997

Asiatic Herpetological ResearchVol. 7, pp. 93-102

The Karyotypes of Megophryinae (Pelobatidae) with a Discussion on theirClassification and Phylogenetic Relationships
Ding-Qi Rao And Da-Tong Yang
Kunming Institute of Zoology, Academia Sinica, Kunming 650223, China
Abstract. -The karyotypes of nine Megophryinae species from China were studied.

has 2n=26 with 5 large,

moderate and 7 small pairs. 8M+3SM+2ST. NF=52,

Megophrys omeimontis Liu

secondary constriction in 6p.
Liu
has
2n=26
with
and
8
5
small
NF=48. Megophrys
5M+5SM+2ST+1T,
Megophrys giganticus
large
pairs.
1

1

minor Stejneger has two different forms. One is 2n=26 with 6 large and 7 small pairs. 8M+2SM+2ST. NF=52;
the otheris

2n=30 with 6 large and 9 small pairs.

The large group is identical to that of the 2n=26 form.

8M+2SM+3ST+2t, NF=56. Megophrys kuatunensis Pope has 2n=24 with 6 large and 6 small pairs.
5M+2M(SM)+2SM+3ST. Megophrys lateralis (Anderson) from Tengchong. Yunnan has 2n=26 with 6 large and7 small pairs, 8M+3SM+2ST, NF=52. Megophrys lateralis (Anderson) from Hekou, Yunnan has 2n=26 with 6
large and 7 small pairs, 5M+1M(SM)+4SM+2ST+1T

Megophrys parva Boulenger has 2n=26 with 6 large and7 small pairs, 4M+5SM+1SM(ST)+3T, one secondary constriction 1Q; few metaphases with 2n=26+l and 4 pair

T chromosomes. Megophrys palpebralespinosa Bourret has 2n+26 with 5 large and 8 small pairs,
5M+4SM+1SM(ST)+2ST+1ST(T). Megophrys daweimontis sp. nov. Has 2n=26, 5 large and 8 small pairs,
6M+3SM+1SM(ST)+1ST+1ST(T), one secondary constriction in 6Q. Ophryophryne microstoma Boulenger has
2n=26 with 6 large and 7 small pairs, 7M+1M(SM)+2SM+1ST+T. The karyotype of O pachyprocta Kouofresembles that of O. microstoma.

Karyotypes of the Megophryinae can be divided into 5+8 and 6+7 types. The karyotype of M. omeimontis
appears to be an intermediate between 5+8 and 6+7. Combining the cytological, morphological and ecologicalcharacters, the classification of Megophryinae can be reviewed. The genus Atympanophrys is probable only

needed to verify its main generic character. Brachytarsophrys and Ophryophryne are valid genera. Megophrxs
can be divided into four groups: 1. M. giganticus, II. M. montana and M. nasuta, III. M. palpebralespinosa andM. daweimontis sp. nov., IV. Other species generally with small-sized bodies and 6+7 karyotypes. M. giganticus
should probable belong to the genus Atympanophrys; a new genus should be erected for groups III and IV.
whereas Megophrys should refer only to group II. A new species, M. daweimontis. is described below.

Keywords:

Megophryinae,

Brachytarsophrys.

Megophrys.

Ophryophryne.

karyotype,classification,phylogenetic relationships, Megophrys daweimontis sp. nov.

Materials and Methods

removed and the ends cut off. The marrow cells werethen washed out with

Megophrys lateralis were collected from Tengchong.
Yunnan in 1991 and Hekou. Yunnan in 1993; M. omeimontis and M. minor from Jingdong, Yunnan in June.
1990 and May, 1991; M. giganticus from Jingdong inMay, 1991; M. palpebralespinosa and M. daweimontis sp. nov. From Kekou, Yunnan in
July 1993; M.
parva and Ophryophryne pachyprocta from Mengla,
Yunnan in May 1993; M. kuatunensis from Chongan
and Dehua, Fujian;
O. microstoma from Hekou.
Yunnan in June 1992 and July, 1993. All the speci-

ground
3: 1

\

c/c

tri-sodium citrate solution,in hypotonic 0.64%

KG solution and fixed insolution of methanol;acetic acid for two periods of

20 minutes each. Slides were prepared by the air-drying

method and stained with 10% Giemsa PBS for

about 20 minutes.

Chromosomes were divided into 4 groups according to the ratio of long arm/short arm in length: M=
metacentric, the ratio is 1-1.7;

SM= submetacentric.

.

mens were brought alive to the laboratory for karyotyping.

Specimens were injected intraperitoneally withcolchicine at a dosage of 20 ug/g body-weight. After
24 to 36 hours the femur and tibia bones were

ST= subtelocentric. 3.0-7.0; T= telocentric,
more than 7.0. Chromosomes designated M, SM andST possess 2 arms and those designated T possess
1.7-3.0;

only

1

arm. The fundamental number (NF) represents

the total

arm numbers for all chromosomes in one

karyotype.

Vol. 7, p. 94

Asiatic Herpetological Research

Results

Nos. 5 and 8 are SM, Nos. 7, 9 and 12 are ST, Nos. 3

and 4 can be

The karyotype of M. lateralis from Tengchong, northwest Yunnan has 2n=26; 6 large and 7 small pairs;
and 12 are SM, Nos. 8 and 13 are ST. theNos. 3.

1

rest are

M and NF=52. There are no satellite or sec-

1

ondary constrictions (Fig. 3c). This karyotype mostly
agrees with that recorded by Wu 1987) for M. latera(lis.

The karyotype of M. lateralis (Anderson) from
Hekou. Yunnan is 2n=26; 6 large and 7 small pairs;Nos. 1.5, 6. 10 and

1

1

are M, Nos. 2. 3, 7. and 12 are

SM, Nos. 8 and 9 are ST, No. 13 is T No. 4 can be M
or

SM and NF=52.

Sometimes one secondary con-striction is close to the centromere of the short arm of

No. 2 (Fig. 3). This karyotype is different from that ofspecimens from Tengchong. Yunnan, especially inNos. 2, 7, 9, and 13, and the location of the secondary
constriction

(it

is

5p in the results of Wu. 1987).

in

We think that a morphological comparison should bemade between the two populations as part of any evaluation of their specific status.

The karyotype of M. omeimontis from Jingdong,
Yunnan has 2n=26; 5 large,
moderate, and 7 small
1pairs; Nos. 3. 7 and 12 are SM. Nos. 8 and 13 are ST.

the rest are

M and NF=52. One secondary constric-tion was near the base of the short arm of No. 6; there
is

1997

no satellite (Fig. 3d).

This karyotype agrees withby Zheng and Wu (1989) for M. omeimontis based upon a population from Mt. Omei inthat reported

Sichuan, but their results indicate that the secondary
constriction was in the long arm of No. 6.

The karyotype of M. giganticus is 2n=26; 5 large
1

3 are ST. No. 8 is

T the rest are

M and NF=48. No satellite or secondary constrictions
were found (Fig. 3e). This karyotype is not the sameas that recorded by Li et al.. ( 1993) for M. giganticus.

Recognition of 5 large and 8 small pairs is the same as
the karyotypes of Brachytarsophrys.

The karyotype of M. minor is variable in specimens from the same locality in Jingdong. One karyotype is 2n=26; 6 large and 7 small pairs; Nos. 7 and 13
are SM, Nos 8 and 12 are ST, the rest areandM

the onlyisItkaryotype where 2n=24 in the Megophryinae. Further
study is needed to evaluate the taxonomy of this species (Fig. 3j).

We found the karyotype of M. parva Boulenger tobe 2n=26; 6 large and 7 small pairs: Nos. 1, 2. 3. 4 and5 are SM, Nos. 6, 7, 8 and 10 are M, Nos.

1

1.

12 andSM and ST and

13 are T, No. 9 varies between

NF=46. One secondary constriction is near the centromere of the long arm of No.
(Fig. 3h). A few
4
with
2n=26+l
andmetaphases
pair of T chromo1

somes were found.

This karyotype

isdifferent thanthat recorded by Li et al. ( 1993) from the same localities,

especially in thearm length for large chromo-

somes, and the location of the secondary constriction(5q in the Li et al. result).

M

The karyotype of
palpebralespinosa Bourret is2n=26; 5 large and 8 small pairs; Nos. 1, 2, 4. 5 and.

12 are M, Nos. 3, 6, 8 and 10 are SM, Nos. 7 and 13

T No.

are ST, No. 9 varies between ST and

1 1

can beSM or ST and NF=52 or 50. No satellites and secondary constrictions were found (Fig. 3g).

The karyotype of M. daweimontis sp. Nov. Is
2n=26; 5 large and 8 small pairs; Nos. 1, 2, 4, 5, 9 and
13 are M. Nos. 3. 6 and 8 are SM, No. 12 is ST, No. 7

varies between ST and

T and No. 10 can be SM or ST

and NF -52 or 50. One secondary constriction is close
to the centromere of the long arm of No. 6.; This kary-

otype is similar to that of M. palpebralespinosa (Fig.
3f).

The

and 8 small pairs are obvious; Nos. 2. 3. 4. 7 and 12
are SM, Nos. 9 and

M or SM and NF=48. No satellites orsecondary constrictions were found.

KaryotypeofOphryophryne

microstoma

Boulenger is 2n=26; 6 large and 7 small pairs; Nos. 1,
2. 4, 5, 6. 8 and 10 are M, Nos. 3 and 7 are SM, No. 9
isST, Nos. 12 and 13 are T, No.

and NF=48.

1 1

can beM or SMNo satellites and secondary constrictions

were found (Fig. 3k). This karyotype is similar to the

6+7 type of some Megophrys species.
The karyotype of Ophryophryne pachyprocta Kou
resembles most closely that of O. microstoma: 2n=26;
The metaphase preparation
not good enough to provide a

6 large and 7 small pairs.
for O. pachyproctais

NF=52. No satellite or secondary constrictions were
found (Fig. 3a). The second karyotype is 2n=30; 6
large and 9 small pairs. The large chromosomes are
identical to those of the 2n=26 form, but the small
ones are different. Among them Nos. 7 and 10 are

plate here.

SM, Nos. 8, 12 and 13 are ST, Nos. 14 and 15 are T.
the rest are M and NF=56 (Fig. 3b).

Omei. Sichuan. In Table 1, its karyotypes come fromtwo populations (Mt. Omei and Mt. Wuliang) andThe karyotype of M. kuatunensis Pope is 2n=24; 6

they are very similar: all are with 5 large, one moderate and 7 small pairs.
One secondary constriction

large and 6 small pairs; Nos.

1,

2, 6, 10 and

1 1

are M,

Discussion
Although Megophrys omeimontis
uted in southwestern China,

its

is

widelytype locality

distribis

Mt.

Asiatic Herpetological Research

1997

occurs in the moderate sized No. 6.
exist.

Two differences

Zheng and Wu (1989) considered No. 6 large,

but it should be called moderate according to the relative length which they measured.

striction

is

in

a

short

arm.

The secondary conProbably the two

populations from Mt. Omei and Mt. Wuliang belong
to one species.
lationVol. 7, p. 95

the 6+7 type sinceisit

has 6 large pairs of chromo-

somes. This species only occurs on Mt. Chongan and
Mt. Daiyun. Dehua. Fujian Province, China, at theeastern edge of the distribution of the Megophryinae.

Three karyotypes occur in M. lateralis (see Table
1

).

Form

I

and

II,

based on the specimens from

Fei et al. ( 1990) recognized the popu-

Tengchong, northwest Yunnan, are generally identical,

from Mt. Wuliang as a separate species, M.

but both of them are different from Form III in chro-

jingdongensis.

mosome No. 2 and chromosome No. 13, and the loca-

The karyotype of Megophrys giganticus is of the
5+8 type, the same as those of the Brachytarsophrystion of the

Further
species and Atympanophrys shapingensis.
research is needed on the systematic positions of M.

but in the base of the short arm of No. 2 in form

giganticus and A. shapingensis.

southeast Yunnan, far

Megophrys kuatunensis is the only Megophryinae
species discovered which has 2n=24.

This karyotype

secondary constriction which

Form

III

is

Species

in

the
I,

III.

based on the specimens from Hekou.
away from Tengchong. The

taxonomic status of the Hekou population needs additional attention after comparing their morphology toother populations.

Table 1. Karyotypes of some Megophryinae species.

isterminal short arm of chromosome No. 5 in form

Vol. 7, p. 96

Species

Asiatic Herpetological Research

1997Vol. 7, p. 97

Asiatic Herpetological Research

1997

Megophrys minor from Jingdong, Yunnan has var-

We suggest that one of the large chromosomes transchromosome and theious karyotypes, but we can't find anatomical differ-ferred a fragment to a smallences between the different specimens with different

large one became small. The small one (T or ST chro-

Karyotype variation is common in Pelobatidae. such as Leptolaylax pelodytoides (Li et al..

mosome) became the No. 6. The transferred fragment

there are various karyotypes in thethe former body type in morphology because the frag-karyotypes.
1991).

same

Becausespecies,itisdifficulttoidentifyaspeciesisthe satellite on the short arm of No. 6 and it retains

ment is not lost.
lost,

distributed species, such as M. lateralis, M. omeimon-

5+8 pattern.

tis and M. minor.

very

stableIn contrast, some Pelobatidae havekaryotypes

withsimplechromosome

If thechromosomal fragment wasthen the karyotype would have resembled theaccording to its karyotype, especially for some widely

All the large sized species have 5+8 type karyotypes. While the species which are generally small

arrangements and secondary constrictions, such as
Oreolaxax, Scntiger and Yibrissaphora. The karyo-palpehralespinosa and M. daweimontis sp. Nov. Havetypes of some genera are variable, such as Leptolaylax

aand some Megophrys species.

Thus, karyotypes canbe used to resolve some taxonomic problems in some
genera, but not in others. In general, karyotypic variation in Megophryinae occur among small chromo-

somes, while the large chromosomes are relatively
invariable.

The karyotypes of Megophryinae species,

which are similar in morphology and habitats, are relatively similar. So karyotypic characters can be usedfortaxonomic purposes

to separate

genera and forevolutionary analyses.

Up to now, karyotypes of 16 species (about 2/3 of

sized

have

5+8

two karyotypicandpattern,patterns.

other

the

MegophrysandMegophrys

Ophryophryne species have a 6+7 pattern.
It is interesting that M. giganticus, Atympanophrys
shapingensis, Brachytarsophrys species. M. nasuta
and M. monticola have not only the same type (5+8)

of karyotype, but also similar morphological charactersand habitats;i.

e.

very large bodies; very large,

wide and flat heads; a hidden tympanum, round digital tips; and usually sit under stones at the edges of
streams during the breeding season (Bourret. 1942;
Inger, 1954; ^966; Liu and Hu, 1961; Tian et al..
1986; Yang et al.. 1991).

total) of Megophryinae have been obtained (Table 1).

They can be generally divided into two types on thebasis of the numbers of large and small chromosomes:
6+7 and 5+8.

Brachytarsophrys feae, B.

sp.

nov.,

Megophrys shapingensis (previously Atympanophrys
shapingensis), M. giganticus, M. nasuta, M. monticola (synonym of M. montana), M. daweimontis sp.
nov.. and M. palpehralespinosa have the 6+7 pattern,whereas M. parva, M. lateralis, M. minor, M. boetOphryophryne microstoma and O. pachyprocta
have the 6+7 pattern. Although M. kuatunensis has 6
large and 6 small pairs it belongs to the 6+7 type. M.
tgeri,

omeimontis has 5 large. 1 moderate and 7 small pairs.

Thus it seems this pattern is intermediate between 5+8
and 6+7 types. In morphology M. omeimontis is similar to species which share the 5+8 pattern.

Morescalchi (1973, 1977) thought that the 5+8
pattern was derived from the 6+7 pattern through thelose of a fragment from one of its large chromosomes(usually No. 6).

The No. 6 chromosome of M. omeimontis is peculiar (see Fig. 3d).
It seems that this chromosome is

The Classification of Megophryinae
The megophryinae is currently divided into 4 genera:
Atympanophrys, Brachytarsophrys, Megophrys, ami
Ophryophryne.
the genus

Some recent researchers think that

Atympanophrysisnot valid becauseitsmain distinguishing character (no tympanum) iswrong and this genus should be a synonym of

Megophrys (Fei et al.

1990).

A. shapingensis has aunique karyotype as well as other morphologicalcharacters. This genus has a 5+8 karyotype, relatively
large sized body, relatively flat and wide head, aden tympanum and round digital tips.
always under stones at the sides of streams.

It

hid-

occursBrachytarsophrys includes three species (Rao andYang, in press): B.feae (previously M.fcae), B. carinB. feae and theensis and a new species from China.

new species have 5+8 karyotypes. It is probable thatB. carinensis also have this kind of karyotype. All the
species in this genus have very large sized bodies,widened and flattened heads, a hidden tympanum,

easily broken at the site of secondary constriction in

shortened legs and horny-bearing eyelids. They occur
under stones at the sides of streams. In our opinion,

the short arm and becomes 5+8 if the separated part is

this genus is valid.

This hypothesis supports the conclusion ofMorescalchi with respect to the evolution of karyo-

lost.

types in Pelobatidae. The karyotype of M. omeimontis should be considered derived from the 6+7 type.

Ophryophryne has three species: O. microstoma,
O. pachyprocta and O. pollard. The former two have

6+7 karyotypes. All the species in this genus havesmall bodies, marrow mouths, small high heads, a dis-Vol. 7, p. 98

Asiatic Herpetological Research

tinct tympanum, and tubercled eyelids.

They are usu-

found on grass and leaves beside very small

ally

streams (this1997

The M. montana group
Brachytarsophrys

somewhatissimilarin cytology, morphology

toand habi-

5+8 karyotype, very large body, wide and flat
head, a hidden tympanum, straight and thick temporalunique among the Megophryinae).
This genus is valid, and it is related to the species oftats:

Megophrys which have 6+7 karyotypes.

fold, transverse shoulder groove, and short legs: but it

Megophiys is a large genus, with about 22 species.
Their body forms are varied. The karyotype patterns
differ. M. giganticus is very large, it has 5+8 karyo-

is different in

istype, an obviously wide and flat head, a hidden tym-

lacking tubercles on the upper eye-lid.

As for M. giganticus, it is narrowly distributed in
Jingdong and Yondge. Yunnan.

isItsimilar in mor-

panum, and round digital tips. The habitats and habits

phology to A. shapingensis with digital tips round,
hidden tympanum, wide and flat hear, straight andfor this species are similar to those of Brachytar-

thick temporal folds, and 5+8 karyotype.

M. giganti-

sophrys and A. shapingensis. M. montana (including
former M. monticola), and M. nasuta all have 5+8cus

and the M.

karyotypes, broad heads, relatively short legs, anindistinct or hidden tympanum similar to Brachytar-

any elongated tubercle on the upper eye-lid and notransverse groove on the shoulder. We suggest that M.

However, they have elongate free dermal

sophrys.

isdifferent than Brachytarsophrysmontana group in having much longer legs, lacking

should be placed

giganticus

in thegenus Atympa-

flaps on the snout and long soft horn-like appendages

nophrys along with A. shapingensis.

on the eyelids. Their habitats and habits are similar to

Megophrys palpebralespinosa and M. daweimonnov. are more similar to the M. lateralis groupeven though they have 5+8 karyotypes. But they have

Brachytarsophrys. The genus Megophrys was
described on the basis of the type, M. montana.

first

Megophrys palpebralespinosa and M. daweimonnov. have 5+8 karyotypes, a small sized bodytubercled eyelids, reddish inner palm tubercles, andwidened and sucker-like digital tips. They are usuallyfound on grass at the sides of small streams. The restof Megophrys. including M. lateralis, M. minor, M.
kuatunensis, M. parva and M. hoettgeri have 6+7
tis sp.

tis sp.

tubercles on the upper eye-lid and reddish inner palm
tubercles,

tympanum, no tubercles on the eyelids, and widened.

B. carinensis, B. feae and B. sp. nov.
(Rao and Yang. 1996). The type species if B. feae.

The genus Ophryophryne is valid and it contains

or roots at the sides of streams or rivers. Although M.
otneimontis seems to be intermediate between the 5+8
similar to

M. lateralisM.

parva in morphology, habitats and habits.
Thus, the species of Megophrys may be divided
into four groups according to their karyotypes,

mor-phology, habitats and habits: 1). M. montana and M.
nasuta: 2). M. giganticus; 3. M. palpebralespinosaand P. daweimontis sp. nov.; 4). M. lateralis, M. omeimontis, M. pan'a, M. minor, M. kuatunensis, M. boet-

and probably including M. uankianensis, M.
kempii, M. baluensis, M. aceras, M. brachykolos andlateralis

The genus Brachytarsophrys is valid and it contains

three species:

isfrom the M.

three species:2.

itdifferent

Based on the above information. Megophryinae isand sucker-like digital tips. They are found on stones

and 6+7 karyotype,isreviewed below:
1karyotypes, are generally moderate or small in bodysize, with regular body forms and head, a distinct

whichgroup. These species may be derived from an ancestor with 6+7 karyotype.

poilani.

O. microstoma. O. pachyprocta and O.

The type species is O. microstoma.

3. The genus Megophrys should be confined to the M.
montana group. It includes two species: M. montana
and M. nasuta. The type species if M. montana.

4. Atympanophrys should be re-named and its diagnosis

should be revised.

The genus should include M.

giganticus.
5.

A new genus Panophrys should be recognized for

the Megophrys lateralis and M. omeimontis group.

Itsomeimontis. This new genus can be

tgeri

type

M. robust a.

distinguished on the basis of small size, tympanum
present, head not very wide and flat, long legs, upper

Because of these differencesinkaryotype andmorphology, we recognize a new genus for the M. lateralis

and the M. omeimontis

Megophrys
group.
should be restricted to the M. montana and M. nasuta
group.

The M. lateralis group is generally small insize, theheads are not wide, tympanum

legs long, digital tips

flat

is

present,

and sucker-like, temporal

fold thin and bent, and a 6+7 karyotype. Frogs of thisgroup are usually found sitting on rocks next to rivers.

species is P.

eye-lids tubercles absent (except for P. palpebralespi-

nosa andP.

daweimontis). temporal fold thin andbent, digital tips flat and sucker-like.

They are usuallyfound on rocks.

As

for the relationships of this subfamily,

branches are recognized (Fig.

1):

two

one for Brachytar-

and Atympanophrys andsophrys, Megophrys,
another for Panophrys and Ophryophryne. The twoVol. 7, p. 99

Asiatic Herpetological Research

1997

E
2-

Figure 1.

Evolutionary tree of Megophrymae.

Figure 2.

Megophrys daweimontis sp. nov. female

X 1.

branches diverged early in megophryine evolutionary
history.

Both branches share almost the same distri-

Description of a New Species

have horn-like tubercles on the eyelids and the head is
In this character they

can be distin-

The Mt. Dawei specimens

have relatively long legs where the heals can overlap

when the flexed legs are held at right angles to theaxis.
The specimens have markings on thebodybacks for heads and trunks, which are different fromthose of M. brachykolos.

These markings are similar

to those similar to those of M. longipes.

According tolongipes has rudimentary webs
between the toes, no metatarsal tubercles, upper eye-Taylor (1962), M.

lids with small horn-like tubercles,of 47

and a body length

mm in males and 65 mm in females: whereas

the specimens from Mr., Dawei lack metatarsal tubercles, have smaller tubercles on the eyelids, lack webs

between the toes, and are shorter (3745

tibiotarsal articulation reaching tip ofsac
in male; toes without rudiment of
vocal
snout;

beyond jaw;

The specimens collected from Mt. Dawei, PingbianCounty, southeast Yunnan are similar to specimens ofM. brachykolos, M. parva and M. longipes., but theyguished from M. parva.

holotype.

Diagnosis: A small tubercle on outer edge of upper
eyelid; vomerine teeth present; snout projecting

bution.

not flattened.

93089) collected at the same place and time as the

mm in males andmm in females). Therefore we recognize the spec-

imens from Mt. Dawei as a separate species.

Megophrys daweimontis sp. nov. (Fig. 2)
Holotype: KIZ 93088, adult male, collected from Mt.
Dawei, Pingbian County, southeast Yunnan, China.,
altitude 1900 m.. July, 1993.
Paratypes: 17 adult males (KIZ 93069-KIZ 93085)
and 3 adult females (KIZ 93086, KIZ 93087, KIZ

web; an inner palm tubercle and very small outerpalm tubercle, a reddish inner metatarsal tubercle;
body length from stiout to vent 34-37 mm in males
and 40-46 mm in females.
Description:
Tongue pyriform; vomerine teeth
headslightly wider than long, depressed;
present;
snout very short; interorbital space concave; tympa-

num distinct, round. Arms long and slender, the first
finger extending

beyond second; palm tubercles red-dish, the inner tubercle is very large and the outer verysmall.

Legs long, the distal end of femur reachingshoulder; tibiotarsal articulation reaching tip of snout;
toes slender, swollen at tips, completely without webbing. Superarticular tubercles lacking; a reddish outer

metatarsal tubercle.

Skin smooth above, with small warts on flanks andtwo pairs of delicate oblique folds on
the scapular region extending posteriorly to waist, apair of folds on the back-side; fold extending from eye
sides of body;

above tympanum to shoulder; upper eyelid with verysmall tubercle on outer edge; lower surfaces smooth.

Olive-brown above, a triangular marking between

marking above shoulder, or X-shaped marking on the back of trunk; blackeyes, followed with V-shaped

marking beneath the vent and behind heel; transverselines above thigh: inner lower surface of legs reddish.

1997

Asiatic Herpetological ResearchVol. 7, p. 100

Measurements of holotype are: 33.5 mm snoutmm head length; 12.5 mm head width; 52.5

1973. Amphibia. Pp. 233-348 In A.

Morescalchi. A.

vent; 12.5

B. Chiarelli and C.

mm hind leg; 18 mm tibia.

and vertebrate evolution. Academic Press, New York.

Body length 33-37 mm in males and 40-

Variation:

Cytotaxonomy

Morescalchi, A., E. Olmo, and V. Stigo. 1977. Trends

Known only from type locality.

Distribution:

Capanna (eds).

46 mm in females.

of karyological evolution in pelobatoid frogs. Experientia 33( 2 ): 1 577-1 567.
1

Schmid. M., L. Vitelli, and R. Batistoni. 1987. Chro-mosome banding in Amphibia, XI.

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Asiatic Herpetological Research

1997

AppendixIlateralis (Tengchong); D. M.
Figure 3. The karyotypes of Megophryinae species: A-B. Megopkrys minor: C. M.
omeimontis; E. M. giganticus; F. M. daweimontis sp. nov.; G. M. palpebralespinosa; H. M. parva; I. M. lateralis(Hekou): J. M. kuatunensis; K. Ophryophryne microstoma.

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