Vol.9, pp. 77-100

Asiatic Herpetological Research

2001

A Review of the Distribution of Diploid, Triploid andTetraploid Green Toads{Bufo viridis Complex) in Asia Including New Data from Iran and Pakistan

Matthias Stock

1,

Daniel Frynta 2 Wolf-Rudiger Grosse
,

1
,

1

Claus Steinlein 3 andMichael Schmid 3

Martin-Luther-Universitdt Halle-Wittenberg, Institutfiir Zoologie, Domplatz 4,

Germany, e-mail: stoeck@zoologie.uni-halle.de.

128 44 Praha, Czech Republic.

*

'

D

-

06099 Halle/ Saale,Charles University, Department of Zoology, ViniCnd 7,

Universitat Wiirzburg, Institutfiir Humangenetik, Biozentrum,D-

Am Hubland,

97074 Wiirzburg, Germany.

A summary of the distribution of green toads containing most of the data published after the discoveryof polyploid forms (1976) including a map, an index and a bibliography are presented and discussed. 21 Asian

Abstract.-type localities of hitherto described nominal green toad taxa are shown. The tetraploids are distributed in highmountains and extremely continental regions with strong climatic shifts. Records of triploid specimens are
situated in supposed contact zones

between the parapatric diploid and tetraploid toads at foothills of Middle

Asian high mountains, and triploid bisexual populations occur in the Karakoram and West-Himalayas. Habitatsof diploids appear to be restricted to lowlands and valley grounds. Polyploids seem to be more resistant. The

methods hitherto used for the determination of the ploidy level and their applicability are evaluated. We show

new data on this species complex from Iran including cytometric, karyological, bioacoustic and morphological
data and we draw taxonomic conclusions for tetraploid Bufo oblongus, diploid Bufo viridis kermanensis, andprobably diploid Bufo kavirensis. New information on the distribution of triploids in northwestern Pakistan basedon flow cytometric measurements is presented. The ploidy level of Bufo latastii is revealed to be diploid.

Key words.- Amphibia, Bufonidae, Bufo viridis complex, Bufo oblongus lectotype, Bufo viridis kermanensis,
Bufo kavirensis, Bufo latastii, Bufo pseudoraddei pseudoraddei, Bufo pseudoraddei baturae, distribution, type
localities, Asia, Iran, Pakistan, chromosomes, ploidy determination, calls, systematics

This paper is a chapter of the doctoral dissertation of M. Stock.

Hence, the present paper was aimed to summarize theIntroduction

recent information about the distribution which is one

Since the discovery of tetraploid forms of the Bufo
viridis complex in the northern Tien Shan (Bachmann
et al„

1978; Mazik et al., 1976) many records of dip-

and tetraploid green toads have been
made, especially on the territory of the former Soviet

loid,

triploid

Union and some few in other countries. New findings
as the detection of possibly all-triploid, gonochoristic

populations of this complex in the Karakorum range ofPakistan (Stock et al., 1998, 1999) provide argumentsthat thenumber of investigations in this species com-plex will increase during the next years because not
only zoogeographic and phylogenetic but also cytoge-

and biochemical questions will be
studied. Since a fundamental summary of the data on
netic, bioacousticof the preconditions for the understanding of the natural historyof these toads. The data on chorology will

also be an important prerequisite for a revision of sys-

tematics and nomenclature in this species complex.

We discuss the hitherto applied techniques ofploidy determination, the size of the (known) range ofpolyploid green toads, any detectable correlation of
their occurrence with climatic/ecological factors, andwe try to enlarge the knowledge about the ploidy levelof taxa hitherto described from various type localities.

Because the knowledge on the ploidy of green
toads from the territory of Iran and Pakistan is very
scarce, we also present and discuss some new, sporad-the territory of the former Soviet Union was done byically collected data from different sources,

Borkin et al. (1986a), the development of knowledge inresults of a field excursion to Pakistan.

this species complex

has been dramatically increased.

and first

Vol. 9, p. 78

Figure 1.

Asiatic Herpetological Research2001

Map of Middle and Central Asia and parts of the Middle East with records of diploid, triploid and tetraploidgreen toads after 1976. For code numbers see appendix.

2001

Asiatic Herpetological ResearchVol. 9, p. 79

Vol. 9, p. 80

20

175
IS

12.5

10

Asiatic Herpetological Research

2001

2001

Asiatic Herpetological ResearchVol. 9, p. 81

Asiatic Herpetological ResearchVol. 9. p. 82

Table 1.

2001

DNA content in diploid toads from Kerman (Bufo viridis kermanensis), diploid toads from Choqua Zanbil(B. wridisssp.), tetraploid toads from Birjand [Bufo ob/ongus), and diploid B. iatastiiUom Skardu measured by flow

cytometry of DAPI stained erythrocytes with chicken nuclei (= 2.34 pg DNA/nucleus) as a standard. SD
deviation.

Kerman, diploid= standard

Asiatic Herpetological Research

2001

35

30

n 25
<D

*»

n
a>

20

v>

3
a.

15

10

Vol. 9, p. 83

Asiatic Herpetological ResearchVol. 9, p. 84

2.31, Borkin et al., 2000). Including the data shown

above, we preliminarily consider (diploid) B. kavirensis tobe a junior synonym of diploid B. viridis ker-

2001

regarding neighboring races" (Eiselt and Schmidtler.

1973) were now explained by the different ploidy levels. The

females of B. oblongus (e.g., Fig. 8B) examfrom Birjand exhibited a coloration patternmanensis. This also agrees with the range presumed

inedby Eiselt and Schmidtler (1973) for their taxon. The
relationships between diploid B. viridis kermanensis

which is very similar to that of the specimen ZISP

and diploid B. viridis ssp. (previously called B. viridis

designatearabicus, see above) require additional investigations.

Nikolsky, 1896, and we consider this taxon according
to the topotypes to be tetraploid. This demonstration

Themorphologicaldifferences

betweenBufo

oblongus (Fig. 8A, B, D) and B. viridis kermanensis(Fig. 8E) which were considered to be "very sharp

1952.1 from the type series (Fig. 8A). We therefore

onomy of green toads in Asia because the name is

en

Q.

« 2

£
- 3

o3

CD

Source

-Q

«> 9

•w

o

v-/

to

.§

«a
of "5

> I
* E"
S
CO

Intercall interval (s) [N of intervals measured]? -D.o .9-

LU(0

Call duration (s) [N of calls measured]^E 2Q.
O
,

If)

oi

M

Fundamental frequency (Hz)!<§

Pulse rate (s' )

-a-

1

<5
CM

"
CD

5 S
E «
•a co

E

>o 5.

Pulse duration (ms)Interpulse interval

(ms)

^= x:
e/>

"S

O

£
E

WaterT ("C)

0)

£?.
"O
O)

°!
Es
2 ^

AirT

CO

Male

*1
° o

Locality (code in Fig. 1)

ahere as the lectotype of B. oblongus,

of tetraploidy for B. oblongus is important for the tax-

El
<S

it

^

2001

Asiatic Herpetological ResearchVol. 9. p. 85

Table 4. Mating call data of triploid Bufo pseudoraddei baturae'xn comparison with Bufo /atasti/Uom India andPakistan.

Asiatic Herpetological ResearchVol. 9, p. 86

2001

Table 5. Evaluation (overview) of methods previously used by different authors for the determination of diploid andpolyploid green toads. For detailed information see text.

Method of determination

Unequivocalness

Applicability in the field

Abbr. in the appendix

Karyotype, karyogram, counting of

yes

no

chrom. count,no, but blood samples canflow cytom.

chromosomes
Flow cytometry

yes, if applied professionallyshortly be stored in ethanol

Microdensitometry (Feulgen stain-

yes, if applied professionallyyes, if blood smears are
stored informaldehyde

microdens., cytom.

5%ing), cytometry

solution

Electrophoresis

no, only prognosis possible

Measurement of erythrocytes

no, only prognosis possible

electrophor.
collection of air dried blooderythr. size

samples easily possible
probably yes, for distinguish-

Mating call analysis

yes

calls

yes

ext. morph.

ing between diploid and polyploid (triploid. tetraploid)

forms, but see B. latastii
External morphology, morphometryno, only prognosis possible

Form and size of the clutchno, often misleading

Evaluation of methods for the identificationand determination of the ploidy in Asian

green toads (Table 5)Some forms/speciesof Middle and Central Asian

Grosse, 1997a). Although better results can be obtained

blood causes a loss of stainability, we recently got goodresults withblood smears which were storedin

5%

formaldehyde solution until staining (our unpublisheddata). The obtaining of blood smears from the tip of theGrosse, 1997a).

The karyological analysis still remains an essential tool. The treatment with colchicine and the prepa-

finger seems to be of very little damage for the animals,

but requires to prevent the pollution of blood samples

with skin mucus.

marrow, spleen or intestine tissue forthe counting of stained

chromosomes (e.g., Schmid,

1978; Macgregor and Varley, 1983) requires the killing of the animals. The obtaining of blood by heart

puncture with micro-syringes (e.g., Schroer, 1996)
and the determination of the ploidy by chromosome

counts from blood cultures (Castellano and Giacoma,

1998) causes little mortality but requires the transportof the toads to the laboratory.

Flow cytometry is another unequivocal technique(Murphy et al., 1997 for review). The preferable storage of blood by freezing in liquid nitrogen is usuallylimited in the field. Ethanol fixation of blood obtained

from anaesthized toads allows a storage for someweeks and the ploidy determination by flow cytometry if refrigeration is possible (e.g.. Stock et al., 1999),

but does not allow the exact determination of absoluteand theerythrocytes
microdensitometrical determination of their DNAstaining

Significant differences in the average erythrocyte
size allow a prognosis of the ploidy level (Stock andDiscrimination
1997a).
problems occur
because some populations exhibit values in the over-

Grosselapping range between diploid and tetraploid toads. In
the classification of triploid individuals

addition,

remains unsolved. In probably all-triploid populations

from Karakoram, the mean erythrocyte areas exhibited significant differences to diploid and tetraploid
populations (Stock et al., 1999), but the variability of
the individual erythrocyte size shows a large overlap-

ping with diploid and tetraploid specimens. Unfortunately, these phenomena also prevent an exact
analysis of the ploidy level of museum and type specimens with the technique of Mercadal (1981).

Results of protein electrophoresis (e.g., Borkin

and Sokolova 1989, Mezhzherin and Pisanets, 1995a,

DNA-contents.
Feulgencontent provides also unequivocal results (Stock andif applied in the laboratory since the storage of air dried

green toads have been partly identified using misleading or ambiguous methods (comp. also Stock andration of bone

clutchof

1995b) can usually not substitute the preceding ploidy
determination since "electromorphs" may cause misinterpretations. Some authors of electrophoretic stud-2001

Asiatic Herpetological Research

Figure 8. A-

Vol. 9. p. 87

Lectotype (present designation) of Bufo ob/ongusN\ko\sky, 1896 (Z ISP 1952.1) from Birjand, E-lran.

B Topotypic tetraploid female Bufo oblongus\xom Birjand. C Diploid male from Gholaman. D Topotypic tetraploid male Bufo ob/ongusUom Birjand. E
Topotypic diploid male of Bufo viridis kermanensis Eiselt & Schmidtler,
1971. F Tetraploid male from Baghestan. G - Holotype of Bufo kavirensis, Andren & Nilson. 1979. H Holotype of
-

-

-

-

-

Bufo luristanicus Schmidt, 1952.

-

describe whether or how they determinedies did not

the

level

ploidy(e.g..

Mezhzherin and

Pisanets.

At any given temperature mating calls of tetraploid toads exhibited longer pulses and interpulse inter-vals resulting in lower pulse rates than diploid toads(Stock 1997a. 1998a). a finding which exhibits parallels in other diploid/tetraploid anuran species (Hyland artificial tetraploids
idae,
Leptodactylidae)

suggesting that differences are caused by polyploidy(Gerhardt. 1994: 317).

For triploid toads from northern Kyrgyzstan (Casnorthern

raploid toads from Central Asia inmany characterswith univariate methods (Stock et al., 1999). A prognosis for the purely morphometric classification of

1991).

tellano et

2001

Asiatic Herpetological ResearchVol. 9, p. 88

1998) and triploid populations from
Pakistan (Stock et al.. 1998. 1999) callal..

diploid, triploid and tetraploid toads is possible withmultivariate

methods and this appears to be one tool

for the prognostic assortment of preserved specimens
in collections. The prognosis can be improved if mor-phometric data are combined with erythrocyte size
data (Stock, 1997b; Stock and Grosse, 1997a; Stock etal.,1999). In any case, the use of exact methods for

ploidy determinationishighlyrecommended(e.g.,

Borkin et al.. 2000).
Pisanets (1987) indicated differences in form andsize of the clutch between diploid and tetraploid toads

parameters were found to resemble those of tetraploidtoads. Therefore, the mating call analysis seems suit-able for distinguishing between diploid and polyploid(triploid. tetraploid)

green toads. However, indepen-dently from polyploidisation. speciation in the B. viri-

complex appears to take place also within eachploidy level and as in other anurans. evolutionary

dis

call

sympatry as well as convergences ofparameters in allopatry can be expected. The lat-terseems to be the case in diploid B.

indivergences

which pulse
suggestedarate datalatastii.

from(Dubois and Martens. 1977)

polyploid

species(Stock,1998a).

Many morphometric traits exhibited differences ofthemeans but values showedlargeintersections

between both diploid and tetraploid toads and morphometric parameters were not suitable for ploidy
determination (Stock. 1997b). The study announced

by Roth (1986) on "about 2000 specimens from thewhole range" has never been published (Roth. pers.

comm.). In advance. Roth (1986) considered morphological characters "useless" for taxonomy but without

determination of the ploidy level of the collection
material. Multivariate analyses confirmed relationships between the ploidy level and the external character syndrome in diploid and tetraploid green toads(Stock, 1997b; Castellano et al.. 1998). Triploid toads

from Kyrgyzstan exhibited more distinct differences
to diploid than to tetraploid

specimens (Castellano et

Triploid populations in the Karakoram
showed significant differences to both diploid and tet-al.,1998).

Figure 9. A, B Lectotype of Bufo /<?/<3s///Boulenger,
1882 (BMNH 1947.2.21.28, formerly 72.4.17.223),
scale for B: 1 centimetre. C, D- Male B. /atast//(ZMB

62721 examined from Skardu, Baltistan. E - Histogram)

obtained by DNA flow cytometry from a mixture of

DAPI stained blood samples with chicken as the standard (a), CV = 2.94%, and diploid B. /atasti/Uom

Skardu (b), CV =-2.74%. Total cell number 22 744.

\

20 40

r—

-1

60

80

r-

100120140160180200

DAPI-tluorescence2001

Asiatic Herpetological ResearchVol. 9, p. 89

Thebeen found above 2000 m a.s.l."). Many papers (sum-character is occasionally misleading (Kuzmin, 1995:

was infre-

marized e.g. by Meinig, 1995) reporting the occurrence of "B. viridis" until 4000
a.s.l. in Central

quently used for the identification of toads for further

Asia, very probably represent data on polyploid toads.

investigations (e.g., Mezhzherin and Pisanets, 1990).

This also may account for Kuzmin's (1999) material

Distribution

dle Asia.

from Turkmenistan (Kurukhaudan

village.

94; Stock and Grosse, 1997a) although

it

12).

m

who reported on B. viridis up to 3200 m a.s.l. in MidIn general, the present

map (Fig.

I)

does not com-pletely reflect the occurrence of green toads

whoseploidy is known but also illustrates the recent degree

Tetraploid toads (white arrows). The
tetraploidtoads are largely distributed in the Middle and Central

of investigation.

Asian mountain systems. They have been describedfrom the Kopet Dagh (5, 6, 8, 11, 12), the Gissar range

The
Diploid toads (black arrows without starlet).
records of green toads found to be diploid in Middle

including its southern spurs (21 to 27, 27a, 30), thewestern Tien Shan (31a, 32, 33, 40), the northern Tien

Asia are distributedinthe Turan Basin and are also

Shan and the Issyk-Kul gorge (39, 51 to 59, 59b, 60),

concentrated

foothills of the high mountains.

the Central Tien Shan (42, 51), the margins of the Tak-at theDiploids have been found along the bottom part of thelamakan desert (43, 50?), the region between Zailiyskiy

Kopet Dagh mountains (7, 8, 10, 1 1); a second group ofrecords is to be seen in the Kafirnigan and Vakhsh river

Alatau and lake Balkhash (60, 37, 37a, 37b), the Dzungarian Alatau (60a, 60b), the eastern Tien Shan (63),valleys (23, 25, 28, 29). The last records seem to exem-and Dzungaria (62, 64, 66 to 72). In the Pamirs, tetrap-

plify a dispersion of diploid toads along the rivers intoloid toads have been found in the western (34, 35) andthe mountain valleys. The western and particularly thein the central parts (45).

northern Tien Shan form a third region with several
records of diploid toads (31a, 36, 38, 39, 59, 59a).

Furthermore, Stock (1998b)
found toads from eastern Pamirs (44) to be tetraploid.

More distant from the mountains researches combined

They were recently described by Fei et al. 1999) as a
new subspecies (T). The present map provides argu-

with ploidy determination have only sporadically been
carried out. Merely some localities with doubtless dip-toads from Eastern Tien Shan to the Dzungarian Gobi

loid toads near the lower Amu-Darya (9, 14) and Zeravshan river (17, 18) are known. The record of diploidin Mongolia. Interestingly, eastern of about 80° E
onlyrecords of tetraploids and obviously no record of dip-toads from the Chingzhal river (61) in Kazakhstan byloid toads have been made. Various records underline

Golubev (1990) appears to be doubtful (Dujsebayeva et

the high ecological plasticity especially of the tetrap-al.,1997). The main range (maps in Dubois and Mar-tens, 1977, Stock et al., 1999) of the diploid B. latastii(

ments for a rather continuous distribution of tetraploid

loid green toads (e.g.

Dujsebayeva et al., 1997). The

dry centers of the large continental deserts Kyzylkum,(N, 49) is the rather humid Kashmir valley and the

Karakum, Muyunkum and Betpak-Dala seem to be

surrounding rather arid mountain ranges.

not suitable as habitats of (tetraploid) green toads but

The small and highly generalized map by Kuzmin

this might be also caused by the recent low

degree of182) and the more detailed one (Kuzmin,

investigation. Considering some records of tetraploids

1999: 479) show either a continuous range or manyin the eastern plane regions, some additional recordsof tetraploid toads in the Turan lowlands can be(1995:

separate symbols of (diploid) B. viridis in Middle Asia

and Kazakhstan. Both maps cannot reflect the detailed
records and/or were not based on karyological studies.

Therefore, Kuzmin (1995, 1999) also announced thatthe distribution requires additional
investigations.

However, Kuzmin's maps refer to the occurrence of

expected in future. The tetraploids in the West of the
known range (Kopet Dagh) may have a rather contin-uous distribution via the mountains of Afghanistanof the Pamiro-Alai-System, but data fromAfghanistan are recently too scarce for a detailed
until those

diploid green toads in the Turan Basin along the rivers

analysis.

whereas the symbols for (diploid) B.

Inner Tien Shan and around the Issyk-Kul appear to

Triploid toads (black arrows with starlet). For the
first time, probable triploidy of three
specimens was

be very doubtful (e.g. see Borkin, 1989).

detected by Bachmann et al. (1978:

In spite of only

viridis in the

few references which contained

data on the ploidy level and on the altitudinal distribution, theoccurrence of diploid toads in Middle Asia

"36% more DNA

than diploid Bufo viridis" ) in toads (presumably) originatingfrom Kabul (48f in Fig.l;

1978:

352,

370).

Triploid

Hemmer et al.,individuals

have

been

m a.s.l. only in rare cases

detected in south-western Turkmenistan (5), in north-(appendix; Castellano et al., 1998: "At least in Kazakhstan and Kyrgyzstan no diploid populations haveern Kyrgyzstan (39) and south-eastern Kazakhstan;
the last record was reported without particular locality

apparently exceeds 1600

2001

Asiatic Herpetological ResearchVol. 9, p. 90(Borkin et al., 1997, 2000) and we did not draw it in
the map. The rare occurrence of triploid individuals in
the Central Pamirs (45) has been concluded

more continental high plains are similar and aretheobviously suitable only for polyploid green toads.

fromAt the moment, the causes for the specific distri-isozyme data (Mezhzherin and Pisanets 1990). Most

bution pattern remain speculation.

of the authors supposed a hybrid origin of triploids asa result of mismatings between diploid and tetraploid

189, 1999) refers to the high degree of temperature
tolerance of tetraploid green toads (-30°C to 45°C). In

toads or only noted the occurrence in contact zones of

the high mountain habitats as well as in the continen-

diploid and tetraploid toads (Borkin et al., 1997,
2000). Preliminarily, Lattes (1997) and Cervella et al.

(1997) indicated that triploids from Kyrgyzstan (Kokjar,

near 39; Castellano et al., 1998) do not originate

tal

Kuzmin (1995:

deserts strong daily and annual variations of tem-

peratures as well as a high mean annual solar radiance
2700-2800 h, in the(e.g. in the northern Asian deserts
interior Tien Shan > 2600 h, see Zlotin, 1997)

might
have caused a genetically caused selective advantage

from hybridizations between diploid and tetraploids
but seem to be closely related forms of the tetraploids.

of the polyploid forms. This may also account for the

Presumably all-triploid populations of green toads
from the Karakoram range and Western Himalayas

missing of diploid toads in the east of the range where
only tetraploids have been detected. The easternby Stock et al.
newasadescribed
andwere(Stocksubspecies(1998)

boundary of the range of tetraploid green toads andthe obviously parapatric occurrence of B. raddei inet al., 1999). The new data from Pakistan (47a, 48b-e)

Northern China and Western Mongolia was discussed

someby Peters (1971: "B. viridis" for tetraploids), Borkin
and Kuzmin (1988), and Stock (1998b). The most(46, 47, 48a) have been discoveredsuggest that triploids,

which form

at least at

localities (e.g.,47) all-triploid gonochoristic populations, are distributed along the upper tributaries of thewesternknown

records of tetraploids (Kopet Dagh,Gilgit river (48b), they live at the Shandur pass (48c,

Khorasan) seem to correspond with the region of tran-

3720 m a.s.l), and also in the Central Hindukush, i.e.

sition from winter-mild, summer-dry steppe climates of

the Chitral valley (48d, e). Interestingly, in the Kara-

Middle Asia and Afghanistan

koram exists obviously an eastern limit of the distribution of triploids situated between their most easternsummer-dry climates of Mediterranean type

record (48a) and the most western one (49) of diploid

Central Iran with fewer than 100

Based only
on morphological characters, Baig (1998) reported onlarge sand or salt areas seem to separate the polyploids

the sympatric occurrence of "Bufo latastii" and "B.
pseudoraddei" in the "Neelam valley" (= Jhelum val-

diploids in the West. Although B. surdus may provide
unexpected karyological data, at the moment, in spiteley) of Azad Kashmir in eastern Pakistan.

of only few studies, records of polyploid green toads inGenerally, the correlation of distributional data on
green toads with global climatic factors appears to beWest of Iran appear to some extent improbable.
This may rather result from the history of distribution

problematic because the local or microclimatic conditions to which the animals are adapted may differ

than from the possible relationship between polyploidy

strongly from the climatic zone in which any locality

The present image of the distribution of diploid andB. latastii in the gorge of the Indus river.

is situated.

Zoogeographic implications. Borkin (1999: 350)
considered "the B. viridis group as an indicator ofsouthern limits of the Palearctic in arid regions".
Mazik et. al. (1976) as well as Pisanets (1978) alreadyreported an allopatric occurrence of diploid and tetraploid green toads because they have different ecologipreferences. This hypothesis is principally
supported by the present map but it seems more
appropriate to call the ranges parapatric (Borkin et al.,cal1997). Many authors (Pisanets and Shcherbak, 1979;

Toktosunov, 1984; Borkin et al., 1986b, c; Borkin andKuzmin, 1988) arranged the tetraploids of the arid
foothill regions in contrast to those of the high mountains. In our opinion, it appears possible that the limit-

ing climatic factors in the high mountains as well as intothe winter-humid,
in

Iran

(Walter and Lieth, 1967; Miiller, 1996). The deserts ofmm rainfall/year andin the East of Central Iran (and Afghanistan?) from thetheand environmental selective pressure.

tetraploid toads possibly also corresponds

to the experi-ence that (especially allopolyploid organisms may
have advantages in regions with extreme or changing
environments (Futuyama 1990: 69

moreresistant"),(Bretagnolle et al.

when

-

colonizing"polyploids arenewhabitats

1998) or were considered to have ahigher competitive ability (e.g., Lumaret et al., 1997).
However, particularly the distribution of diploid plant

more
species and their polyploid relatives probably
historical factors thanof
influencethereflects
stronglyof their current ecological requirements (Bretagnolle etal.

1998) or adaptedness. Morescalchi (1990) stated:"In amphibians, changes in genome size (...), far frombeing random, they are related to metabolic andin thegenetic factors which are of crucial importanceanimals".
theseof
adaptive strategies
onto-Vol. 9, p. 91

Asiatic Herpetological Research

2001

Balletto,

Acknowledgments
We thank Prof. Dr. J. Martens, Mainz, for mating callfromrecordings

northern

and

Iran,

Dr.

T Dujse-Eremchenko, C. Giacoma, A.

Lattes, G.

Odierna. 1999. Sistematica e filogenesi nel complesso

T.
di

CYDOK

E.L, L. Borkin, S. Castellano,

bayeva, V.

Bufo viridis Laurenti, 1768. Rev. Idrobol. 38 (1/2/

image
Klapperstiick, Halle, for using the
Dr.
N.
St.
PetersProf.
Ananyeva,
analyses system.

3):

burg, Dr. R. Giinther, Berlin, Dr. C. McCarthy, London, Dr. G. Nilson, Goteborg, Dr. J. B. Rasmussen,

Iran. Tehran University Publ. 2250. (in Farsi).

K0benhavn, for loaning preserved specimens. Chromosome photographs were kindly prepared by G.
Hesse, Wurzburg. The basic computer-map was created by H. Nagel, Halle, according to our concept.
Many thanks to Dr. K. J. Baig, Islamabad, Dr. L. Ya.

Borkin, St. Petersburg, Dr. S. Castellano, Torino, Prof.

T Papenfuss,

Dr. A. Dubois, Dr. A. Ohler, Paris, Dr.

Schmidtler, Munich, and Prof. Dr. E.

F.

Berkeley, J.

Zhao, Chengdu, for help with getting literature. Dr. S.
L. Kuzmin, Moscow, kindly allowed to use his database "Amphibians of the former USSR".

Khatiri,

the

field

Gorgan, for their kind hospitality, R.
Dressel, Dresden, for optimism and assistance during

work

in

Pakistan,Dr.

D.

Baloutch, M. and H. G. Kami. 1995. Amphibians ofBedriaga, J. V. 1898. Wissenschaftliche Resultate dervon N. M. Przewalski unternommenen Reisen. Zoologischer Theil. vol. Ill, part 1. Amphibien und Reptilien. Kaiserliche Akademie der Wissenschaften, 1sted. St. Petersburg.

1981. Osobennosti differencialnikh

Birstein, V. Ya.

okrasok khromosom Anura na primere khromosom
nekototrikh vidov Rana Bufo. Zoologicheskii Zhuri

nal 60 (2):246-256. (In Russian).
Blair,

W. F. 1972: Evolution in the genus Bufo. Uni-

versity of Texas Press, Austin.

M. Stock is very grateful to the family of M. andM.

199-220.

Bogart, J. P. 1972. Karyotypes. Pp. 171-232 In W. F.
Blair (ed.), Evolution in the genus Bufo. Univ. Texas
Press. Austin, London.

Lamatsch,Wurzburg, for help with flow cytometry, and T. Dieterich, Greifswald, for a toad from Tengiz Lake (Kaza-

Boulenger, G. A. 1882. Catalogue of the Batrachia
Salientia in the collection of the British Museum. 2nd
ed. London.

khstan).

Boulenger, G. A. 1891. Descriptions of new Oriental
reptiles and batrachians. Annals and Magazine of Nat-

Literature Cited
Afrasiab, S. R. and H.A. Ali. 1988.

A new record of

toad Bufo surdus Boulenger (Amphibia, Bufonidae)
from Iraq, with preliminary key for Iraqi Amphibia.
Bulletin Iraq National Museum 8 (1): 115-123.

Andren, C. and G. Nilson. 1979. A new species oftoad (Amphibia, Anura, Bufonidae) from the Kavir
Desert, Iran. Journal of Herpetology 13 (1):93-100.

The Times Atlas of the World.
Edition.
London, Bartholomew andComprehensive
Times Books, Harper Collins:pl. 1-123 + l-222pp.
Anonymous.

1993.

Ataev, E. A. 1987. K ekologii danatinskoii zhaby na

Kyurendage. Izvestija Akademii Nauk Turkmenskoi
SSR. Seriya biologiceskikh nauk 3:67-69. (In Rus-

ural history (6) 7:279-283.

Borkin, L. Ya. 1989. Uber den Fund einer albinotisKaulquappe der tetraploiden Krotenart Bufochendanatensis Pisanetz. 1978 aus

dem Issyk-Kul'-See,

Kirgisien. Salamandra 25:122-124.

Borkin, L. Ya. 1999. Distribution of amphibians inNorth Africa, Europe, and the former Soviet Union,
pp. 329-420. In Duellman, W. E. (ed.) Patterns of distributionof Amphibians. John Hopkins University

press. Baltimore, London.

Borkin, L. Ya., I. A. Caune, E. M. Pisanets and Y. M.

Rozhanov. 1986a. Karyotype and genome size in the
Bufo viridis-gwup. pp. 137-141. In Rocek, Z. (ed.),Studies in Herpetology. Proceedings of the 6th Ordi-sian).

nary General Meeting of the Societas Europea Herpe-

Bachmann, K., A. Konrad, E. Oeldorf and H. Hem-

tologica. Prague.

mer. 1978. Genome size in the green toad (Bufo viridis group). Experientia 34:331-332.

The Amphibian Fauna of Azad
Jammu and Kashmir with new record of Paa liebigii.
Baig, K.

J.

1998.

Proceedings

Pakistan

Academyof

Sciences

35(2): 1 17-121.
Balletto, E., M. A. Cherchi and J. Gasperetti.

Bufo

viridis iz

idiploidnye populyacii zhab gruppy
Mongolii. Doklady Akademii NaukSSSR 287 (3):760-764. (In Russian).
Borkin, L. Ya., J. M. Rozanov, Ch. Terbish and I. A.

Caune.
1985.

Amphibians of the Arabian peninsula. Fauna of Saudi
Arabia 7:318-392.

Borkin, L. Ya., Ch. Terbish and I. A. Caune. 1986b.

Tetraploidnaya

1986c.

kariologia,
Raspostranenie,
izmenchivostch
sonomitcheskoe polozhenie
i

tak-

zhab

gruppy Bufo viridis v Mongolii. pp. 120-143. In Akademiya Nauk SSSR, Institut evolyutsionnoi mor-

Asiatic Herpetological ResearchVol. 9. p. 92

fologii

iekologii zhivotnych imeni A. N. Severtsova(ed.): Gerpetologicheskie issledovaniya v Mongolskoi

narodnoi respublike. (In Russian).

Borkin L. Ya., and T. M. Sokolova 1989. Electrophoretic comparison of diploid and tetraploid toadsof the Bufo viridis group (preliminary data), pp. 3638. In Szczerbak, N. N. (ed.), Problems of Herpetol-ogy. The 7th USSR Herpetol. Conf. Abstracts. Nauk-2001

1'Himalaya occidental (Cachmire et Ladakh). Bulletin
Societe Zoologique de France 102:459-465.

Duda, P. L., and K. Opendar. 1971. The karyotype of
Bufo sp. from Kashmir. Chromosome Information
Service 12: 18-20.

Dujsebayeva, T, S. Castellano, C. Giacoma, E. Balletto, and G. Odierna. 1997. On the distribution of the

ova Dumka Press, Kiev (In Russian).

Bufo viridis complex (Anura. Bufonidae) in Southern
Kazakhstan. Asiatic Herpetological Research 7:27-

Borkin, L. Ya., and S. L. Kuzmin. 1988. [Amphibians

31.

of Mongolia, species account], pp. 30-197. In E. I.
Vorob'eva and I. S. Darevski (eds.), Amphibians andEiselt, J. and J. F. Schmidtler. 1971. Vorlaufige Mittei-

of Mongolian Peoples Republic. General

Reptilesproblems. Moskva, Nauka. (In Russian).
Borkin, L. Ya., A. E. Vinogradov, Z. K.

Museums Wien 75:383-385.
Brushko andKuzmin. 1995. New records of tetraploid toads

S. L.

(Bufo viridis group) from Alma-Ata and Taldy-Kurgan region, Kazakhstan. Russian Journal Herpetology
2 (2): 174- 175.
Borkin, L. Ya.,

V K. Eremchenko andlung uber zwei neue Subspecies von Amphibia salienaus dem Iran. Annalen des Naturhistorischen

tia

Eiselt, J. and J. F. Schmidtler. 1973. Froschlurche ausdem

Iran unter Beriicksichtigung ausseriranischer
Populationsgruppen. Annalen des Naturhistorischen
Museums Wien 77:181-243.

Fei Liang, Ye Chang yuan, Huang Yong zhao, Chen
J.

M. Rozanov.

Xiao nuan. 1999. Taxonomic studies on Bufo viridis

1997. Distribution of and contact zones in the diploid-fromtetraploid Bufo viridis complex in Central Asia. p. 26,

Research. 20 (4): 294-300.

In Z.

Rocek andS.

Hart (eds.), Herpetology '97,

Abstracts of the Third World Congress of Herpetol-ogy 2-10 August. 1997, Prague, Czech Republic. Prague.

Borkin, L. Ya., J. M. Rozanov, and S. N. Litvinchuk.

2000.

Nuclear

DNA content in some green toads(Bufo viridis complex) of Turkey and Iran. Russian
Journal of Herpetology 7 (in press).
Bretagnolle, E, E Felber, E G. Calame and P. Kupfer.
1998. La polyloidie chez les plantes. Article desynthese. Botanica Helvetica 108 (l):5-37.

Carevsky, S. ,,1925". 1926. Notes on some batrachians from the Palearctic region. Ezhegodnik zoo-

logiceskogo muzeya. Leningrad 26 (l/2):74-78.
Castellano, S., C. Giacoma, T. Dujsebayeva, G. Odi-

and E.

erna

Balletto.

1998.

Morphometrical andacoustical comparison between diploid and tetraploidgreen toads. Biological Journal of the Linnean Society
63:257-281
Castellano, S. and C. Giacoma. 1998. Stabilising anddirectional female choice for male calls in the Euro-pean green toad. Animal Behaviour 56 (2):275-287.

DMAAC

-

Defense Mapping Agency Aerospace cen-

1986-1993. Tactical pilot charts (TPC). Scaleter.

A to D, F6 A to D, F7 A to D, G5 A to D, G-6 A to D, G-6 A to D. Different issues
1:500.000. Sheets F5

after.

west

China (Amphibia: Anura).

Zoological

Fikhtman, B. S. 1989. K diagnostike zelenykh zhab
kompleksa Bufo viridis Kirgisii. Pp. 263-264. In
Shcherbak, N. N. (ed.), Voprocy gerpetologii. Naukova Dumka. (In Russian).
Frynta, D., J. Moravec, J. Cihakova, J. Sadlo, Z. HodPitule, and L.
kova, M. Kaftan, P. Kodym, D. Krai,V

Shejna. 1997. Results of the Czech biological expedition to Iran. Part 1. Notes on the distribution ofamphibians andActa Societas Zoologica

reptiles.

Bohemica 61:3-17.
Futuyama,

D.

J.

1990.

Evolutionsbiologie.

Birkhauser. Basel, Bosten, Berlin.

Gerhardt, H. C. 1994. The evolution of vocalization infrogs and toads. Annual Review Ecology and Systematics 25:293-324.

Golubev, M. L. 1990. New records of amphibians andreptilesin

Kazakhstan(USSR). Vestnik Zoologii

9:76-78. Kiev. (In Russian).

Gruber, U. 1981. Notes on the Herpeto fauna of Kash-mir and Ladakh. British Journal Herpetology 6: MSISO.

Hemmer, H., J. F Schmidtler and W. Bohme.

1978.

Zur Systematik zentralasiatischer Griinkroten (Bufoviridis -Komplex) (Amphibia, Salientia. Bufonidae).
Abhandlungen Museum Tierkunde Dresden 34:349384.

1986. St. Louis, Missouri.

Dubois, A. and J. Martens. 1977. Sur les crapauds dugroupe de Bufo

viridis(Amphibiens, Anoures) deKashchenko, N. E 1909. Gady, sobrannye sredneaziatskimi ekspediciami prof.

V V Sapozhnikovav.

Vol. 9, p. 93

Asiatic Herpetological Research

2001

V and E. M. Pisanets. 1990. Genet-

1902-1906 i. 1908 gg. Ezhegognik Zoologicheskogo

Mezhzherin, S.

Muzeya imp. Bd. 14:119-130.
Khan, M. S. 1997. A new toad of the genus Bufo fromicheskaya struktura i proischozhdenie aziatskikh poli-

the foot of Siachin Glacier, Baltistan, Northeasternzhab Bufo danatensis Pisanets. 1978.
Nauk Ukrainskoi SSR, Serya B,
Akademii
Dokladychimiceskie
biologiceske. Nauki
Geologiceskie,
ploidnikhiPakistan. Pakistan Journal Zoology 29:43-48.

Kryukov, V. I., V. T. Rzhepakovskii and V. P. Astapov.
1985. Kariotipicheskaya charakteristika zhab semivyborok iz Tadzhikistana. Doklady Akademii Nauk
Tadzhikskoi SSR XXVIII (10): 593-595. (In Russian).

V

Ni, L. Ya. Borkin and L.V.
Kudryavcev. B. N.. V.
Solov'eva. 1988. Soderzhanie RNK belka v kletkakh(8): 71-73. (In Russian).

Mezhzherin, S.

V and E. M. Pisanets. 1991. Genet-

divergenciya

icheskaya

zelenykh zhab Bufo

kompleksacomplex fauny SSSR.

predstavitelei

viridisDoklady Akademii Nauk 317 (10): 222-226. (In Russian).

itetraploidnykh zhab gruppy Bufo viridis. Citologia 1 1:1324-1331. (In Russian).

diploidnykhi

Kuzmin S.

L.

1995. Die

Amphibien RuBlands undBrehm-Biicherei.
Neue

Gebiete.
angrenzenderWestarp Wissenschaften, Magdeburg.

Kuzmin S. L.

1999.

Mezhzherin, S.

V and E. M. Pisanets. 1995a. Genetic

structure and origin of the tetraploid toad Bufo danat-

ensis Pisanets, 1978 (Amphibia, Bufonidae) from Central

Asia: Description of biochemical polymorphism

and comparision of heterozygosity levels in diploid andtetraploid species. Genetica 31 (1): 43-53. (In Russian).

The amphibians of the former

Soviet Union. Sofia. Pensoft. 538 pp.

Mezhzherin, S.

V and E. M. Pisanets. 1995b. Genetic

structure and origin of the tetraploid toad Bufo danat-

Leviton A. E., Gibbs jr. R. H., Heal E., Dawson C. E.

ensis Pisanets, 1978 (Amphibia, Bufonidae) from Cen-

1985. Standards in herpetology and ichthyology: Part

tral

I.

Standard symbolic codes for institutional resource

collections in herpetology and ichthyology.

Copeia1985: 802-832.

Lumaret, R., J. L.Guillerm, J. Maillet, R. Verlaque.
1997. Plant species diversity and polyploidy in islandsof natural vegetation isolated in extensive cultivatedlands. Biodiversity and Conservation 6 (4): 591-613.

Macgregor, H. C. and J. M. Varley. 1983. Working
with animal chromosomes. A. Wiley-Interscience
Publication. Chicester, New York, Brisbane, Toronto,

Singapore.

Mazik, E. Yu., B. K. Kadyrova and A. T. Tokotosunov.
1976. Osobennosti kariotipa zelenoi zhaby (Bufo viridis) v Kirgizii. Zoologiceskii Zhurnal

55:1740-1742.

Asia:

Differentiation

forms andof geographic

genetic relationship between diploid and tetraploid species. Genetica 31 (3): 342-352. (In Russian).

Mocquard, F. 1910. Voyage de M. le Dr. Louis Villant
Pelliot). Bull. Mus.

dans l'Asie centrale (Mission
Hist. Paris 16:145-153.

Morescalchi, A. (1990): Cytogenetics and the problem of Lissamphibian relationships. Pp. 1-19 In E.

Olmo (ed.). Cytogenetics of Amphibians and Reptiles.
Advances in Life Sciences. Birkhauser, Basel.
Miiller, M. J. 1996. Handbuch ausgewahlter Klimasta-

tionen der Erde. Forschungsstelle Bodenerosion derUniversitat Trier, Mertesdorf (Ruwertal) 5 (5th ed.).
Trier.

(In Russian).

L. A. Lowcock, S. Cheryl, I. S.
R. D. MacCulloch, and D. E.
N.
Orlov,
Darevsky,

Meinig, H. 1995. Beitrag zur Hohenverbreitung derWechselkrote (Bufo viridis). Zeitschrift fur Feldherpe-

Upton. 1997. Flow cytometry in biodiversity surveys:
methods, utility and constraints. Amphibia-Reptilia

tologie (Bochum) 2:91-95.

18:1-13.

Mercadal,

1981. Determinacion del nivel deMurphy, R. W.,

ploidia en ejemplares preservados del genero Cer-

Nikolsky, A. M. 1 896. Diagnoses reptilium et amphibiorum in Persia orientali a N. Zarudny collectorum.

atophrys. Amphibia-Reptilia 3/4:205-212.

Annuaire

1971a. Die Amphibien und Reptilien
West-Pakistans. 2. Nachtrag. Senckenbergiana biolog-

Imperial des Science, St. Petersburg 1:369-372.

I.

T.

Mertens, R.

ica, 52:7-15.

du

Musee

Zoologiquedel'Academie

Nikolsky, A. M. 1897. Presmykajushchiyasya, amfibii
iryby, sobrannyya N. Zarudnym" v' vostochnoi Per-

Annuaire du Musee Zoologique de l'Academie

Mertens, R. 1971b. Zur Kenntnis von Bufo luristani-sii.

cus (Salientia, Bufonidae). Salamandra. 7:83-84.

Imperial des Science, St. Petersburg 2:306-361.

Meyer, F. 1991. Beobachtungen an Bufo danatensis(Pisanetz, 1978) in der Mongolei (Amphibia, Anura:
Bufonidae). Zoologische Abh. Mus. Tierkd. Dresden
46: 157-164.

Nikolsky, A. M. "1899", 1900. Reptiles, amphibiens
poissons recueillis pendant le voyage de Mr. N.

etZarudny en

1

898 dans la Perse. Annuaire du MuseeAsiatic Herpetological ResearchVol. 9, p. 94

2001

Zoologique de l'Academie Imperial des Science, St.

Roth, P. and P. Rab. 1986. Karyotype analysis of thePetersburg 4:375-417.

Bufo viridis-Group: Systematic implications. Pp. 131136. In Rocek, Z. (ed.) Proceedings of the 6th Ordinary General Meeting of the Societas Europea Herpe-

Orlova, V. F. and V. K. Uteshev. 1986. The tetraploid
toad of the Bufo viridis group from Dzungarian Gobi.

Mongolia, pp. 137-141. In Rocek, Z. (ed.), Studies in
Herpetology. Proceedings Ord. General. Meet. Societ.

tologica. Prague.

Europ. Herpetol.: Prague.

banding studies in the natterjack toad, Bufo calamita.
335-338. In Van Gelder, J. J., H. Strijbosch and P. J.

Peters, G. (1971): Materialien zur Okologie und Ver-

breitung der Amphibien in der Mongolei. Mitteilungen Zoologisches Museum Berlin 47: 315-348.

M.

Pisanets, E.

O novom poliploidnom vide

1978.

zhab Bufo danatensis Pisanets.

sp. n. iz

Turkmenii.

Doklady Akademii Nauk Ukrainskoj SSR,

Ser.

B,

Pisanets, E. M. 1987. Razlichiya v kladke ikpry zele-

noi i danatinskoi zhab. Vestnik Zoologii 6:80-81. (InRussian).

M.

Bufo

viridis

-

i

of

the

Societas

Europea Herpetologica.

Nijmegen.

Schmid,M.

Amphibia.

I.

1978.

Chromosome

bandingM.

plex) from Eurasia (Anura:Bufonidae). Pp. 357-360.
In Korsos, Z. and I. Kiss (eds.). Proceedings 6th Ord.

General. Meet. Societ. Europ. Herpetol.: Budapest.

M. 1992b. Poliploidiya

iotkloneniya vuzhab
Bufo danatenmorfologii gonad
tetraploidnikhsis (Amphibia: Anura:

Bufonidae). Citologija i gene-tika 26:35-39. (In Russian).

Schmidt, K. P. 1952. Diagnoses of new amphibians
and reptiles from Iran. Nat. Hist. Misc. Chicago Ac.

Schmidt, K.

P.

Iran. Vidensk.

1955. Amphibians and reptiles from
Medd. Dansk. Naturh. Foren 117:193-

207.

surdus; mit einem Schlussel und Anmerkungen zu den

ubrigen Kroten Irans und West-Pakistans. Salamandra
5 (3/4): 113-123.
Schneider, H. and E. M. Egiasaryan. 1995. The mating call of the green toad, Bufo viridis viridis (Anura,
Amphibia), in Kazakhstan. Russian Journal Herpetol-ogy 2:1-4.
Schroer, T 1996. Morphologie und Ploidiegrade vonWasserfroschen aus unterschiedlichen Populations-

Nordost-Polenin(Anura,systemenzelenykh zhab (Amphibia, Anura) fauny SSSR.
Vestnik Zoologii 4:1 1-16. (In Russian).

Zeitschrift fur Feldherpetologie 3:133-150.

tika

Pisanets, E. M. and O. I. Vasilenko. 1995. Reproductive age of di- and tetraploid toads (Bufo viridis turan-nakhodki zemnovodnykhpresmykayushchikhsya v
Kazakhstane. Vestnik Zoologii 1:70-72.
i

Srednei Azii

(In Russian).

47 In Ananjeva, N. B. (ed.) Abstracts of the 2nd Asian

Stock,

Hepetol. Meet. Ashgabad.

cytologischeE.

vneshei

sp.

,

S.

i

nov.

V Mezhzherin and N. N.

Issledovanya

morfologii

Bufonidae)
sis

M.
1996.

aziatskikh

po gibridizacii
zhab (Amphibia:

iopisanie novogo vida Bufo shaartusienDopovidi nacional'noi akademii nauk

Ukraini 6:147-151. (In Russian).
Roth, P. 1986. An overview of the systematics of thegroup in Middle and Central Asia. Pp.
127-130. In Rocek, Z. (ed.) Proceedings of the 6th
Ordinary General Meeting of the Societas Europea
BufoviridisHerpetologica. Prague.

Ranidae).

Shcherbak, N. N. and M. L. Golubev. 1981. Novye

ensis and B. danatensis) in South Turkmenia. pp. 46-

Shcherbak.

Chromo-soma 66:361-388.

Pisanets, E. M. and N. N. Shcherbak. 1979. Sistema-

Pisanets,

inConstitutive heterochromatin and nucle-

Schmidtler, J. J. and J. F. Schmidtler. 1969. fiber Bufo

1992a. Evidence of hybridization
between di- and tetraploid toads (Bufo viridis - com-

Pisanets, E.

M. Bergers (eds.). Proceedings 4th Ordinary General
MeetingSci. 93:2.

Novye dannye po kariologii
Complex (Amphibia, Bufonidae)(In Russian).

E.

and P. Rab. 1987. Sequential chromosome

1991.

voprocy proischozhdeniya aziatskikh tetraploidnykh
zhab. Herpetological Researches 1:41-50. Leningrad.

Pisanets,P.

olus organizer regions in Bufo and Hyla.

Geol., chim. i biolog. Nauki 3:280-284. (In Russian).

Pisanets, E.

Roth,

M.

i

1995. Morphologische, ethologische undUntersuchungen zum taxonomischenundStatus ditetraploider Griinkroten (Bufo viridis

Komplex)inMittelasien.

Diploma Thesis, Martin-

Luther-University Halle- Wittenberg, 107 pp.

M.

1997a. Mating call differences between
anddiploid
tetraploid green toads (Bufo viridis comStock,plex) in Middle Asia In Rocek, Z. and Hart, S. (eds.)
Herpetology '97, Abstracts of the Third World Con-

gress of Herpetology 2-10 August.

1997, Prague,

Czech Republic.

M. 1997b. Untersuchungen zur Morphologie
und Morphometrie di- und tetraploider Griinkroten(Bufo viridis -Komplex) in Mittelasien (Amphibia:
Stock,

2001

Anura:

Asiatic Herpetological Research

Bufonidae).

Zoologische

AbhandlungenStaatliches Museum Tierkunde Dresden 49:193-222.

Stock,

M. 1998a. Mating

call

differences between

I.
1997. Geography and organization ofmountainhighecosystems in the former USSR. pp.

Zlotin, R.

133-159. In Wiegolaski, F E. (ed.) Ecosystems of the

diploid and tetraploid green toads (Bufo viridis com-world,

plex) in Middle Asia. Amphibia-Reptilia. 19:29-42.

Amsterdam.

Stock, M. 1998b. Tetraploid toads (Bufo viridis complex) from north-western China and preliminary taxo-

nomic conclusions for Bufo nouettei Mocquard, 1910.
Zeitschrift fur Feldherpetologie (Bochum) 5:139-166.

Stock, M. and W.-R. Grosse. 1997a. Erythrocyte sizeand ploidy determination in green toads {Bufo viridis

complex) from Middle Asia. Alytes (Paris) 15:72-90.
Stock, M. and W.-R. Grosse. 1997b.

New data fromthe Bufo viridis complex in Central Asia. In Rocek, Z.

and Hart, S. (eds.) Herpetology '97, Abstracts of theThird World Congress of Herpetolgy 2-10 August.
1997. Prague, Czech Republic.
Stock, M., W.-R. Grosse, M. Schmid and C. Steinlein.
1998. Ploidy level diversity in the Bufo viridis com-plex of Central Asia Zoology (Jena). Suppl. 1:76.
Stock, M., M. Schmid, C. Steinlein and W.-R. Grosse.
1999. Mosaicism in somatic triploid specimens of the

Bufo viridis complex in the Karakoram with examination of calls, morphology and taxonomic conclusions.
Italian Journal of Zoology 66 (3): 215-232.

Toimastov, S.
iz

S.

1989. Osobennosti morfologii zhabokrestnostei oz. Jashil'kul' na Pamire. Izvestiya

Akademii Nauk Tadzhikskoi SSR, otdelenie biologicheskikh nauk 4 (1 17):43-48. (In Russian).

Toktosunov, A. T. 1984. Ecological basis of altitudeadaptation of the vertebrates of Tian-Shan.
Publ. House. Leningrad. (In Russian).

Nauka

Toktosunov, A. T. and E. Yu. Mazik. 1977. Napravlennost' zhab v Tyan' -Shane, pp. 203-204. In Darevski, I.
S. (ed.) Voprocy gerpetologii. Nauka. (In Russian).

Ullerich, F.-H. 1966. Karyotyp und DNS-Gehalt von
B. viridis, B. bufo x B. viridis und B.
calamita (Amphibia, Anura). Chromosoma 18:316342.

Bufo bufo,

Walter, H. and S.-W. Breckle.

1986. Okologie derErde. Vol. 4. Stuttgart. Gustav Fischer.
Walter,H.

and H. Lieth.

1967.

Klimadiagramm-

Weltatlas. 3rd. ed., Gustav Fischer, Jena.

Wu Min and Zhao Yajiang. 1987. A preliminary studyof the karyotype of Bufo viridis Laurenti in Xinjiang.

Zoological Research, Kunming 8 (4):339-342. (In
Chinese with English summary).

Zhao, E.-M. and K. Adler.

1993. Herpetology ofChina. Oxford, Ohio, USA, SSAR:l-522pp.

Vol. 9, p. 95

vol.

3:

Polar and alpine

tundra.

Elsevier,Vol. 9, p. 96

2001

Asiatic Herpetologieal Research

Apppendix 1
L\£'

PloidyTopographic description

SourceMethod2001

IS

Asiatic Herpetological ResearchVol. 9, p. 97

Vol. 9, p. 98

37a

Asiatic Herpetological Research

2001

,,
34

Vol. 9, p. 99

Asiatic Herpetological Research

2001

.

n

Kyrgyzstan. Issyk-kul. N-bank near Sary-Kamysh.

1670

m a.s.l.. 42°29'N,76°20'E

Stock ( 1997a), Stock and Grosse (1997a).
ISlock (1998a)—al.:„ —n.
eryuir. size, cansVol. 9, p. 100c-,Asiatic Herpetological Research

Tadzhikistan, SW-part Beshkent desen, near Shaartus. PisanetM 1991 1, Pisanets el al (1996),
Kuzmin (1999). see also number 29
type localitv of Bufo shaarmsiensis

2001

•aasA