Vol. 7. pp. 68-79

Asiatic Herpetological Research[997

Taxonomic Review of the Megophryid Frogs (Anura: Pelobatoidea)

Amy Lathrop
Centre for Biodiversity and Conservation Biology of the Royal Ontario Museum, 100 Queen's Park, Toronto.
Ontario, Canada M5S 2C6 and Department of Zoology, University of Toronto
recently undergone notable reorganization. Oncesubfamilies, Pelobatinae and Megophryinae. it now consists of only the former. The family

Abstract.-

The family Pelobatidae hascomposed of two
Megophryidae has

recently been elevated to represent Asian members once in the family Pelobatidae; megophryids

are diagnosedby three synapomorphies.

A phylogenetic analysis of the Archeaobatrachia by Cannatella (1985) included three species from two
the articulation
genera within Megophryidae. Paraphyly in the genus Megophrys was supported by one character,
of
the
that
alternativedata
indicatedof the urosacrum. A re-analysis of Cannatella's (1985)
relationships
patterns
were plausible and that the homology of the urosacral articulation should be examined inamong megophryids
more detail.

Key words: Amphibia, Anura, Pelobatoidea. Megophryidae, taxonomy

Noble (1922), reflected on the problems involved

Introduction
The classification of anurans has been a matter ofmuch recent debate. Several different opinions exist
about their relationships (see Ford and Cannatella.
1993 for a review of the recent work on anuran relationships), and no fewer than three different taxono-

mies are currently in use (Duellman and Trueb, 1986;

Dubois 1985, 1986; Ford and Cannatella, 1993). Previous classifications (prior to Duellman. 1975) havebeen based on a single complex of characters. As awhen alternative character schemes were
result,derived, earlier classifications were found to be incongruent.

with constructing a classification based on either asingle character or a complex of characters. However,thetrend continued(1967)

provided

until

Tihen (1965) and Inger
based on multiple

classifications

characters. Subsequently, others have adopted a similar

approach to the classification of anurans (Kluge

andFarris, 1969; Lynch, 1973; Duellman, 1975; Laurent. 1979; Dubois. 1986). More recently. Ford ( 1989,1993) undertook an analysis of the Neobatrachia, in

an attempt to identify the phylogenetic position of
Dendrobatidae among all anurans. Similarly, Cannatella (1985) generated a phylogeny for the Archaeo-choice of characters used to construct each classifica-

Both of these thorough phylogenetic
in a phylogeny for all major
culminatedanalysesand Cannatella. 1993). They
of
(Ford
frogs
groupstion (Noble [1922, 1931] vertebral centrum, pectoral

have applied contemporary techniques to generate abatrachia.

The source of thegirdle,thighincongruities have been themusculature,and dentition;

Griffiths[1963] hyolaryngeal structures, skull and limb mor-reflecting natural groupings (de Queiroz
and Gauthier. 1992). i.e., evolutionary history.

taxonomyphology, and reproduction and development, vertebral

column; Hecht [1963] andtypes).

Starrett[1973] tadpole

Each classification directly reflected the char-acters used in the particular study. For example, theproposed by Hecht (1963) in which
Orton*s (1957) larval characters were used to construct a classification that differed markedly from oneclassification

developed the same year based on adult morphology(Griffiths, 1963).

The history of these characters andtheir role in the classification of anuransisreviewedin the works of Lynch (1973) and Duellman (1975).

Anuran classification has progressed since Reig
(1958) divided the Anura into four suborders: Amphicoela, Aglossa, Archaeobatrachia. and Neobatrachia.

Based on the analysis of Ford and Cannatella 1993),
Amphicoela and Archaeobatrachia are artificialgroupings. However. Neobatrachia and Aglossa(

thegroupings (the latter now called
observaPipoidea). They confirmed Lynch *s (1973)
tion that members of Pelobatidae represent a transireflectnaturaltionalgroup

between

the

basal

anuransandAsiatic Herpetological Research

1997

Characters that support the family Pelobatidae (Duellman and Trueb, 1986), superfamily
Table 1

Neobatrachia; this implies that they possess an intermediate number of primitive character states.

.

Pelobatoidea (Ford and Cannetella. 1992) which
includes pelobatines and megophryines and thegenus Pelodytes.
ossified intervertebral discs present

procoelous centrum

Cannatella (1985) found that the family Pelobatidae. composed of two subfamilies. Megophryinae

and Pelobatinae. represented an artificial grouping,
and that each subfamily must be elevated to familial
status to remove the conflict. Currently the two families arenatella.

presacrals

I

and

II

not fusedatlantal cotyles of Presacral

Vol. 7, p. 69

placed in a superfamily, Mesobatrachia (Can1985). and is composed of pelobatoids
Pelodytidae, and(Pelobatidae.
I

closely juxta-

Megophryidae) andpipoids (Rhinophrynidae and Pipidae).

To

posed

the

date,

evolutionary

relationshipsof theMesobatrachia have been addressed only in the phyloribs absent

sacral diapophysis greatly expandedsacrum fused to coccyx (moncondylar articulation with the coccyx in megophryine and some

genetic analysis of Cannatella1993).

tella,

Within(1985. Ford and Canna-

Mesobatrachia.

the

the

relationships of Pelobatidae (Scaphiopus and Spea)

have been partially resolved by Cannatella (1985).
However, within the Megophryidae, only genealogi-transverse processes on the proximal part ofcal relationships of Scutiger (Yea et al.. 1992), andScutiger with Oreolala.x (Yang and He, 1980) has
been examined. In Cannatella"s 1985) analysis of thethe coccyx (often incorporated into the sacral

Archaeobatrachia (hereafter referred to as

diapophysis)

ease of discussion), conclusions were made regarding

eopelobatines)(

thepectoral girdle arciferal

relationships

DCC forof two genera of Megophryidae,

Megophrys and Leptobrachium. However, there werecartilaginous omosternum

errors in the coding of the characters in DCC's analy-

osseous sternum

sis that have serious implications on the compositionof the family. Consequently, herein. I will review thescapula not overlain by the clavicle

taxonomic history of the family Megophryidae, andparahyoid absent

provide a re-analysis of Cannatella's data relevant toPelobatoidea (Pelodytidae, Pelobatidae. Megophry-cricoid ring incomplete dorsallyconclude, by discussing the need for a phylogenetic classification that includes all the genera ofidae).

astragalus and calcanium are fused only proximally and distally

two tarsalia
m. sartorius is not discrete from the m. semi-

tendinosus and the end of the latter inserts
ventral to the m. gracilis
m. glutaeus magnus has an accessory tendon

Megophryidae.

Taxonomic History of Megophryidae
Themegophryid,

Megophrys montana, wasspecies of megophryids were

Boulenger's(

known at the time of1882) comprehensive account of known

amphibians. Since then, 78 additional species havebeen described, and undoubtably more will be added
as southeast Asia is more thoroughly investigated.

The(

amplexus is inguinal

first

described by Kuhl and Van Hasselt (1822). Only fourm. adductor magnus lacks an accessory head
pupil vertical

I

first

efforts

to classify

anurans was Cope

1865). He grouped taxa on the condition of the pec-and the vertebral column. In the taxonomic practices of the past, grouping of organisms

toral girdle

were based on similarity, often creating polyphyletic
type IV larvae
trigeminal and facial ganglia are fused
diploid number of chromosomes is 26 (24 in

Leptolalax pelodytoides)

assemblages of taxa. The family Asterophrydidae
Gunther 1859. a polyphyletic group, was the first family that

ily

included a megophryid. Members of this fam-included

threegeneraofmegophryids

Asiatic Herpetological ResearchVol. 7. p. 70{Megalophrys, Xenophrys, and Leptobrachium), as
well as a microhylid, and a myobatrachid.

Noble (1922, 1931) provided groupings based on
multiplecharactersofmegophryids were placed

adult

the
in

morphology;

the suborder

Anomo-

1997

Taxonomic History of the Genera
The generic taxonomy of Megophryidae suffers froma convoluted history and temporal instability ofitsgenera. The first generic name was proposed for thetype species. Megophrys montana Kuhl and van Has1822. Shortly thereafter, Wagler (1830) publishedcoela, family Pelobatidae. In these works, the family

selt

Pelobatidae included three subfamilies: Megophryi-

name as Megalophrys for the genus
This
Megophrys.
spelling continued for a century, andnae (Megophrys, Leptobrachium, Nesobia, Scutiger,
Aelurophryne, Leptobrachella), Pelobatinae (Scaphi-

thegeneric

appeared as the original generic name for seven spein monographic reviews of anuran taxonomy

opus, Spea, Pelobates. Peladytes), and Sooglossinae

cies(Nesomantis and Sooglossus). Noble (1926) believed(Boulenger, 1908; van Kampen, 1923). In his review
of reptiles and amphibians of the Malay Peninsula.

that theSooglossinae possessed

the evolutionaryallsignificant pelobatid characteristics (maxillary teeth,
arciferal pectoral girdle,

procoelous vertebrae, a sin-gle coccygeal condyle, and the particular arrangementof the pectoral muscles). However, it showed a different arrangement of the thigh musculature thought tobe derived with respect to Asian pelobatids. Not satisfied

with the characters used by Noble, Griffiths(1959) removed Sooglossinae from the family Pelo-

Smith (1930) corrected

thisunjustified

emendation

but without comment to the previous error.

From

this

genus.

singleMegophrys(Meg-alophrys). Boulenger (1882) recognized three forms
on the basis of the presence or absence of vomerine
teeth and the type of vertebrae (procoelous or opistho-

coelous).

He

split

the

group

into

three

genera:batidae and placed it in its own family, Sooglossidae.

Xenophrys (Giinther. 1865), Megalophrys (Wagler.
1830), and Leptobrachium Tschudi. 1838. Boulenger

tinae,

The classification of the Pelobatidae (viz. PelobaMegophryinae. and Pelodytinae) was maintained for nearly 40 years. Lynch (1973) removed

Megophrys because the characters that he previously
had used to separate them varied intergenerically and

Peladytes from the family Pelobatidae, and erected awere not useful for distinguishing the genera.

new family for this one genus (Pelodytidae) on thebasis of limb characteristics thatwere derived inde-

pendently from those of pelobatids.

Duellman and Trueb (1986) provided a phyloge-

later(1908)

regrouped

Ceratophrys

all

three

Gravenhorst.

into

1829.

the

genus

was used

todescribe another form of Megophrys montana. Ceratophrys. a genus of large-headed leptodactylid.
occurs exclusively in South America and bears only aresemblance

Megophrys. Other junior

netic tree that placed megophryids within Pelobatidae,

superficial

as the sister group to Pelodytidae (see Tableof synapomorphies uniting Pelodytidae and Pelobatidae). This arrangement has been the convention

synonyms of Megophrys are Ixalus (Anderson. 1871 ),once a genus of Rhacophoridae. The description of
the species was based on juvenile material ofsince Lynch's (1973) work.

Megophrys

Cannatella(

for a list

1985) performed a phylogenetic anal-

ysis of the Archaeobatrachia

batidae

1(Megophryinae

dard

Pelobatinae)

was

typic

with respect to Pelodytidae and thus
removed megophryids from Pelobatidae and erected a

new family, Megophryidae. Despitea recent effort to(Boulenger,1908);

Gorham907 placed Megophrys montana in a monogenus. Pelobatrachus. However, he did notprovide any unique characters to distinguish it from

Pelo-paraphyletic

lateralis1966) considered this species a nomen dubium. Bed-thatand found

and

(

to

the

(

1

other

)

known megophryid genera (Xenophrys.

Megalophrys.

Leptobrachium).

Therefore,

was

it

disseminate this information (Ford and Cannatella.

returned to Megophrys (Boulenger, 1908). Brachytar-

1991 ) and Fu etsophrys and Atympanophrys, were described by Tianand Hu (1983). The latter. Atympanophrys, (Greek

1993), with the exception of Truebal.,((submitted), the application has not received wide

acceptance. Recent taxonomic accounts have grouped
megophryids as a subfamily of Pelobatidae (Dubois.
1986; Duellman. 1993; Fang. 1985; Yea. et al., 1992:
Fei and Yea, 1990, 1983; Frost, 1985; Guan-Fu, et al.,

1993; Inger and Stuebing. 1991; Huang, 1985; Kuo,
1985; Wu, et al.. 1993; Zhao and Adler. 1993; andothers).

[atympano = without tympanum]), was a monotypic
genus described from a specimen of Megophrys shapingenesis in which the authors report thatallstruc-

tures associated with the middle ear (stapes, tympanic

annulus) were absent. Brachytarsophrys was thought
to be distinct because of its peculiar shaped head andhigh neural spines of the vertebrae. Dubois (1986)
reviewed the three genera, Megophrys. Brachytarsophrys. and Atympanophrys. He concluded that thethree different evolutionary states that Tian andHuAsiatic Herpetological Research

1997(19S3) referred to were neither sufficient nor important enough to deserve generic ranking and thus madethem subgenera of Megophrys. A later investigation of
the type series of Atympanophrys revealed that all

structures of the middle ear were present and that thethemiddle ear in one specimen
wasaresultof improper handling of the(SM00042)

absenceoftype (Fei et al., 1991, p. 4). The genus Atympanophrys

was considered a junior synonym of Megophrys (Fei1991 ). Brachytarsophrys is still recognized as a

et al.,

subgenus of Megophrys (Duellman, 1993).

Boulenger (1887,p.

405) reported

Amphibia,

preoccupied by an earlier name, Scutig-

1802, for a genus of centipede. As a
Boulenger erected a new generic name. Coph-era Latreille.
result,ophryne, for these forms.

presence of a deeply notched tongue, teeth, a tympanum which was concealed under the skin, and a distinct
fortheeustachiantubes).
openings
Consequently. Myers and Leviton (1962) adopted
Procter's (1922) taxonomy, but because Scutiger pre-

ceded

all

Aelurophryne.

Himalayan

megophryids

were placed in Scutiger.
Ophryophryne, a genus described by Boulenger(1903), was originally placed in the Bufonidae
because it possessed similar external characteristics(horizontal pupil and an edentate maxilla). Boulenger

that Scutiger

Theobald, 1868 (Chinese megophryids), as a genus ofisVol. 7. p. 71

He also pointed out that(1903) suggested that Ophryophryne was the evolulink

tionary

between the pelobatids and bufonids.

However, Noble (1926) examined the thigh musculature of two species of Ophryophryne and concluded

were diminutive Megophrys. Ophryophryne
remained a distinct genus, because most of the variathat they

many of the characters of Cophophryne appear to betion between Megophrys and Ophryophryneintermediate between pelobatids and bufonids (viz.
absence of maxillary teeth, expanded sacral diapophy-

head region. Dubois (1980) stated that ".

ses).

Because of these characteristics, it was consid-

ered a member of the family Bufonidae.

.

.

is

in thethe strong

resemblance between Megophrys and Ophryophryne(to the point thatitisprobably impossible to deter-mine to which group one would place the specimen

gle species, A. mammata (= Bufo mammata Giinther.

without the head!) indicates that they probably do not
merit the status of distinct genera"In this work,1896) that could be distinguished from Cophophryne

Dubois relegated Ophryophryne to the status of sub-by the absence of a posteriorly-notched tongue. These
two genera (Aelurophryne and Cophophryne) could

earlier

Boulenger (1919) erected Aelurophryne for a sin-1

be distinguished from bufonids by the presence ofvertical pupils (a character found in many pelobatids).

.

genus of Megophrys. Dubois (1986) re-examined his
comparisons (Dubois, 1980) and considered
the differences to be distinctly different, thus return-ing generic status to Ophryophryne.

Thus, Boulenger (1919) placed these two genera in
the Pelobatidae. Procter (1922) followed Boulenger'sin a1950) placed some species of Leptohrachium
new genus, Vibrissaphora. to represent a small

Liu((1919) familial arrangement. However, he consideredgroup that possessed keritanized nuptial excrescences

Cophophryne and Aelurophryne to be congeneric, andplace
Cophophryne as a junior synonym ofalong the margin of the maxilla. Liu et al. (1973)
examined members of Vibrissaphora and Leptohrach-

Aelurophryne.

ium hasseltii and found that the only difference
between the two genera was the presence of the max-

Noble (1931) recognized both Scutiger Theobald

Vibrissaphora. This character did not(= Cophophryne Boulenger) and Aelurophryne (Bou-illary spine inlenger, 1919). He distinguished the two by the pres-seem significant enough to warrant two separate gen-ence of short maxillary teeth in Scutiger and theabsence of dentition in Aelurophryne. Pope and Bor-

era.

(1940)

ing

and

Liu

(1950)

also

recognized

Aelurophryne in their surveys of Chinese amphibians.
Liu ( 950) further characterized the two genera by the1

Therefore, Vibrissaphora was returned to LeptoSome authors still1980).
(Dubois,hrachiumrecognize Vibrissaphora as a distinct genus (Zhao andAdler. 1993).

Nesobia (van Kampen. 1923) was a name given topresence of a tympanum and a small opening for theeustachian tube in Aelurophryne. and the absence of

small

these characters in Scutiger. After having re-examined

genus was not questioned by Dubois (1980) becausematerial was not available for examination. However,
because the name Nesobia was preoccupied, a genus

four species fastigiate described by Liu (1950). MyersandLeviton(1962) argued that Scutiger andcould
not be distinguished in the way of
Aelurophryne
Liu, Pope and Boring, and Noble had proposed. They

inhabitthat possess horizontal pupils andNatuna Island. The taxonomic status of thisspeciesof molluscs, a new name was applied to this group(Leptobrachella Smith 1925).

found that three of the four Scutiger forms possessed
a combination of characteristics that were supposed to
be present

in either

Scutiger or Aelurophryne(e.g..

English translation of the original text in French.

Asiatic Herpetological ResearchVol. 7. p. 72

1997

Table 2. Genera of Megophryidae and their known synapomorphies.

Leptobrachella (Borneo and the Bunguran [=Natuna]

Ophryophryne northern Vietnam and southern

Islands)

China)(

very reduced sternum*maxillary teeth absent

pointed expansion at tips of digits
ventral skin of digits sharply delimited

horizontal pupil

vomerine teeth absent

no denticles on larval oral discnuptial excrescences absent

Megophrys (southern, eastern, and southeastern Asia)*vertical pupil

tadpole with funnel-shaped oral disclarvae with median anal tube

Leptobrachium (Southern China to the Philippines,
Indochina, and Sunda Is. to Bali)

vertical pupil

vomerine teeth absent
snout not extending beyond lower jaw

Scutiger (high altitudes of southwestern China, northern Burma. Mepal, and northern India)

tongue strongly notched posteriorly

maxillary teeth reduced or absent

two well developed metacarpal tubercles

rough skinvertebrae proceolous

tympanum obscured

omosternum cartilaginous

vertical pupil

sternum with a bony style*horizontal pupilcharacter is not unique to the genus

Leptolalax (Burma and southern China through Thailand and Vietnam to Malaya and Borneo)(no unique synapomorphies)

Hu et al., erected Carpophrys (Sichuan Institute of
Biology, 1977) as a generic name for a single species,
Leptobrachium oshanensis. In a draft of the second
edition (1977) of the Chinese amphibians (Liu, et al.,1966). Liu described three species of Leptobrachiumthatwere distinct in which he placed them in a new

genus. Carpophrys.
In this document Liu included a diagnosis of Car-

closely related to the larger subset of Leptobrachium.
but distinct. Dubois 1981 pointed out that the genus()

name Carpophrys was not valid because the originaldescription was published anonymously and the typespecies for the genus was not designated (following
13b, Art. 14. International Code of Zoological
Nomenclature, 1985). Thus, he created two subgenera
Art.

for the groups—Leptolalax

for the three species, andpophrys and the type species for the genus (Leptobrachium pelodytoides). This draft was distributed

Leptobrachium for the larger subset (in the genus
Leptobrachium). Later, Dubois (1983) elevated Lep-

among Chinese colleagues and has been cited in sev-

tolalax to generic status. Feieral Chinese publications (after 1977), (personal com-gatedmunication. Jinzhong Fu. 1995). Dubois (1980)
reviewed the status of Leptobrachium and agreed with

Carpophrys

Liu that three species (L. gracile, L. pelodytoides, andL. oshanensis')

were distinct from other species ofthevalidityoftheand Ye (1992) investi-

twogeneric

names.

Leptolalax. and concluded that
although a description of the genus existed and a typespeciesandwas designated, Leptolalax is valid because

Carpophrys was not published in an official publicaInternational

Code of Zoological

Leptobrachium based on the position of the axillary
glands, appearance of the skin on the dorsum, unpigmented eggs, and other relative proportions of thetion

body. He thought these three species were likely to be

species divided into six genera (Leptobrachella, Lep-:

tobrachium. Leptolalax, Ophryophryne, Megophrys,
and Scutiger). Each genus is supported by a set ofDubois ( 1980) used the specific epithet oshanense

because as he noted (B.G. 476 in a footnote) that theoriginal genus name. Leptobrachium, is neuter in gender, and must be amended to agree with Leptolalax.

(Art.

8a. 1,

Nomenclature, 1985).
Currently, the family Megophryidae consists of 83

unique synapomorphies (Table 2) except for LeptolaMy investigation into the literature has providedlax.

no unique synapomorphies for Leptolalax. An addi-

Asiatic Herpetological Research

1997

Vol. 7. p. 73

Majority rule

Leiopelma
Discoglossidae

Leptobrachium

100

Megophrys aceras
Megophrys montana

100

100

Pelobates cultripes

67

100

Pelobates syriacus

67

100

Pelobatesfuscus

Pelobates varaldii

Spea bombifrons
Spea hammondii

100
100
100

Spea intermontana
100

Spea multiplicata
Scaphiopus couchii

100

Scaphiopus holbrookii

100

Scaphiopus hurterii
Pelodytes
Figure1.

Majority rule consensus tree depicting the relationships of Pelobatoidea. In this phylogeny,

Megophrys ismonophyletic and multiple trees result from the ambiguous relationships of Pelobates. The clade, Megophryidae isbolded.

tional genus, Oreolcdax, erectedby Myers and Levi-

ton (1962) to represent Chinese Scutiger-iike forms

possess maxillary teeth. Recently. Fei and Ye(1989) provided additional evidence to distinguish
Oreolalax from Scutiger, and this motion has beenthat

followed by some authors (Fu and Murphy, in press;

sidered to be uniquely derived: loss of most of thehyale of the hyoid and an ossified episternum. For
unknown reasons, the latter character was notincluded as a synapomorphy by Ford and Cannatella(1993).

Wu et al.. 1993; Guan-Fu et al., 1993; Fei and Yea,

Among the 83 species of megophryids, DCC
included only three species from two of the six gen-1990). Other sources do not recognize the genus Oreolalax (Dubois. 1979. 1986; Duellman, 1993; Frost,era.

1985).

available to study" (p 275). He went on to say that his

Evidence of Monophyly and OutgroupTaxa

He reported that "relationships within the mego-

phyrines are poorly defined

.

.

.

because few taxa were

results suggest that Megophrys was paraphyletic withthe single species of Leptobrachium used in his analy-

The association of megophryids with pelobatids andsis; this

Pelodytes (= Pelobatoidea of Cannatella. 1985) has
gone largely unchallenged through the history of anu-

sacrum and the coccyx). The character
was said to be a reversal to the plesiomorphic condi-

ran taxonomy. Previously recognized as Pelobatidae

tion in one species of Megophrys. Coding of this char-(authors prior to1973) or Pelobatidae plus Pelo-dytidae (Lynch, 1973), the group is supported by several synapomorphies (Table

1).

Although DCC was fairly confident about thesynapomorphies of the Pelobatoidea. themonophyletic nature of megophryids does not remain

definitiveincontestable.

The results published in his thesis list

ten synapomorphies for the group; four are reversals
to the plesiomorphic condition, four represent conver-gences with other anuran groups, and two were con-

was defined by a single character (the articu-

lation of theambiguous at best, the character state isprobably not homologous to the primitive condition
found in discoglossids and Ascaphus. Cannatella
acter

is

alluded to this problem

inhis section

on character

analysis (p. 127).

Becausetheresultsof

DCC's

analysishavebrought into question the monophyly of Megophrys, I
re-investigated the issue.

Vol. 7, p. 74

Asiatic Herpetological Research

1997

Leiopelma
Discoglossidae

Leptobrachium

Megophrys aceras
Megophrys montana
Pelobates culthpes
Pelobates syriacus
Pelobates varaldii
Pelobatesfuscus

Spea bombifrons
Spea hammondii
Spea intermontana
Spea multiplicata
Scaphiopus couchii
Scaphiopus holbrookii
Scaphiopus hurterii
Pelodytes
Figure 2. Phylogeny of Pelobatoidea. The relationships of Pelobates is unresolved. The character UROSACRM
when evaluated as ordered with its original four states (Cannatella, 1985) constrains Megophrys to be paraphyleticwith Leptobrachium. The clade. Megophryidae is bolded.

Materials and Methods
I

coglossus, the sister species of the clade Pelobatoidea.

examined the data set of Cannatellato(1985) in orderunderstand the evolution of the morphologicalcharacter within pelobatoids. My evaluation of the
taxa in the clade Pelobatoidea (viz. Leptobrachium,

and Leiopelma as a secondary outgroup.

Results
Three most parsimonious trees (MPTs) were resolved

Megophrys. Pelobates, Pelodytes, Scaphiopus. andSpea) included 43 of 181 characters for 17 of 42 taxa

when the data were run as unordered (65 steps; CI -of archaeobatrachians. Two characters associated withmonophyletic, with Leptobrachium as its sister group.
Differences among the MPTs occurred within the

COLUMEL1 (0, absent; 1, present; 2,greatly elongated) and COLUMEL2 (0, normal size;
thestapes:reduced) were combined to removed inapplicablecharacter states. The character
was coded
1.

COLUMEL

as follows (0. absent;1.

present, normal, extending.723; RI = .860; Fig.

clade of Pelobates.

1).

In all cases, Megophrys wasCharacter states for Pelobates

were invariant, except for the COLUMELLA and the

CRICOID, both of which were autapomorphic for P.
veraldiiand P. fuscus. respectively. These autoapo-

reduced). Uninformative characters at this level of themorphies were responsible for the differing topologies
among Pelobates. When all the characters wereanalysis were removed; of thetreated as ordered, as in theentire length of ear cavity; 2, greatly elongated; 3,181

characters.

137

characters were not informative in the analysis of

Pelobatoidea(Appendix I). Phylogenetic analyses
were performed using Swofford's (1993) Phylogenetic Analysis Using Parsimony program (PAUP). A
heuristic search was performed using simple stepwise
addition only minimal trees were retained and trees
were constructed using branch swapping option oftree

bisection-reconnection.

steepest

descent,

andholding all most parsimonious trees. Characters wereordered and polarized by outgroup comparison(

Watrous and Wheeler, 1981) using the outgroup Dis-

DCC analysis, six MPTs

were resolved, each 73 steps long (CI = .699; RI =.850).

Three of trees were identicalunordered analysis (Fig.
topologies,

Megophrys

1).

isto those in the

Among the other threeparaphyletic with Lepto-

brachium. and the relationships of Pelobates were notresolved (Fig. 2).

One character used by DCC was the articulationof the sacrum and the coccyx (UROSACRM).

DCC

had doubts about the polarity and homology of thischaracter. Although similar in configuration, he ques-1

Asiatic Herpetological Research997

The systematics of megophryids is poorly under-

tioned whether the condition in Megophrys montana

was homologous

Vol. 7. p. 75

stood. Our ideas of the relationships within the familyof Leiopelma. Cannatella
coded this character as homologous, but with reserva-

have not changed much since Noble's (1926) evolu-

When the character was either left unordered ortionary hypothesis. Without a comprehensive phylo-

tion.

to

that

genetic

ing three topologies were identical to those Fig.

prove to be unstable;

the articulation of thesacrum1.

in

Megophrysmontana is not homologous to that of Leiopelma, andIfdifferent

from that seen in Megophrys aceras, then 5character states are required to describe the variationin

UROSACRM.

rather than the original 4 states as in1)

the analy-

theresulting

we can basetaxonomywillalikelyit
may include paraphyletic
groupings. Although Dubois (1980) constructed aclassification to serve as a starting point for futurestudies, his conservative scheme is not based on aphylogeny, but primarily on phenetic clusterings without regard for evolution of the characters.

DCC. When UROSACRM was coded as five orderedor unordered character states (Appendix

hypothesis

classification,

which

from

removed completely from the analysis. Megophrys
was resolved as a monoprn letic group and the result-

To understand the relationships between the genera

of

and

Megophryidae,

theplacementof

sis resulted in three trees identical to those of the pre-megophryids within the pelobatoids. a thorough phy-

unordered analysis of the original data set(ordered = 73 steps; CI = .712: Rl = .854; unordered =logenetic analysis of the family

analysis should be performed in a similar manner as

66 steps; CI = .727; RI = .859).

Cannatella's

vious(1985)

study,

andisnecessary. This

theclassification

should be derived directly from the phylogeny (Wiley.

Discussion
DCC'sphylogeny

depicts

the

two

of

species

Megophrys
paraphyletic with respect to Leptobrachium. However, my re-analysis demonstrates that
as

there

is

Megophrysanalternative

arrangement

in

which

monophyletic. The condition of theis

UROSACRUM (e.g.. syncondrotic or synovial, monocondylar or bicondylar) has been used as a diagnostic
character for some groups of anurans (Nicholls, 1916;

1981 ). Employing a phylogenetic classification would
ensure that the taxonomy was consistent, functional,

and maximized information content. In this way, theknowledge that is gained from this interesting groupof frogs will produceausefulclassification

thatclearly reflects the evolutionary paths of each species.

AcknowledgmentsI

am grateful to J. R Bogart. for his loan of literature

Lynch. 1973; Cannatella. 1985: Ford.
1989) and avoided by others (Duellman and Trueb.

on short notice. R. MacCulloch helped translate theFrench. J. Fu translated, discussed, and clarified much

Noble (1922) cautioned against its utility as aof the Chinese literature that would still continue to bediagnostic character for pelobatids (Megophryidae.
Pelobatidae. and Pelodytidae). Furthermore, withina mystery to me. C. Huang was kind enough to allow

Griffiths. 1963;1986).

Megophryidae, specifically Megophrys, there has

thebeen several reports of variation in the urosacral
fusion (Kluge, 1966). Thus, because UROSACRM
has the ability to make Megophrys paraphyletic, theme to use her draft translation of Fei et al.,'s (1991)
am grateful to S.
Key to Amphibians of China.
I

Monks, R. Murphy, and D. Upton for their helpful
suggestions which have improved the quality of thispaper. This study was supported by the Natural Sci-

articulation of the coccyx to the sacrum must be
examined more closely to determine its true homol-

ences and Engineering Research Council (NSERC) of

ogy.

is

Wiens (1989) illustrated the difference in development between a fused bicondylar articulation and one
that

is

not fused, and suggested that these arrange-ments may have arisen via several different developmental pathways.

What appears to be two different

conditions

Megophryidae

style

in

the

separate;joint—sacrum andsynchondrotic

in

uro-

Megophrysmontana and Leptobrachium hasseltii; sacral fusion—

Megophrys aceras
may have been formed fromtwo very similar developmental pathways. Ontogein

netic studies of the formation of thesacrum and thecoccyx may reveal homologous states that would not
be identified by examining just the adult condition.

Canada Grant A3 148 awarded to R. W. Murphy. This
contribution XXX from the Centre for Biodiversity

and Conservation

Museum.

Biologyof theRoyal

Ontario

Asiatic Herpetological ResearchVol. 7. p. 76

Appendix

1997

I

Data matrix of characters from Cannatella 1985) to include only Pelobatoidae and the outgroup taxa, Leiopelma
and Discoglossus. (A) Leiopelma, (B) Discoglossus pictus, (C) Leptobrachium. (D) Megophrys aceras, (E) M.
(

montana, (F) Pelobates cultripes, (G) P. syriacus. (H) P. varaldii, (I) P. fuscus, (J) Spea bomifrons, (K) S. hammondii, (L) S. intermontana. (M) S. multiplicata. (N) Scaphiopus couchii. (O) S. holbrookii. (P) S. hurterii. (Q)
Pelodaytes punctatus. See original work (Cannatella. 1985) for character coding.

characters

ADDLONG

taxa

ABCDEFGHI

J

KLM

1997

Asiatic Herpetological ResearchVol. 7. p. 78

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