).
June 2009] HERPETOLOGICA 139
PHC?s misrepresentation of the Godfray
and Knapp (2004) article goes further?it is an
overview article that refers to Mace (2004; in
the same journal issue) regarding the tension
between taxonomy and conservation. Mace
(2004) clearly focuses the problem of taxo-
nomic confusion at the species level, because
issues such as species boundaries, subspecies
(whatever this term may mean), and Evolu-
tionarily Significant Units may cause signifi-
cant confusion in the process of formal
conservation listings and in courts of law.
Mace (2004) makes no mention of any
possible taxonomic chaos above the species
level. Indeed, the IUCN adoption of the
generic taxonomy so criticized by PHC
indicates that the realm of conservation
embraces, rather than decries, taxonomic
efforts above the species level. Mace (2004)
makes an elegant plea for conservationists and
taxonomists to work together on matters
related to species concepts and relevant
taxonomic issues. Ironically, by presuming
they understood the basic issue at hand and
neglecting the primary literature, PHC have
nominated themselves as another example of
Ivory-Tower elitism with respect to the
increasingly crucial field of amphibian con-
servation.
In this day of web-based sources of
information (e.g., AmphibiaWeb [maintained
by D. B. Wake and supported, in part, at least
historically by the University of California,
Berkeley and in part, by NSF funds awarded
to the AmphibiaTree work-group] and Am-
phibian Species of the World [maintained D.
R. Frost and supported by the American
Museum of Natural History]), no one has to
wonder to which natural population a partic-
ular amphibian name applies. Therefore, the
justification for authoritarian nomenclatural
stability, never strong, has evaporated, and
systematists have increased freedom to ramp
up the rates of discussion, naming of taxa,
testing of hypotheses, and promotion of
research advocating new taxonomies. Indeed,
taxonomists are not confused by nomencla-
tural changes and see them for what they are,
aspects of the discussion among systematists
that promote work on various taxonomic
groups, shed light on new questions, compel
irritated systematists to write Points of View
and to defend their territories with new data
and larger studies, and respond to the notions
of what taxonomists think are useful within the
canon of monophyly.
As noted above, non-systematists in the
conservation world are not confused by
changing nomenclature, as evidenced by
IUCN?s adoption, through its Red List of
Threatened Species (http://www.iucnredlist.
org/amphibians) (also supported by Conser-
vation International and NatureServe) of the
taxonomy criticized by PHC at the first
possible opportunity (well before ITIS, and
with AmphibiaWeb as a counter-example).
Like Amphibian Species of the World and
AmphibiaWeb, the online IUCN Red List can
be searched by common synonyms (all syno-
nyms in Amphibian Species of the World)so
as to obviate any possibility of the confusion that
PHC suggest is such a serious problem. In
addition, the ongoing revolution in amphibian
taxonomy has made its way into textbooks
written by critical scientists knowledgeable
about the ongoing issues (e.g., Vitt and Cald-
well, 2009), which assures that an unfamiliar
taxonomy will not remain unfamiliar for long.
We recognize that PHC may regard Am-
phibian Species of the World and the IUCN
Red List as inherent sources of confusion,
inasmuch as having adopted the Frost et al.
(2006) taxonomy, they are not consistent with
the taxonomy employed by AmphibiaWeb,
which largely adheres to their viewpoint in
??Rana?? and ??Bufo??, although it does not
formally address subgenera nor allow for
them to be searched for explicitly. Presum-
ably, AmphibiaWeb will evolve to better meet
PHC?s approach in the near future. If so, then
we will have to question whether employing
two nomenclatural systems (ICZN, 1999;
PhyloCode) in a single database that does
not reveal to the user which system governs
what names will reduce or increase taxonomic
confusion (Sluys et al., 2004). Regardless,
suppose that Amphibian Species of the World,
AmphibiaWeb, and the IUCN Red List never
are totally in agreement. Can that be demon-
strably bad when differences of opinion are
what drive the whole scientific process (Hull,
1988)?
Regardless of the points made by PHC and
discussed more fully below, we expect that
140 HERPETOLOGICA [Vol. 65, No. 2
large genera (such as Rana and Bufo, sensu
PHC) will continue to be partitioned in an
effort to render tractable units that invite
additional work and that will guide non-
systematically-informed comparative biolo-
gists toward cogent comparisons much more
effectively than do arcane cladograms. For
recent examples, see the partition of Manti-
dactylus (Glaw and Vences, 2006); redelimita-
tion of several genera of Ranidae, including
??Rana?? (Che et al., 2007); the continuing
struggle to understand relationships within
Rhacophoridae, resulting in generic redelimi-
tations (e.g., Grosjean et al., 2008); recent
attempts to develop generic naming conven-
tions that are congruent with the age of a
taxon (e.g., Hydromantes, Atylodes, and Spe-
leomantes partitioned out as genera from
former Hydromantes by Vieites et al.,
2007?a partition supported by AmphibiaTree
funds and co-authored by one of the PIs, but
not reflected in AmphibiaWeb); and changes
in family-groups, such as the partition of the
monophyletic groups Hylidae and Microhyli-
dae based on taxon age (Bossuyt and Roelants,
2009). Partitioning has been a useful standard
practice for the entire history of systematics
and is the reason that there are more than
three amphibian genera (i.e., Rana Linnaeus,
1758, Salamandra Laurenti, 1768, and Caecilia
Linnaeus, 1758), the minimum required to
provide for named monophyletic genera for
frogs, salamanders, and caecilians, rather than
the.500 genera of living amphibians currently
recognized (Frost, unpublished data). The
putative confusion resulting from taxon parti-
tioning decried by PHC has been part and
parcel of systematics since its beginning, reflects
the growth of systematics as it deals with more
and more species, and has not presented any
credible impediment to communication.
The standard familial classification of am-
phibians prior to Frost et al. (2006) was
formulated in the 1970?s (e.g., Duellman,
1975; Heyer, 1975; Heyer and Liem, 1976;
Kluge and Farris, 1969; Laurent, 1980 ??1979??,
1986; Liem, 1970; Lynch, 1971, 1973;
McDiarmid, 1971) when the number of
species of amphibians known was less than
half of that known today. For instance, in the
1970?s Dendrobatidae contained approximate-
ly 70 species, but by the time of Grant et al.?s
(2006) revision, the number had grown to
about 250. Such growth without taxonomic
restructuring makes for unwieldy groups that
do not invite additional work. Ranks such as
subgenus are rarely encountered and easily
misunderstood by non-systematists and are
unlikely to be commonly used or reflected in
secondary literature; as a result they will have
little impact on non-systematic comparative
literature. Therefore, we think the recom-
mendation by PHC to extensively employ the
subgenus category to ameliorate the discom-
fort of name changes for English-speaking
North Americans is doomed to failure, first,
because taxonomic nomenclature is an inter-
national endeavor played on a global field
without national preference, and second,
because this use of subgenera in practice
equates to concealing diversity from the non-
systematics community.
PHC may assert that phylogenetic defini-
tion of name placement is an important part of
the answer to the problem that they perceive,
but even they have not been able to stick to
their own definitions. For example, if the
definition of Hylidae of Ford and Cannatella
(1993) and Cannatella and Hillis (2004) is
applied to our current understanding of the
phylogeny of frogs (e.g., Frost et al., 2006;
Grant et al., 2006), minimally the following
must be placed within Hylidae (to the serious
distortion of any traditional understanding of
that taxon): Aromobatidae, Bufonidae, Cen-
trolenidae, Ceratophryidae, Cycloramphidae,
Dendrobatidae, Hylodidae, Leiuperidae, and
Leptodactylidae. But, Darst and Cannatella
(2004) saw this coming and redefined Hylidae
by implication to exclude Hemiphractinae
because they showed the latter taxon to be
far from other nominal hylids. This kind of
redelimitation is necessary if one is concerned
about keeping content and diagnosis as stable
as possible with respect to the traditional use
of the term Hylidae. This one example
suggests to us that the efforts of PHC to
stabilize nomenclature through definition of
names will largely be obviated by what the
future will bring: an onslaught of new people,
new ideas, epistemological considerations, and
ever denser taxon sampling.
PHC also suggest that an extremely broad
external review of the scientific names to be
June 2009] HERPETOLOGICA 141
used in a Names list would prevent such lists
from being controversial and therefore, in
their view, make them much more useful. Of
course, being noncontroversial would militate
against any kind of change (and certainly
would retard the adoption of new approaches,
such as Phylogenetic Taxonomy, which PHC
are promoting). This is similar to the implica-
tion by Hillis (2007) who noted that the name
Rana pipiens had enormously more hits in
Google than did Lithobates pipiens and that
this was evidence of how confusion could be
propagated by name changes and critical
literature lost. However, Dubois (2007: 399?
400) provided evidence that this superficially
reasonable viewpoint, if applied for more than
a brief period would have severely negative
effects (citation dates modified to reflect our
Literature Cited):
??I had the curiosity to repeat his Google
searches and I found different, but similar
results for the same nomina (e.g., about
369 000 results for Rana pipiens versus 151
for Lithobates pipiens), but I also obtained
other interesting results with other nomina:
e.g., about 20 900 references for Rana kuhlii
versus 1130 for Limnonectes kuhlii; about
18 700 results for Rana limnocharis versus
858 for Fejervarya limnocharis;orabout
19 400 references for Rana breviceps versus
403 for Sphaerotheca breviceps. However, the
three latter species, first removed from Rana
by Dubois (1981, 1987 ??1986??, 1992), are
now universally accepted as belonging, not
only in other genera, but also in other
subfamilies (Dubois, 2005a) or even families
(Frost et al., 2006). Following Hillis? (2007)
suggestion would require to come back to the
obsolete, although long prevalent, ranid tax-
onomies of Boulenger (1918, 1920a, b), Inger
(1954, 1966, 1968) and many others. As
another example, a Google search for Tomop-
terna breviceps produced 1550 results, i.e.,
much more than Sphaerotheca breviceps, which,
to please Google users, would require ignoring
the works of Glaw et al. (1998) and Vences et
al. (2000), as well as all subsequent works,
which confirmed their main results.??
In our view the PHC viewpoint in applica-
tion would likely help retard progress in the
United States compared to the forward
motion of the rest of the amphibian biologist
community. The international community of
amphibian conservation biologists and system-
atists are getting species and monophyletic
groups named and nomenclaturally sorted
(e.g., Alam et al., 2008; Antunes et al., 2008;
Barrio-Amoro?s et al., 2008; Biju and Bossuyt,
2009; Bossuyt and Dubois, 2001; Chaparro et
al., 2008; Cruz et al., 2008; Das, 2008; Fei et
al., 2008; Gu? nther, 2008; Ko?hler et al., 2008;
Kuzmin, 2008; Li et al., 2008; Manamendra-
Arachchi and Pethiyagoda, 2005; Padial et al.,
2008) at an increasing rate compared to that of
the USA. Further, we doubt if imposing an
additional level of social inertia by English-
speaking North Americans on international
nomenclature would be seen by the interna-
tional community as positive. Would Chinese
authors who recognize Lithobates (ca. 50
species), thereby underscoring the distinctive-
ness of Rana (the Rana temporaria group and
close relatives, ca. 50 species) from its close
Asiatic relative, Pseudorana, (2 species) (see
Che et al., 2007), want their taxonomy judged
by people who have no substantive knowledge
of the systematic issues in Asian ranids? We
doubt it. Moreover, progress is rapid, due to
the increasingly easy acquisition and analysis
of large amounts of data and the availability of
rapid publication outlets such as Zootaxa.
TAXONOMIC PROBLEMS OF SPECIAL CONCERN
PHC singled out two of our taxonomic
treatments (Frost et al., 2008b) as problems of
special concern: one regarding American
bufonids and the other concerned with
American ranids. They suggested specific
nomenclatural remedies to reduce the taxo-
nomic confusion that they asserted was
created by our treatment of these groups,
which we here address in some detail. We
apologize to the general reader at this point
because to answer adequately the accusations
in PHC requires very specific and technical
responses regarding phylogenetic reconstruc-
tion and evidence (both of which PHC have
played fast and loose) and the PhyloCode.
Moreover, the questions raised by PHC
cannot be restricted to North America,
although that is PHC?s preferred arena,
142 HERPETOLOGICA [Vol. 65, No. 2
inasmuch as revisionary name changes have
global effects.
??BUFO??
First, we (Frost et al., 2008b) erred in
listing one generic name for some North
American toads. The seniority of Incilius
Cope, 1863, over Ollotis Cope, 1875 ??1876??,
and Cranopsis Cope, 1875 ??1876?? is clear and
was independently discovered by PHC, Frost
et al. (2009), and Jeff Boundy (pers. comm. to
DRF). We regret the error. However, errors
happen unless one never makes any taxonom-
ic changes?seemingly the position taken by
Pauly and Cannatella and close associates.
This was the case with ??Bufo?? in which the
obvious and documented paraphyly of ??Bufo??,
sensu lato (Graybeal and Cannatella, 1995;
Pauly et al., 2004) went unaddressed, at least
until Frost et al. (2006).
PHC?s discussion of ??Bufo?? taxonomy
revolves around a reanalysis of the Frost et
al. (2006) data purporting to provide PHC
with the basis to consider Rhinella (sensu
Chaparro et al., 2007), Incilius, and Anax-
yurus as subgenera, along with the Bufo bufo
group (Eurasian) within a redelimited Bufo.
Analysis of evolutionary tree structure rests on
(a) assumptions and concomitant operations
of analysis (e.g., transformation cost func-
tions); (b) data; and (c) selection of terminal
taxa. Minor changes in any of these three
categories can have major effects, so what
constitutes a test of a phylogenetic hypothesis
is epistemologically an important area of
discussion. In our view if an analytical
technique is being discussed, b and c must
remain constant, or be augmented, not
reduced. In this case Pauly and Cannatella
have changed all three, called it a reanalysis of
the data from Frost et al., 2006, and then
made a big deal of the differences between
the two analyses. Cannatella and Pauly?s
unpublished reanalysis is based on reducing
the amount of evidence under consideration
from that employed by Frost et al. (2006) to
just 12S and 16S mtDNA data; the exclusion
of the problematic Rhinella margaritifer
(discussed by Pauly et al., 2004); not looking
for the best overall solution of the two nested
NP-complete problems; and, seemingly not
basing their analysis on denser taxon sampling
nor including all the relevant data available
(e.g., data from Chaparro et al., 2007; Clarke,
2001; Cunningham and Cherry, 2004; Frost et
al., 2006; Graybeal, 1997; Graybeal and
Cannatella, 1995; Matsui et al., 2007; Men-
delson et al., 2005; Pramuk, 2000, 2002, 2006;
Pramuk et al., 2001; Pramuk and Lehr, 2005;
Pramuk et al., 2008; Wasonga and Channing,
2007). A reanalysis of a subset of Frost et al.?s
data and terminals does not constitute a
scientific test of the Frost et al. (2006)
hypothesis. Moreover, it is not evident how
the Cannatella-Pauly analysis extends beyond
the studies of Pramuk (2006, 2008) and
Chaparro et al. (2007), who provided far more
evidence. PHC do not explain this nor how
fewer data are better than more, particularly
given that increasingly dense taxon sampling is
needed to apprehend additional homoplasy
(Wheeler, 1992; Zwickl and Hillis, 2002).
Perhaps the Cannatella and Pauly manuscript,
which we have not seen, contains substantially
more substance than implied by PHC and we
have misunderstood the intent of their cri-
tique. We hope that their new bufonid
manuscript will address a number of genera
not sampled by Frost et al. (2006) that appear
to be nested within traditional ??Bufo?? (i.e.,
Adenomus, Altiphrynoides, Andinophryne,
Bufoides, Churamiti, Crepidophryne, Laur-
entophryne, Nimbaphrynoides, Parapelo-
phryne, Pseudobufo, and Sabahphrynus,as
well as the hugely interesting and unallocated
??Bufo?? stomaticus and ??Bufo?? orientalis
groups). The inclusion of all of these diverse
taxa within Bufo, together with substantial
amounts of new data, could significantly alter
the names of the species associated with those
genera as well as the traditional concept of
Bufo as a similarity cluster.
PHC misrepresent Frost et al. (2006)
regarding the partition of Bufo (sensu lato),
in part by not presenting the entire Frost et al.
(2006) tree of bufonids, which is reproduced
in Fig. 1. It shows in dark gray what Frost et
al. would have had to call Bufo in order to
meet PHC?s criterion of United-States-first
taxonomic stability. The difference between
PHC?s tree (their Fig. 1A) and the original
Frost et al. tree, is that an African clade
composed of Capensibufo, Stephopaedes (now
in Mertensophryne), and various groups of
June 2009] HERPETOLOGICA 143
FIG. 1.?Bufonid section of the Frost et al. (2006) tree, modified to show in light gray what taxa would minimally have
to be included within Bufo to retain all former members of Bufo within that taxon (Cranopsis of Frost et al., 2006, is now
Incilius). The dark gray polygon represents the smallest group of taxa that would have to be included to retain all North
American bufonid taxa within norminal Bufo. Note the placement of ??Bufo?? margaritifer, the type species of Rhinella,
and excluded from the reanalysis of PHC. The status of Chaunus with respect to Rhinella is covered in text.
144 HERPETOLOGICA [Vol. 65, No. 2
African ??Bufo?? (e.g., including what are now
Vandijkophrynus and Amietophrynus) in-
trudes between Bufo sensu stricto and the
American clade composed of Anaxyrus, In-
cilius, and Chaunus, with ??Bufo?? margaritifer,
far from the Anaxyrus?Incilius?Chaunus
clade in the Frost et al. (2006) tree. As noted
above, ??Bufo?? margaritifer (the type species of
Rhinella) was excluded from PHC?s rework of
Frost?s data (their Fig. 1A), an astonishing
omission that likely allowed PHC to obtain the
results they did from their reanalysis (see
discussion by Pauly et al., 2004, regarding this
terminal taxon). Had Frost et al. (2006)
wished to retain North American bufonids as
Bufo, they minimally would have had to make
Capensibufo and Stephopaedes synonyms or
subgenera, a taxonomic change that Frost et
al. (2006) did not believe would be embraced
by those working with the African taxa.
Further, Frost et al. (2006) considered
American ??Bufo??, with about 130 species, to
be unwieldy. Had Frost et al. (2006) wished to
retain all former ??Bufo?? as Bufo (as they noted
in their 2006 paper they would have had to
include every taxon they studied, as well as
those reasonably placed there on the basis of
other literature, within the lighter gray
polygon, effectively placing all bufonids other
than the atelopodines, Melanophryniscus,
Dendrophryniscus, and likely Truebella, with-
in an extremely large genus Bufo.
Beyond the peevish and arguable criticisms
of the methods employed by Frost et al.
(2006), the PHC reanalysis provides PHC
with the tree that they apparently prefer, i.e.,
a monophyletic group composed of Anaxyrus,
Incilius,andRhinella, with Bufo (sensu
stricto) as its sister taxon. They go on (Pauly
et al., 2009: p. 124) to write:
??Similarly, results from Pauly et al. (2004),
Pramuk (2006), and Pramuk et al. (2008),
indicate that (1) the New World Clade sensu
Pauly et al. (2004) is very strongly supported;
this clade includes all of the North American
species of the list; and (2) the New World
Clade is closely related to, or the sister-group
of, the Bufo bufo species group.??
We agree that the papers they cite suggest
that the American taxa Rhinella, Incilius,and
Anaxyrus form an inclusive group. We cannot
agree that all of the relevant papers (including
Chaparro et al., 2007, not mentioned by PHC)
signal that the Eurasian Bufo (sensu stricto) is
the sister taxon to this inclusive group, or even
especially closely related to it without a
simultaneous analysis of all evidence and taxa
so far presented.
The topology presented by Pauly et al.
(2004) on the basis of ca. 2.5 kb of mtDNA
suggests that minimally the sub-Saharan
African taxon Amietophrynus (38 species)
would fall into their redelimited Bufo as well.
(This is substantially similar to what would be
required in the Frost et al., 2006, tree [Fig. 1],
formulated on the basis of many more data,
assuming that Chaparro et al., 2007, are
correct in nesting Rhinella margaritifer within
former Chaunus.)
The preferred topology of Pramuk (2006),
on the basis of 4 kb of nuDNA (POMC; RAG-
1) and mtDNA (12S, tRNA
val
, 16S) as well as
83 morphological characters suggested that
the tropical Asian Phrynoidis (2 species), but
not Amietophrynus, would have to be includ-
ed in this group. Importantly, the topology
reported by Chaparro et al. (2007), but not
cited by PHC, based on about 4 kb of mtDNA
(12S mtDNA) and nuDNA (POMC and RAG-
1) suggested that Phrynoidis (2 species), the
Indonesian Ingerophrynus (11 species), trop-
ical Asian Duttaphrynus (6 species), and
African Schismaderma (1 species) would have
to be included in that group.
The only topology that corroborates the
narrow conclusion of PHC, based on a subset
of the data provided by Frost et al. (2006), is
the topology of Pramuk et al. (2008), based on
2.5 kb of mtDNA (12S, tRNA
val
, 16S) and
nuDNA (RAG-1, CXCR-4) but excluding the
morphology and one nuDNA gene (POMC)
from Pramuk (2006). Normally, we would
assume that analyses based on the most data
and with the fewest assumptions would be
best; PHC do not specify the optimality
criterion they used to judge goodness of trees.
This example illustrates PHC?s lack of
concern for ??taxonomic chaos?? in any global
sense. Rather, they seem to want to stabilize
names of United States? amphibians without
regard to effects that their action may have on
names of species on the rest of the planet.
June 2009] HERPETOLOGICA 145
Had they really been interested in maintaining
net-positive nomenclatural stability for all of
the species previously in Bufo, they would
have opted for placing all taxa that fall into the
lighter polygon into Bufo (Fig. 1) as was
previously suggested by Smith and Chiszar
(2006). That still would have necessitated
placing many taxa unfamiliar to most North
Americans and Europeans into Bufo, such was
Ansonia, Nectophryne, Didynamipus, Necto-
phrynoides, Pedostibes, Pelophryne, Schisma-
derma, and Wolterstorffina, a taxonomy that
likely would have been rejected by the
workers from Middle and South America,
Africa, and tropical Asia, outside of PHC?s
audience of concern.
The approach suggested by PHC and
putatively supported by their reanalysis of a
subset of the Frost et al. (2006) data and taxa,
requires, minimally, that Duttaphrynus, Epida-
lea, Ingerophryne, Nannophryne, Phrynoidis,
Pseudepidalea, Peltophryne, and Rhaebo re-
main distinct from Bufo in order to achieve the
narrow objective of constructing an exclusive
group named Bufo (i.e., ,BUFO.
6
), composed
of Rhinella (sensu Chaparro et al., 2007),
Incilius, Anaxyrus, and the Bufo bufo group.
If Cannatella and Pauly can demonstrate
that Rhinella, Incilius, and Anaxyrus form an
exclusive monophyletic group with Bufo (sensu
stricto), the group will contain at least 145
species (including Anaxyrus: 22 species; Bufo:
.13 species; Incilius: 33 species; Rhinella:77
species). We are skeptical that Spanish- and
Portuguese-speaking America will employ this
taxonomy, even if it is used by some Amphi-
biaTree North Americans. South American
workers will still have Nannophryne (the
former Bufo cophotis group) and Rhaebo (the
former Bufo guttatus group), so preservation of
Bufo (sensu PHC) will not meet any perceived
need in that productive academic community.
We expect them to continue to use Rhinella for
that geographically meaningful unit, likely even
partition it into more tractable monophyletic
units, which will require English-speaking
North Americans, who possess only a fraction
of the entire New Word bufonid fauna, either
to employ Anaxyrus and Incilius,orto
stubbornly retain a taxonomy that countenanc-
es paraphyly and disregards South American
scientific viewpoints.
??RANA??
The discussion by PHC of Rana and
Lithobates represents Hillis? third attempt
(see also Hillis and Wilcox, 2005, and Hillis,
2007) to convince the systematics community
that his heterodox approach to ranid nomen-
clature is compatible with both Phylogenetic
Classification (de Queiroz and Gauthier, 1990,
1992, 1994; i.e., the PhyloCode [www.
phylocode.org]) and the International Code
of Zoological Nomenclature (ICZN, 1999).
PHC do not mention that Dubois (2006b,
2007) already demolished Hillis? claims of
meaningful compatibility, nor do they refer-
ence Dubois?s papers in their Point of View,
but his criticisms are serious and have merit,
and Hillis? nomenclatural recommendations
are clearly outside of the realm of standard
nomenclatural practice. Consequently, the
anuran English names subcommittee did not
think Hillis? recommendations required dis-
cussion in the English Names list.
Moreover, PHC did not mention Che et
al.?s (2007) phylogenetic analysis and revision
of Ranidae (sensu Frost et al., 2006), which
strongly influenced discussions of ranid tax-
onomy within the anuran English names
subcommittee. In large part, that work
supersedes the results of Frost et al. (2006)
and Hillis and Wilcox (2005). Che et al. (2007)
recognized Lithobates (sensu Frost et al.)
because to do otherwise requires the distinc-
tive Chinese genus Pseudorana to be placed in
synonymy or treated as a subgenus of Rana.
Stuart (2008), studying a different gene,
subsequently confirmed this phylogenetic
result. With the exception of his coauthorship
with Bossuyt et al. (2006), in which no definite
name changes were made, Hillis has repeat-
edly focused just on the New World species to
the exclusion of the global ranid fauna.
Because the anuran English names subcom-
mittee judged that the optimal taxonomy
should be based on large, global, phylogenetic
6
We employ Dubois? (2007) method of denoting names
applied under the Phylocode and restrict the traditional
italics for those generic and species names applied under
the International Code of Zoological Nomenclature
(ICZN, 1999). Because these names are applied under
different and mutually incompatible rules, they need to be
distinguished.
146 HERPETOLOGICA [Vol. 65, No. 2
studies pertaining to the entire group, rather
than on studies of smaller geographic compo-
nents (e.g., just the Americas) we were led to
adopt the taxonomy of Che et al. (2007; which
strongly modified the Frost et al., 2006,
taxonomy, at least in the Old World), which
is reflected in Frost et al. (2008b), in the
online Amphibian Species of the World
(2009), and in the IUCN Red List?but not
in AmphibiaWeb.
The PHC-suggested subgeneric classifica-
tion (their Fig. 2A) is a direct descendant of
the taxonomy proposed by Hillis and Wilcox
(2005); we reproduce that here in our Fig. 2.
Names in that system were attached not only
to near-terminal subtaxa within American
Rana, but also to other interior stems. Hillis
and Wilcox explicitly stated (2005: 303) that
??? the clade names presented in Appendix B
are all unranked (i.e., they are not assigned to
categories such as section or subgenus)??, but
now these clade names are proposed as
subgenera (PHC?s Fig. 2). Originally these
names were to be construed, either implicitly
or explicitly, within Phylogenetic Nomencla-
ture, which is predicated on naming groups by
various methods of pointing to ancestral
stems, and thereby formulating a rule (a
??definition??) for placement of a name on a
particular monophyletic part of a larger
cladogram. In this system, groups are not
delimited by content or particular synapomor-
phies. Further, taxon-ranks, as regulated by
the International Code of Zoological Nomen-
clature (ICZN, 1999) are excluded.
Hillis and Wilcox (2005) asserted that use of
these names was consistent with the Interna-
tional Code. Dubois (2006b) disagreed citing
incompatibility in principle and also suggested
that none of Hillis and Wilcox?s (2005) new
names was available for zoological taxa under
the Code. Hillis (2007) countered with evi-
dence for the legitimacy of names coined by
Hillis and Wilcox (2005) under the Interna-
tional Code (ICZN, 1999)?except for the
ones that remained unavailable for zoological
nomenclature under even the most generous
reading of the Code (i.e., ,LAURASIARANA.,
,NOVIRANA., and ,STERTIRANA.) which
Hillis (2007) neglected to report (Dubois,
2007). Moreover, Dubois (2007) explained in
great detail that even though identical names
might be applied in both systems to taxa that
were identical in content, neither system
required this.
In response to the criticisms of Dubois
(2006b), and as noted by Dubois (2007), Hillis
(2007) formulated the taxonomy currently
accepted by PHC for North American ranids,
simply by dumping the more interior clade
names that were problematic (i.e., unavailable
under the Code: ,LAURASIARANA., ,NOVI-
RANA., ,RANULA.; or junior synonyms of
other ??subgeneric?? names employed by Hillis:
,TORRENTIRANA.5Zweifelia).
Rana has a long history of partition, from
being the only genus of frogs (Linnaeus, 1758)
to being the fourth largest group of ranid frogs
(Che et al., 2007; Frost, 2009). Surprisingly,
neither Hillis and Wilcox (2005), nor Hillis
(2007), nor PHC defined ,RANA. within
Phylogenetic Nomenclature, presumably be-
cause the bulk of Old World ??Rana?? (e.g., at
least in the last 15 years what is now
partitioned among Babina, Clinotarsus, Glan-
dirana, Huia, Hylarana, Meristogenys, Odor-
rana, Pelophylax, and Pseudorana) of other
authors (e.g., Cai et al., 2007; Che et al., 2007)
was beyond the scope of their studies. This
raises the question of whether or not PHC
also intend to include these taxa among their
subgenera in order to preserve Rana as it was
conceived of in the late 20th and early 21st
centuries. One presumes that PHC would
restrict ,RANA. to Raninae sensu Bossuyt et
al. (2006), but even this would allow most
contained groups other than Staurois, such as
the large genera Pelophylax (22 species),
Odorrana (51 species), and Hylarana (86
species), as well as the distinctive Pseudorana
(2 species), all non-North American taxa, to be
included within ,RANA.. AmphibiaWeb lists
under Rana various species that are currently
in Babina, Clinotarsus, Glandirana, some
Hylarana [e.g., Rana milneana and R. wa-
liesa], some Odorrana, Pelophylax, and Pseu-
dorana, so we suspect that the intention is to
consider these to be in Rana as well, although
the message from AmphibiaWeb is mixed and
may not reflect PHC?s intention. Fortunately,
the limits of Rana have been recently
delimited by Che et al., (2007: 12) in their
global study of ranids as the taxa subtended
(or implied to be subtended) by their Stem G.
June 2009] HERPETOLOGICA 147
(The following will unavoidably confuse read-
ers not firmly grounded in both the ICZN and
PhyloCode.) We follow Che et al. (2007) in
this determination and define the converted
taxon name ,RANA. as applying to the
smallest monophyletic group including both
Rana temporaria Linnaeus, 1758, and Rana
aurora Baird and Girard, 1852. This renders
both ,RANA. (PhyloCode) and Rana (ICZN)
subjective synonyms of Hillis and Wilcox?s
(2005) ,LAURASIARANA. (which under the
Code is a nomen nudum and an objective
synonym of Aurorana Dubois, 1992, through
the shared type species). This renders the
name Rana/,RANA. valid and of coextensive
content (sensu Frost et al., 2006) and Che et
al., 2007) under both the International Code
and the PhyloCode.
This leaves a conundrum which we do not
resolve (and which illustrates the inherent
conflict of the ICZN and the PhyloCode) with
respect to the content equivalency of Litho-
bates and ,NOVIRANA. and ,LITHOBATES.
and Zweifelia. Hillis and Wilcox (2005)
defined ,LITHOBATES. as applying to the
Rana palmipes group (now the Lithobates
palmipes group), but Frost et al. (2006) and
Che et al. (2007) applied the name Lithobates
to the largest endemic clade of American
ranids, coextensive in content with ,NOVI-
FIG. 2.?The original taxonomy tree of Hillis and Wilcox (2005) showing all taxa named by them and incorrectly
suggested to be consistent with the International Code of Zoological Nomenclature (1999).
148 HERPETOLOGICA [Vol. 65, No. 2
RANA. of Hillis and Wilcox (2005), although
this name is, unfortunately, an objective
synonym of Zweifelia Dubois, 1992, under
the ICZN.
As is obvious from the previous paragraph,
clarity in all future discussion of nomenclature
requires that workers state clearly which rules
of nomenclature they are following (e.g., see
Marjanovic and Laurin, 2008). In particular,
we agree with Dubois (2007: 400?401) that
the use of systems of nomenclature alternative
to the hugely dominant International Code
(see Frost et al., 2006: 143) should be clearly
noted in publications. That should prevent
anyone from thinking that authors have
complied with the International Code, given
that the conflation of the different systems can
result in such serious misunderstanding (Sluys
et al., 2004). We therefore endorse Dubois?
(2005b, 2006a, b, c) practice of giving all
names applied under the PhyloCode an
appearance different from that of names
applied under the International Code (e.g.,
,PANTHERANA. versus Pantherana, respec-
tively). The PhyloCode deserves careful and
reasoned consideration, and while it does have
some properties that we find attractive,
inherent stability is not one of them. Any
compromise between the PhyloCode and the
ICZN that is not based on clear principles will
result in something less satisfactory than
either. We hope that editors of biological
publications will adopt this approach as
well, thereby avoiding the taxonomic confu-
sion than can occur when alternative systems
of nomenclature use identical names in
fundamentally different ways to the detriment
of communication. Further, we encourage
Hillis and Cannatella to adopt a single
nomenclatural system for their future publi-
cations and a format for the names that
will identify that system and thereby eliminate
(or greatly diminish) the risk of any misun-
derstanding and taxonomic confusion that we
find evident in their Point of View. The
decision to embrace the International Code
of Zoological Nomenclature in subsequent
editions of Scientific and Standard English
Names lists or to abandon it for an alternative
nomenclatural system must be made by the
governing boards of the SSAR, HL, and
ASIH.
Acknowledgments.?We thank S. D. Biju, J. Campbell,
A. Channing, B. Crother, D. Cundall, R. de Sa?,D.
Diveley, J. Faivovich, M. Foster, T. Grant, L. Grismer, R.
Reynolds, and W. Wheeler for providing comments on
drafts of this manuscript under a very short deadline. Any
errors of logic or fact are ours alone.
LITERATURE CITED
ALAM, M. S., T. IGAWA,M.M.R.KHAN,M.M.ISLAM,M.
KURAMOTO,M.MATSUI,A.KURABAYASHI, AND M. SUMIDA.
2008. Genetic divergence and evolutionary relationships
in six species of genera Hoplobatrachus and Euphlyctis
(Amphibia: Anura) from Bangladesh and other Asian
countries revealed by mitochondrial gene sequences.
Molecular Phylogenetics and Evolution 48:515?527.
ANTUNES, A. P., J. FAIVOVICH, AND C. F. B. HADDAD. 2008.
A new species of Hypsiboas from the Atlantic Forest of
southeastern Brazil (Amphibia: Anura: Hylidae). Co-
peia 2008:170?190.
BAIRD, S. F., AND C. GIRARD. 1852. Descriptions of new
species of reptiles, collected by the U.S. Exploring
Expedition under the command of Capt. Charles
Wilkes, U.S.N., First Part?including the species from
the western part of America. Proceedings of the
Academy of Natural Sciences of Philadelphia
6:174?177.
BARRIO-AMORO
?
S, C. L., F. J. M. ROJAS-RUNJAIC, AND E. E.
INFANTE-RIVERO. 2008. Tres nuevos Pristimantis (An-
ura: Strabomantidae) de la sierra de Perija?, estado
Zulia,Venezuela. Revista Espan? ola de Herpetolog??a
21:71?94.
BIJU, S. D., AND F. BOSSUYT. 2009. Systematics and
phylogeny of Philautus Gistel, 1848 (Anura, Rhaco-
phoridae) in the Western Ghats of India, with
descriptions of 12 new species. Zoological Journal of
the Linnean Society 155:374?444.
BOSSUYT, F., R. M. BROWN,D.M.HILLIS,D.C.
CANNATELLA, AND M. C. MILINKOVITCH. 2006. Phylogeny
and biogeography of a cosmopolitan frog radiation: Late
Cretaceous diversification resulted in continent-scale
endemism in the family Ranidae. Systematic Biology
55:579?594.
BOSSUYT, F., AND A. DUBOIS. 2001. A review of the frog
genus Philautus Gistel, 1848 (Amphibia, Anura, Rani-
dae, Rhacophorinae). Zeylanica. Colombo 6:1?112.
BOSSUYT, F., AND K. ROELANTS. 2009. Anura. Pp. 357?364.
In S. B. Hedges and S. Kumar (Eds.), The Timetree of
Life. Oxford University Press, New York, New York,
U.S.A.
BOULENGER, G. A. 1918. Aperc?u des principes qui doivent
regir la classification naturelle des espe`ces du genre
Rana. Bulletin de la Socie?te? Zoologique de France
43:111?121.
BOULENGER, G. A. 1920a. A monograph of the American
frogs of the genus Rana. Proceedings of the American
Academy of Arts and Sciences 55:413?480.
BOULENGER, G. A. 1920b. A monograph of the South
Asian, Papuan, Melanesian and Australian frogs of the
genus Rana. Records of the Indian Museum 20:1?226.
CAI, H., J. CHE,J.PANG,E.ZHAO, AND Y.-P. ZHANG. 2007.
Paraphyly of Chinese Amolops (Anura, Ranidae) and
phylogenetic position of the rare Chinese frog, Amolops
tormutus. Zootaxa 1531:49?55.
June 2009] HERPETOLOGICA 149
CANNATELLA, D. C., AND D. M. HILLIS. 2004. Amphibians:
leading a life of slime. Pp. 430?450. In J. Cracraft and
M. J. Donoghue (Eds.), Assembling the Tree of Life.
Oxford University Press, Oxford, U.K.
CARAMASCHI, U., AND J. P. POMBAL,JR. 2006. A new species
of Rhinella Fitzinger, 1826 from the Atlantic Rain
Forest, eastern Brazil (Amphibia, Anura, Bufonidae).
Pape?is Avulsos de Zoologia. Sa?o Paulo 46:251?259.
CHAPARRO, J. C., J. M. PADIAL, AND I. DELARIVA. 2008.
Two sympatric new species of Phrynopus (Anura:
Strabomantidae) from Yanachaga Chemillen National
Park (central Peruvian Andes). Zootaxa 1761:49?58.
CHAPARRO, J. C., J. B. PRAMUK, AND A. G. GLUESENKAMP.
2007. A new species of arboreal Rhinella (Anura:
Bufonidae) from cloud forest of southeastern Peru.
Herpetologica 63:203?212.
CHE, J., J. PANG,H.ZHAO, G.-F. WU, E.-M. ZHAO, AND
Y.-P. ZHANG. 2007.Phylogeny of Raninae(Anura: Ranidae)
inferred from mitochondrial and nuclear sequences.
Molecular Phylogenetics and Evolution 43:1?13.
CLARKE, B. T. 2001. Towards a natural classification of
African toads (Anura: Bufonidae): past progress and
future prospects. African Journal of Herpetology
50:19?30.
COLLINS, J. T., AND T. W. TAGGART. 2009. Standard
Common and Current Scientific Names for North
American Amphibians, Turtles, Reptiles, and Crocodil-
ians. Fifth edition, Available at http://www.naherpetology.
org/nameIntro.asp. Center for North American Herpe-
tology. Lawrence, Kansas, U.S.A.
CROTHER, B. 2009. Are standard names lists taxonomic
straightjackets? Herpetologica 65:129?135.
CROTHER, B. I (Ed.). 2008. Scientific and standard English
names of amphibians and reptiles of North America
north of Mexico, with comments regarding confidence
in our understanding. 6th edition. Herpetological
Circular no. 37:1?94.
CRUZ, C. A. G., R. N. FEIO, AND L. B. NASCIMENTO. 2008.
A new species of Phasmahyla Cruz, 1990 (Anura:
Hylidae) from the Atlantic rain forest of the states of
Minas Gerais and Bahia, Brazil. Amphibia-Reptilia
29:311?318.
CUNNINGHAM, M., AND M. I. CHERRY. 2004. Molecular
systematics of African 20-chromosome toads (Anura:
Bufonidae). Molecular Phylogenetics and Evolution
32:671?685.
DARST, C. R., AND D. C. CANNATELLA. 2004. Novel
relationships among hyloid frogs inferred from 12S
and 16S mitochondrial DNA sequences. Molecular
Phylogenetics and Evolution 31:462?475.
DAS, I. 2008. Two new species of Pelophryne (Anura:
Bufonidae) from Gunung Murud, Sarawak (northwest-
ern Borneo). Raffles Bulletin of Zoology. Singapore
56:435?443.
DE QUEIROZ, K., AND J. A. GAUTHIER. 1990. Phylogeny as a
central principle in taxonomy: Phylogenetic definitions
of taxon names. Systematic Zoology 39:307?322.
DE QUEIROZ, K., AND J. A. GAUTHIER. 1992. Phylogenetic
taxonomy. Annual Review of Ecology and Systematics
23:449?480.
DE QUEIROZ, K., AND J. A. GAUTHIER. 1994. Toward a
phylogenetic system of biological nomenclature. Trends
in Ecology and Evolution 9:27?31.
DI CANDIA, M. R., AND E. J. ROUTMAN. 2007. Cytonuclear
discordance across a leopard frog contact zone.
Molecular Phylogenetics and Evolution 45:564?575.
DOMINGUEZ, E., AND Q. D. WHEELER. 1997. Taxonomic
stability is ignorance. Cladistics 13:367?372.
DUBOIS, A. 1981. Liste des genres et sous-genres
nominaux de Ranoidea (Amphibiens Anoures) du
monde, avec identification de leurs espe`ces types;
consequences nomenclaturales. Monitore Zoologico
Italiano. Nuova Serie, Supplemento. Firenze 15:
225?284.
DUBOIS, A. 1987 ??1986??. Miscellanea taxinomica batra-
chologica (I). Alytes. Paris 5:7?95.
DUBOIS, A. 1992. Notes sur la classification des Ranidae
(Amphibiens anoures). Bulletin Mensuel de la Socie?te?
Linne?enne de Lyon 61:305?352.
DUBOIS, A. 2005a. Amphibia Mundi. 1.1. An ergotaxon-
omy of Recent amphibians. Alytes. Paris 23:1?24.
DUBOIS, A. 2005b. Proposed Rules for the incorporation of
nomina of higher-ranked zoological taxa in the Inter-
national Code of Zoological Nomenclature. 1. Some
general questions, concepts and terms of biological
nomenclature. Zoosystema. Paris 27:365?426.
DUBOIS, A. 2006a. Incorporation of nomina of higher-
ranked taxa into the International Code of Zoological
Nomenclature: some basic questions. Zootaxa 1337:
1?37.
DUBOIS, A. 2006b. Naming taxa from cladograms: a
cautionary tale. Molecular Phylogenetics and Evolution
42:317?330.
DUBOIS, A. 2006c. New proposals for naming lower-
ranked taxa within the frame of the International Code
of Zoological Nomenclature. Comptes Rendus. Biolo-
gies. Paris 329:823?840.
DUBOIS, A. 2007. Naming taxa from cladograms: some
confusions, misleading statements, and necessary clar-
ifications. Cladistics 23:390?402.
DUELLMAN, W. E. 1975. On the classification of frogs.
Occasional Papers of the Museum of Natural History,
University of Kansas 42:1?14.
ELMER, K. R., AND D. C. CANNATELLA. 2008. Three new
species of leaflitter frogs from the upper Amazon
forests: cryptic diversity within Pristimantis ??ocken-
deni?? (Anura: Strabomantidae) in Ecuador. Zootaxa
1784:11?38.
FAIVOVICH, J., C. F. B. HADDAD,P.C.D.A.GARCIA,D.R.
FROST,J.A.CAMPBELL, AND W. C. WHEELER. 2005.
Systematic review of the frog family Hylidae, with
special reference to Hylinae: a phylogenetic analysis
and taxonomic revision. Bulletin of the American
Museum of Natural History 294:1?240. [available for
anonymous download at http://digitallibrary.amnh.org/
dspace/handle/2246/462].
FEI, L., C. YE,J.JIANG, AND F. XIE. 2008. Two new species
of the Ranidae from China, with phylogenetic relation-
ships of Hylarana (Sylvirana) nigrovittata group
(Amphibia, Anura). Acta Zootaxonomica Sinica/ Dong
wu fen lei xue bao. Beijing 33:199?206.
FORD, L. S., AND D. C. CANNATELLA. 1993. The major
clades of frogs. Herpetological Monographs 7:94?117.
FOUQUET, A., P. GAUCHER,M.BLANC, AND C. M. VE
?
LEZ-
RODRIGUEZ. 2007. Description of two new species of
Rhinella (Anura: Bufonidae) from the lowlands of the
Guiana Shield. Zootaxa 1663:17?32.
150 HERPETOLOGICA [Vol. 65, No. 2
FROST, D. R. 2009. Amphibian Species of the World: An
Online Reference. Version 5.3. (12 February 2009).
Electronic Database accessible at http://research.amnh.
org/herpetology/amphibia/index.html. American Muse-
um of Natural History, New York, New York, U.S.A.
FROST, D. R., T. GRANT,J.FAIVOVICH,R.H.BAIN,A.HAAS,
C. F. B. HADDAD,R.O.DE SA
?
,A.CHANNING,M.
WILKINSON,S.C.DONNELLAN,C.J.RAXWORTHY,J.A.
CAMPBELL,B.L.BLOTTO,P.E.MOLER,R.C.DREWES,
R. A. NUSSBAUM,J.D.LYNCH,D.M.GREEN, AND W. C.
WHEELER. 2006. The amphibian tree of life. Bulletin of
the American Museum of Natural History 297:1?370.
[Available for anonymous download at http://
digitallibrary.amnh.org/dspace/handle/2246/5781].
FROST, D. R., T. GRANT,J.FAIVOVICH,R.H.BAIN,A.HAAS,
C. F. B. HADDAD,R.O.DE SA
?
,A.CHANNING,M.
WILKINSON,S.C.DONNELLAN,C.J.RAXWORTHY,J.A.
CAMPBELL,B.L.BLOTTO,P.E.MOLER,R.C.DREWES,
R. A. NUSSBAUM,J.D.LYNCH,D.M.GREEN, AND W. C.
WHEELER. 2008a.IsThe Amphibian Tree of Life really
fatally flawed? Cladistics 24:384?395.
FROST, D. R., R. W. MCDIARMID, AND J. R. MENDELSON,
III. 2008b. Anura: Frogs. Pp. 2?12. In B. Crother (Ed.),
Scientific and standard English names of amphibians
and reptiles of North America north of Mexico, with
comments regarding confidence in our understanding.
Herpetological Circular No. 37:1?94.
FROST, D. R., J. R. MENDELSON, III, AND J. B. PRAMUK.
2009. Further notes on the nomenclature of Middle
American toads (Bufonidae). Copeia 2009:418?419.
GAFFNEY, E. S. 1979. An introduction to the logic of
phylogeny reconstruction. Pp. 77?111. In J. Cracraft
and N. Eldredge (Eds.), Phylogenetic Analysis and
Paleontology. Columbia University Press, New York,
New York, U.S.A.
GAFFNEY, E. S., H. TONG, AND P. A. MEYLAN. 2006.
Evolution of the side-necked turtles: the families
Bothremydidae, Euraxemydidae, and Araripemydidae.
Bulletin of the American Museum of Natural History
300:1?698.
GLAW, F., AND M. VENCES. 2006. Phylogeny and genus-
level classification of mantellid frogs (Amphibia,
Anura). Organisms, Diversity & Evolution 6:236?253.
GLAW, F., M. VENCES, AND W. BO
?
HME. 1998. Systematic
revision of the genus Aglyptodactylus Boulenger, 1919
(Amphibia: Ranidae), and analysis of its phylogenetic
relationships to other Madagascan ranid genera (To-
mopterna, Boophis, Mantidactylus, and Mantella).
Zeitschrift fu? r Zoologische Systematik und Evolutions-
forschung 36:17?37.
GODFRAY, H. C. J., B. R. CLARK,I.J.KITCHING,S.J.MAYO,
AND M. J. SCOBLE. 2007. The web and the structure of
taxonomy. Systematic Biology 56:943?955.
GODFRAY, H. C. J., AND S. KNAPP. 2004. Introduction. In
Taxonomy for the twenty-first century. Philosophical
Transactions of the Royal Society of London. Series B
359:559?569.
GOEBEL, A. M., T. A. RANKER,P.S.CORN, AND R. G.
OLMSTEAD. 2009. Mitochondrial DNA evolution in the
Anaxyrus boreas species group. Molecular Phyloge-
netics and Evolution 50:209?225.
GRANT, T., D. R. FROST,J.P.CALDWELL,R.GAGLIARDO,
C. F. B. HADDAD,P.J.R.KOK,D.B.MEANS,B.P.
NOONAN,W.E.SCHARGEL, AND W. C. WHEELER. 2006.
Phylogenetic systematics of dart-poison frogs and their
relatives (Amphibia: Athesphatanura: Dendrobatidae).
Bulletin of the American Museum of Natural History
299:1?262. [available for anonymous download at http://
digitallibrary.amnh.org/dspace/handle/2246/5803].
GRAYBEAL, A. 1997. Phylogenetic relationships of bufonid
frogs and tests of alternate macroevolutionary hypoth-
eses characterizing their radiation. Zoological Journal of
the Linnean Society 119:297?338.
GRAYBEAL, A., AND D. C. CANNATELLA. 1995. A new taxon
of Bufonidae from Peru, with descriptions of two new
species and a review of the phylogenetic status of
supraspecific bufonid taxa. Herpetologica 51:105?131.
GROSJEAN, S., M. DELORME,A.DUBOIS, AND A. OHLER.
2008. Evolution of reproduction in the Rhacophoridae
(Amphibia, Anura). Journal of Zoological Systematics
and Evolutionary Research 46:169?176.
GU
?
NTHER, R. 2008. Descriptions of four new species of
Choerophryne (Anura, Microhylidae) from Papua
Province, Indonesian New Guinea. Acta Zoologica
Sinica/ Dong wu xue bao. Beijing 54:653?674.
HEDGES, S. B., W. E. DUELLMAN, AND M. P. HEINICKE.
2008. New World direct-developing frogs (Anura:
Terrarana): molecular phylogeny, classification, bioge-
ography, and conservation. Zootaxa 1737:1?182.
HEYER, W. R. 1975. A preliminary analysis of the
intergeneric relationships of the frog family Leptodac-
tylidae. Smithsonian Contributions to Zoology 199:
1?55.
HEYER, W. R., AND D. S. S. LIEM. 1976. Analysis of the
intergeneric relationships of the Australian frog family
Myobatrachidae. Smithsonian Contributions to Zoology
233:1?29.
HILLIS, D. M. 2007. Constraints in naming parts of the
tree of life. Molecular Phylogenetics and Evolution
42:331?338.
HILLIS, D. M., AND T. P. WILCOX. 2005. Phylogeny of the
New World true frogs (Rana). Molecular Phylogenetics
and Evolution 34:299?314.
HULL, D. L. 1988. Science as a Process: An Evolutionary
Account of the Social and Conceptual Development of
Science. University of Chicago Press, Chicago, Illinois,
U.S.A.
ICZN. 1999, International Code of Zoological Nomencla-
ture. Fourth edition. International Trust for Zoological
Nomenclature, London, U.K. [available online at http://
www.iczn.org/iczn/index.jsp].
INGER, R. F. 1954. Systematics and zoogeography of
Philippine Amphibia. Fieldiana. Zoology 33:183?531.
INGER, R. F. 1966. The systematics and zoogeography of
the Amphibia of Borneo. Fieldiana. Zoology 52:1?402.
INGER, R. F. 1968. Amphibia. Exploration du Parc
National de la Garamba, Mission H. de Saeger.
Bruxelles 52:1?190.
KLUGE, A. G., AND J. S. FARRIS. 1969. Quantitative
phyletics and the evolution of anurans. Systematic
Zoology 18:1?32.
KO
?
HLER, J., F. GLAW, AND M. VENCES. 2008. Two
additional treefrogs of the Boophis ulftunni species
group (Anura: Mantellidae) discovered in rainforests of
northern and south-eastern Madagascar. Zootaxa
1814:37?48.
KUHN, T. S. 1962. The Structure of Scientific Revolutions.
University of Chicago Press, Chicago, Illinois, U.S.A.
June 2009] HERPETOLOGICA 151
KUZMIN, S. L. 2008. [On nomenclature of the Siberian
Newts, Salamandrella Dybowski, 1870 (Caudata: Hy-
nobiidae)]. Izvestiya Samarskogo Nauchnogo Centra
Rossiskaya Akademii Nauk 10:447?452.
LAURENT, R. F. 1980 ??1979??. Esquisse d?une phylogenese
des Anoures. Bulletin de la Socie?te? Zoologique de
France 104:397?422.
LAURENT, R. F. 1986. Sous Classe des Lissamphibiens
(Lissamphibia). Pp. 594?797. In P. Grasse? and M.
Delsol (Eds.), Traite? de Zoologie. Anatomie, Systema-
tique, Biologie. Volume 14 (Batraciens). Fascicule 1B.
Masson, Paris, France.
LAURENTI, J. N. 1768. Specimen Medicum, Exhibens
Synopsin Reptilium Emendatum cum Experimentis
Circa Venena et Antidota Reptilium Austriacorum.
Joan. Thom. nob. de Trattnern, Wien, Austria.
LEHR, E., J. B. PRAMUK,S.B.HEDGES, AND J. H. CO
?
RDOVA.
2007. A new species of arboreal Rhinella (Anura:
Bufonidae) from Zanachaga-Chemille?n National Park
in central Peru. Zootaxa 1662:1?14.
LI, J.-T., J. CHE,R.H.BAIN, E.-M. ZHAO, AND Y.-P.
ZHANG. 2008. Molecular phylogeny of Rhacophoridae
(Anura): a framework of taxonomic reassignment of
species within the genera Aquixalus, Chiromantis,
Rhacophorus and Philautus. Molecular Phylogenetics
and Evolution 48:302?312.
LIEM, D. S. S. 1970. The morphology, systematics, and
evolution of the Old World treefrogs (Rhacophoridae
and Hyperoliidae). Fieldiana. Zoology 57:vii + 145.
LIMA, A. P., M. MENIN, AND M. C. D. ARAU
?
JO. 2007. A new
species of Rhinella (Anura: Bufonidae) from Brazilian
Amazon. Zootaxa 1663:1?15.
LINNAEUS, C. 1758. Systema Naturae per Regna Tria
Naturae, Secundum Classes, Ordines, Genera, Species,
cum Characteribus, Differentiis, Synonymis, Locis.
10th Edition. Volume 1. L. Salvii, Stockholm,
Sweden.
LYNCH, J. D. 1971. Evolutionary relationships, osteology,
and zoogeography of leptodactyloid frogs. Miscella-
neous Publication. Museum of Natural History, Uni-
versity of Kansas 53:1?238.
LYNCH, J. D. 1973. The transition from archaic to
advanced frogs. Pp. 133?182. In J. L. Vial (Ed.),
Evolutionary Biology of the Anurans: Contemporary
Research on Major Problems. University of Missouri
Press, Columbia, Missouri, U.S.A.
MACE, G. M. 2004. The role of taxonomy in species
conservation. Philosophical Transactions of the Royal
Society of London. Series B 359:711?719.
MACIEL, N. M., R. A. BRANDA
?
O,L.A.CAMPOS, AND A.
SEBBEN. 2007. A large new species of Rhinella (Anura:
Bufonidae) from Cerrado of Brazil. Zootaxa 1627:
23?39.
MANAMENDRA-ARACHCHI, K., AND R. PETHIYAGODA. 2005.
The Sri Lankan shrub-frogs of the genus Philautus
Gistel, 1848 (Ranidae: Rhacophorinae), with descrip-
tion of 27 new species. Contributions to Biodiversity
Exploration and Research in Sri Lanka. Raffles Bulletin
of Zoology, Supplement 12:163?303.
MARJANOVIC, D., AND M. LAURIN. 2008. A reevaluation of
the evidence supporting an unorthodox hypothesis on
the origin of extant amphibians. Contributions to
Zoology. Amsterdam 77:149?199.
MATSUI, M., P. YAMBUN, AND A. SUDIN. 2007. Taxonomic
relationships of Ansonia anotis Inger, Tan, and
Yambun, 2001 and Pedostibes maculatus (Mocquard,
1890), with a description of a new genus (Amphibia,
Bufonidae). Zoological Science. Tokyo 24:1159?1166.
MCDIARMID, R. W. 1971. Comparative morphology and
evolution of frogs of the neotropical genera Atelopus,
Dendrophryniscus, Melanophryniscus and Oreophry-
nella. Science Bulletin. Natural History Museum of Los
Angeles County 12:1?66.
MENDELSON, J. R., III, B. L. WILLIAMS,C.A.SHEIL, AND
D. G. MULCAHY. 2005. Systematics of the Bufo coccifer
complex (Anura: Bufonidae) of Mesoamerica. Scientific
Papers. Natural History Museum, University of Kansas
38:1?27.
PADIAL, J. M., J. C. CHAPARRO, AND I. DELARIVA. 2008.
Systematics of Oreobates and the Eleutherodactylus
discoidalis species group (Amphibia, Anura), based on
two mitochondrial DNA genes and external morphol-
ogy. Zoological Journal of the Linnean Society
152:737?773.
PADIAL, J. M., S. REICHLE,R.W.MCDIARMID, AND I. DELA
RIVA. 2006. A new species of arboreal toad (Anura:
Bufonidae: Chaunus) from Madidi National Park,
Bolivia. Zootaxa 1278:57?58.
PAULY, G. B., D. M. HILLIS, AND D. C. CANNATELLA. 2004.
The history of Nearctic colonization: molecular phylo-
genetics and biogeography of the Nearctic toads (Bufo).
Evolution 58:2517?2535.
PAULY, G. B., D. M. HILLIS, AND D. C. CANNATELLA. 2009.
Taxonomic freedom and the role of official lists of
species names. Herpetologica 64:115?128.
PRAMUK, J. B. 2000. Prenasal bones and snout morphology
in West Indian bufonids and the Bufo granulosus
species group. Journal of Herpetology 34:334?340.
PRAMUK, J. B. 2002. Combined evidence and cladistic
relationships of West Indian toads (Anura: Bufonidae).
Herpetological Monographs 16:121?151.
PRAMUK, J. B. 2006. Phylogeny of South American Bufo
(Anura: Bufonidae) inferred from combined evidence.
Zoological Journal of the Linnean Society 146:407?452.
PRAMUK, J. B., C. A. HASS, AND S. B. HEDGES. 2001.
Molecular phylogeny and biogeography of West Indian
toads (Anura: Bufonidae). Molecular Phylogenetics and
Evolution 20:294?301.
PRAMUK, J. B., AND E. LEHR. 2005. Taxonomic status of
Atelophryniscus chrysophorus McCranie, Wilson, and
Williams, 1989 (Anura: Bufonidae) inferred from
phylogeny. Journal of Herpetology 39:610?618.
PRAMUK, J. B., T. ROBERTSON,J.W.SITES,JR., AND B. P.
NOONAN. 2008. Around the world in 10 million years:
biogeography of the nearly cosmopolitan true toads
(Anura: Bufonidae). Global Ecology and Biogeography
17:72?83.
SLUYS, R., K. MARTENS, AND F. SCHRAM. 2004. The
PhyloCode: naming of biodiversity at a crossroads.
Trends in Ecology and Evolution 19:280?281.
SMITH, H. M., AND D. CHISZAR. 2006. Dilemma of name-
recognition: why and when to use new combinations of
scientific names. Herpetological Conservation and
Biology 1:6?8.
STUART, B. L. 2008. The phylogenetic problem of Huia
(Amphibia: Ranidae). Molecular Phylogenetics and
Evolution 46:49?60.
152 HERPETOLOGICA [Vol. 65, No. 2
STUART, S. N., M. HOFFMANN,J.CHANSON,N.COX,R.
BERRIDGE,P.RAMANI, AND B. YOUNG (Eds.). 2008.
Threatened Amphibians of the World. Lynx Editions,
Barcelona, Spain; International Union for the Conser-
vation of Nature, Gland. Switzerland; Conservation
International, Arlington, Virginia, U.S.A.
VENCES, M., F. GLAW,J.KOSUCH,I.DAS, AND M. VEITH.
2000. Polyphyly of Tomopterna (Amphibia: Ranidae)
based on sequences of the mitochondrial 16S and 12S
rRNA genes, and ecological biogeography of Malagasy
relict amphibian groups. Pp. 229?242. In W. R. Lour-
enc?o and S. M. Goodman (Eds.), Diversite? et
Ende?e?misme a` Madagascar/Diversity and Endemism
in Madagascar. Socie?te? de Bioge?ographie, Paris,
France.
VIEITES, D. R., M.-S. MIN, AND D. B. WAKE. 2007. Rapid
diversification and dispersal during periods of global
warming by plethodontid salamanders. Proceedings of
the National Academy of Sciences of the United States
of America 104:19903?19907. [supplementary data
available at www.pnas/org/cgi/104/50/19903/suppl/DC1
#ST].
VITT, L. J., AND J. P. CALDWELL. 2009. Herpetology: An
Introductory Biology of Amphibians and Reptiles.
Third Edition. Academic Press, Burlington, Massachu-
setts, U.S.A.
WASONGA, D. V., AND A. CHANNING. 2007. Description of
the tadpole of Mertensophryne lonnbergi (Anura:
Bufonidae) from the highlands of Kenya. Salamandra
43:239?244.
WHEELER, W. C. 1992. Extinction, sampling, and molec-
ular phylogenetics. Pp. 205?215. In M. J. Novacek and
Q. D. Wheeler (Eds.), Extinction and Phylogeny.
Columbia University Press, New York, New York,
U.S.A.
ZWICKL, D. J., AND D. M. HILLIS. 2002. Increased taxon
sampling greatly reduces phylogenetic error. Systematic
Biology 51:588?598.
.Accepted: 20 May 2009
.Associate Editor: Alicia Mathis
June 2009] HERPETOLOGICA 153