Asiatic Herpetological Research

December 1993

Vol. 5, pp. 105-108

Karyotype Information on some Toad Agamas of the Phrynocephalus
guttatus Species Group (Sauria, Agamidae) of the former USSR.

VALANTINA V. MANILO AND MICHAEL L. GOLUBEV
Instituteof Zoology, Academy of Sciences, Kiev, Ukraine

Abstract. -Karyotypes of several toad agamas of the Phrynocephalus guttatus species group (sensu law)were investigated in specimens from a variety of localities of the former USSR. Differences in thediakinetic stage of meiosis have been observed, permitting distinctions among three groups of species. Theforms guttatus, moltschanovi, kushackewitschii, and alpherakii comprise Group I; P. guttatus salenskyi
represents the second group; and P. versicolor hispida represents Group III.

Key words:

Reptilia, Sauria, Agamidae, Phrynocephalus guttatus, Kazakhstan, Middle Asia, Precaucasus,

karyology.

40.

USSR

fauna: la- P. g.
Scheme of distribution of forms of P. guttatus species group of the former
guttatus; lb- P. g. moltschanovi; II- P. g. kushackewitschii; III- P. g. alpherakii; IV- P. g. salenskyi;
V- P. versicolor hispida; VI- P. guttatus spp. (the numbering of populations is in accordance with the data

FIG. 1.

in table 1).

Introduction

The first and only extensive karyologicalinvestigation of the agamid lizard genus
Phrynocephalus Kaup is the work of
Sokolovsky (Sokolovsky, 1974; 1977).
Karyotype characteristics permitted therecognition of five groups. The "guttatus"group included two species, P. guttatus(Gmel.) and P. versicolor Str. Thesespecies have a diploid number of 46, allThe
are telocentric.
into12couldbedivided
pairskaryotypesof macrochromosomes and 1 1 pairs ofchromosomes

microchromosomes. Approximately 50%of the metaphase plates in P. guttatuscontained satellite chromosomes on the firstpair of chromosomes, but these were neverobserved in the P. versicolor karyotype.

© 1993 by Asiatic Herpetological Research

The specimens examined came fromDaghestan (P. guttatus) and Central Gobi,
Mongolia (P. versicolor) and were believed
to represent the nominative forms of bothspecies.

The systematics of the P. guttatus groupbased on external morphologicalcharacteristics is extremely complicated andremains unclear. At various times theforms alpherakii Bedr., moltschanovi Nik.,kushackewitschii Bedr., salenskyi Bedr.,
etc. have either been included in P.
guttatus, sensu stricto or treated as related

The forms bogdanowi Bedr.,and paraskiwi Semenov etBedr.,
hispidaal. have been assigned to P. versicolor.
species.

1909; Nikolsky,1915;(Bedriaga,Terentjev and Chernov, 1949; Peters,Asiatic Herpetological ResearchVol. 5 p. 106m\i H HIi•w•finVnolif)

fl«AH HiM~»>•---*--FIG 3. Diakinetic stage of meiosis of five forms of
P. guttatus species group, s. lato: I- all elements

stinoDecember 1993

are ring- or stick-shaped; (guttatus, moltschanovi,kushackewitschii, alpherakii); II- one or twoelements are cross-shaped (salenskyi); III- two andmore (up to four) elements are cross-shaped^*%\v(hispida).

-

performed by Giemsa stain (2% solution)
in 0.01
sodium-phosphate buffer (pH6.8) for 20-30 minutes. After washing indistilled water, the preparations werepassed through alcohols and xylols(orthoxylol) and subsequently embedded in
Canadian Balsam.
In excess of 30
fromeach form weremetaphase platesaBiolam 1-212
investigated usingMFIG. 2

The karyogramme of Phrynocephalus

guttatus salenskyi.

1984; Semenov et al. 1987). Golubev(1989) suggested that P. guttatus and P.
versicolor from Kazakhstan areconspecific. Karyotype details of the forms
listed above have never before been
examined. The purpose of this study is todetermine whether karyotype information
will aid in our understanding of thesystematicsandevolutionof

Phrynocephalus.

Methods
Between 1989-1991 we collected
specimens of nearly all listed forms of both
species of Phrynocephalus inhabiting theterritory of the former USSR with the
exception of P. v. bogdanowi from theextreme south of Tuva (Central Asia) andP. guttatus ssp. from Turkmenistan(Fig. 1and Table 1). Chromosome samples wereprepared from cellular suspension of bonemarrow, blood, and testis. We used asmear method and a method known as"digging out" in conformity withprocedures described by Ford andHamerton (1956) and McGregor andVarley (1986) as partially modified byManilo (1986). Chromosomal staining wasmicroscope.

Metaphase plates of spermatogonialdivision, spermatocyte I (diakinesis) andspermatocyte II (metaphase II) bivalents
were investigated in testis preparations.
Chromosome morphology is describedaccording to the classification proposed by
Levari etal. (1964).

Results and Discussion

Our data support

thefindingsofSokolovsky (1974, 1977). The diploid
number is uniformly 46 and theFundamental Number (NF) is 46.
Inseveral forms {guttatus, moltschanovi,kushackewitschii) we noted satellite
chromosomes on several plates; whereas inother forms (alpherakii, salenskyi) we saw
no evidence of satellites. The revelation ofthis structure largely depends on thedegreeof spiralization.
will

be foundpossible that satellites
the latter forms with

It isinfurther investigation andmaterial.

more extensive

December 1993

TABLE 1.

Asiatic Herpetological ResearchVol. 5 p. 107

Localities, sample sizes, and taxa of Phrynocephalus guttatus s. lato populations collected andinvestigated in this study (numbering of populations is given in accordance with the data shown in Fig.

No

1.

Asiatic Herpetological ResearchVol. 5 p. 108

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