I

Asiatic HerpetoloRical Research

June 1<W5

Social Organization

Vol. 6, pp. 97-110

and Demography in the Rock Agama, Stellio caucasius

EUGENY N. PANOV AND LARISA Y. ZYKOVA
Severtsov Institute of Animal Evolutionary Morphology and Ecology, Russian Academy of Sciences,

Leninsky Prospect 33, Moscow 117071 Russia,

Abstract.

-We studied the social organization and demography of the rock agama Stellio caucasius

in anatural population, located in Gobustan (eastern Azerbaijan, approximately 60 km south of Baku) and in an

introduced population in the small Karadag Range near Krasnovodsk (western Turkmenistan).

We found thatthese populations are highly stable with low turnover. This appears to be the result of delayed reproduction,

longevity and a sedentary life style. Population growth is relatively slow due to high juvenile mortality andlow immigration rates from adjacent subpopulations. The age structure of all subpopulations studied was

dominated by older age classes. Rock agamas exhibit those natural history and population characteristic of a

K selected species.
Key words: Agamidae, age, Azerbaijan, density dependence, K-strategy, Lacertilia, ontogenetic trajectories,
polygyny, population dynamics, social behavior, spacing, Stellio caucasius, survivorship, mortality,
territoriality, translocation, Turkmenistan.

FIG.

1.

Adult (10+ years old) male Stellio caucasius

perched on basking site (left); adult female S.

caucasius (right).

Introduction
Long-term mark and recapture studies ofnatural populations have become importantin recent decades due to their great power indemonstrating population parameters.
These investigations permit the testing of
hypotheses concerning the mechanisms
The
governing population parameters.
ability to follow particular individuals© 1995 by Asiatic Herpetological Researchthrough time reveals the scale of behavioral
heterogeneity within the local population. It
also allows the examination of these
population parameters as a function of age
or changing social status.

This approach has already gained firm
position in the population studies of birds
and mammals, but it has only recently beenbroadly

employedinreptilestudies.

Vol. 6, p. 98

June 1995

Asiatic Herpetological Research

populations are characterized by high andMost individuals
very high density.
demonstrate strong home area fidelity.
Adults live in small social groups in which
the lizards form long lasting pair bonds(Panov and Zykova, 1985; 1989).

This

paper

examinesthesocialinteractions within local settlements of rockagamas and analyzes the role of social

behavior as a regulator of demographic
processes.

Methods and Materials
Rock agamas are large lizards with an
overall length of up to 30 cm. and weighing
up to 160 gm. Males are on average larger
than females and have a heavier build.
General background color is a mixture ofgray, brown and olive with a darker, dull

FIG. 2. Regional distribution (cross hatched area)of Stellio caucasius (upper plate).
sites inSpecific study

Azerbaijan and the Turkmen Republic:1)

Gobustan; 2) Krasnovodsk; 3) Bol'shoi Balkhan
Range; 4) Kyurendag Range; 5) lower Sumbar
River; 6) Parkhai Gorge; 7) Kalaligez; 8) Aidere(lower plate).

Although this method is used in modern
herpetology rather frequently, it is oriented
mainly toward answering the questions of
traditional population ecology (dynamics ofnumbers and sex-age composition of local
populations, modes of spacing in the context
of resource utilization by communities andspecies, etc.). In behavioral ecology, andparticularly with respect to the fates ofparticular individuals and their social
relationships, the reptiles in general andlizards in particular remain poorly studied(for review see Semenov, 1989).

The rock agama, Stellio caucasius, is an
ideal subject for the study of behavioral
aspects of population organization in reptiles
that arestrategists. This is a long-lived

K

lizard attaining an age of ten or more years(Zykova and Panov,1991).

Many

spotted pattern on the back and sides.
During the breeding season males differ
from females by having black on the breast,
contrasting with pale or pinkish-gray on thethroat (Fig. 1). In males, there is light grayepidermal holocrine gland in the center ofthe blackish-gray belly. These lizards are
typical inhabitants of stony landscapes,
although some populations have become
adapted to the life on the steep slopes of clay
canyons or even at the margins of the sandy
desert (Ananyeva and Ataev 1984; Panov,Zykova, Glauzer and Vasil'ev, 1987).

The bulk of data presented here wasobtained during a comparative study of two
populations of rock agama: a natural
population, located in Gobustan (eastern
Azerbaijan, approximately 60 km south ofBaku) and in an introduced population in thesmall Karadag Range near Krasnovodsk(western Turkmenia) where rock agamas
were known to be absent earlier (Fig. 2). In
the second area, on 17 May 1985, we
introduced into an abandoned quarry 13
adult males, 19 adult females, 13 two year
old lizards and 25 juveniles born in thepreceding year. All these animals werecaught in the Blocky Balkan Range situatedsome 160 km from the introduction site.
The latter lies in view of Krasnovodsk
Plateau known to be a part of rock agama

Asiatic Herpetological Research

June 1995

1

«"2

c'3

Vol. 6, p. 99

94

FIG. 3. Territory structure in Gobustan (A) and Aidere (B): 1)

territory boundaries; 2) breeding males; 3)

subordinate males; 4) females.

The
1985).
tothe
introduction site is similardrysemidesert habitats of rock agamas found in
Gobustan. The predominately stony surface
is covered by very sparse grasses, but trees
and bushes, in contrast with Gobustan, are

23 May1988, 5 - 10 April and 28 April
and
10-11
15-25
1989,
SeptemberMay
Some
1990 (total of 103 days).
observations were made during short visits
in the summer and fall from 1985 to 1989.

wholly absent. The experimental plot was

Important additional data were obtained
during the course of field studies conducted
on two marked populations in western
Kopetdag near Kara-Kala settlement (SjuntAn
Khasardag State Reserve).
observational plot in Parkhai Gorge wasinspected in the spring months of 1986 andbreeding

range(Ataev,situated in a broad, dry valley with steep
slopes broken by narrow ravines and rifts.

The numerous cavities and cracks under andbetween the stones provided abundant
temporary and permanent shelters for the
lizards.

Prior to releaseallintroduced

agamas were marked by toe clipping.
In Gobustan, on the natural plot of 0.65

marking andcapturing,observations were conducted in April 1987
and 1988; in spring of 1986 and 1989 we
performed censuses and selective capture of
lizards. In Krasnovodsk, on the introduced
population, field work was carried out 24
April 6 May 1986, 22 25 March and 30- 18
May 1987, 26 April - 17 May
April
hectares,1989 and in September - October 1986 1988; a population in the Kalaligez area was1987 (total
investigated in the fall of 1984of 25 days).
In all of the above study plots we carriedout total censuses of lizards in the areas
under study. Most of the agamas observedon the plots were captured. They were
measured with a ruler and calipers according
to standard procedures, weighed andVol. 6, p. 100cAsiatic Herpetological Research

June 1995

Asiatic Herpetological Research

June 1995

Results and Discussion

16.3

Vol. 6, p. 101

m 2 on average), for details see Panovand Zykova (1985).
Spacing Patterns in Rock Agama
Settlements

The baseline of spacing pattern of awhole social organization in Rock Agama isa mosaic of mutually exclusive territories

owned by mature adult males.

Maturefemales have either mutually exclusive oroverlapping home ranges situated within
territories of those males with which thegiven female is tied by the family bonds.
The adult female, as a rule, does not leave
the territory of "her" male. Therefore, theterritory of adult breeding male, if there isfemale(s) living in it (which is notnecessarily the case) in the same time theterritory of pair or a family group defended
as a whole by the breeding male only.

The home ranges of immature lizards
one and two years of age may lie bothwithin the territory of the family group or
outside in the neutral zones separating suchterritories. The home range of an immature
male during the first years of life within an
adult male territory, shifts to the periphery
this territory as the immature male
becomes older. In contrast, the home rangeofof an immature female adjacent to that of an
adult male shifts with time toward the male's
territory.

Formation of Territorial Structure

We followed the establishment ofterritorial structure in the course of our long-term observations on the Krasnovodsk
Here 13 male
introduced population.
into
an area thatwerereleased"founders"
a
have
would
population of a
supported
density comparable to that in natural
Males werecolonies of rock agamas.
which
and
crevicesholes
into
released
deep
seemed to us to be similar to hollows

normally used by rock agamas as theirpermanent dens.

However, contrary to our expectations,the majority of introduced males left the area

where they had been released, and moved
away at distances ranging from 60 to 500
m. Only three males remained in proximity
to the release site by the next spring. At thattime the territory of only one male (N 56)
overlapped the release site. The boundaries
of two other males (NN 36 and 31)
territories were 60 and 100 m respectively
from the release site (Fig. 4 c, d).

The process of territory establishment
adjacent to the release site is shown in Fig. 4c-f.
During 1986, the year following
introduction, adult male territories werecommonlarge and had no common boundaries (Fig.
4d). Since male rock agamas do not patrol
the borders of their territories (as, forboundaries. The size of territories in suchsaturated habitats depends on the substrateexample, males of steppe agama, Trapelus
sanguinolentus, do (see Panov and Zykova,

and local food abundance (Fig. 3). In
barren habitats of Gobustan with low
microrelief (absence of talus mounds, inparticular) the size of adult male territories
was found to vary from 100 to 210 m 21986), it is difficult to locate precisely theboundary between territories and, therefore

In saturated habitats all suitable space isamong adult males, so thatsharedterritories

neighboringhave

m on average). In the middle(140.0 15.8
altitudes of2

theWestern

Kopetdag

Mountains (Canyon Ai-Dere, see Fig. 3B)
where the climate is more humid and thevegetation is rich and diverse and thesubstrate includes jumbles of fallen rocks
and boulders, the size of territories was±found to range from 28 to 136 m 2 (94.0

to estimate exactly the real territory size.

The greatest territory diameter was estimatedin 1986 on the experimental plot as 140 m,with the width of the neutral zones
separating neighboring territories as some

20-60m. The small, indistinctly demarcatedhome ranges of three immature lizards(males NN 30 and 61, and female N 41, allless than two years old) were situated in
Otherthese neutral zones (Fig. 4e).
immatures approximately two years oldestablished their home ranges within the
territories of mature males, as well as fourVol. 6, p. 102

Asiatic Herpetological Research

lizards born already in place of introductionin preceding year.

In

1988,

the

third

year

after

introduction, most of the area that in 1987

was neutral zone was shared among
relatively young males of about four yearsof age (NN 30 and 61) and three years ofage (N 1 15 already born on the introductionsite). Although these males apparently had
attained maturity, they seemed to bebachelors at that time. Three year old maleN 1 15 and four year old female N 74 wererepeatedly seen in 1988 in close proximity(10-15 m) to each other, but we did notwitness immediate contacts between them.
In

1989,

the

fourth

year

after

introduction, slight changes in territorial
structure were observed (Fig. 4e). Therewas some tendency toward clumping ofterritories toward each other, possibly
because of the increasing density of lizards.
Male N 61 (about five years old) left hisadolescent bachelor home range andoccupied the territory of deceased male N 31
and established a bond with his former
mate, a female of eight or nine years of age.
The corpse of male N 31 indicated that hehad died between June 1988 and April
1989.
Relatively young male N 115
occupied the territory of five to six year old
male N 4a after he disappeared. Female N
74 moved into the territory of 10 to 11 year
old male N 3.

June 1995

often move greater distances from the den,up to several tens of meters.

During the day adult males range more
The several posts of a male areconnected by a network of relatively
The territory ispermanent pathways.
utilized unevenly with some points locatedalong the pathways receiving regular use
while others, situated at some distance fromwidely.

the pathways being visited only occasionallyor not used at all during a given season.

This pattern of territorial use can resultchanges in territorial boundaries. For
example, by comparing the positions "c","d", and "e" in Fig. 4, one can see that in1988 the border of the territory of male N 36
shifted 30-40 m eastward from its positionin 1986. Such shifts are possible in nonin

saturated habitats only.

In established rockagama settlements with dense populations
the boundaries remain constant from year toyear.

In relatively sparse Krasnovodsk
population the cases of mutual intrusions byneighboring males into neutral zones

separated their territories and even into
peripheral parts of these territories
themselves are possible. The above said
does not hold in respect to male"pretenders", or "satellites". Their homeranges may broadly overlap the peripheral
parts of two or more territories of adult"resident" males (see, for example,

Dynamics of Space Utilization Within aHome Area
During the hot periods of the year, eachindividual had at least one permanent den
and one to several observational posts (usedalso as basking sites) where it spentconsiderable time, except during periods offoraging. In individuals about one year oldor younger the den and basking site were thesame or immediately adjacent. Usually the
shelter was situated under a boulder or rockon which the lizard basked. Foraging
activities of juveniles and yearlings takeplace within a radius of several meters of theindividual dens. Many subadults of both
sexes and some adult females behave in asimilar way, although during foraging theyhomeranges of male-pretenders NN 30 and 61 in
Fig. 4). In saturated habitats home rangesof satellites are practically always situatedwithin the territories of resident males.

By the winter the agamas leave theirsummer home areas and migrate tocommunal hibernation shelters. Migrations
begin when air temperatures are relatively
high (25° C and above). Communal winterdens may be situated up to 500 m from anindividual's home area. On 23 March 1987

on the Krasnovodsk experimental plot,when only a few agamas had returned totheir summer areas from the communal
hibernation shelter (daily temperatures
ranged from 2° C to 16.5° C) we found inthat shelter males NN 21, 27, 36, 61,

June 1995

Vol. 6, p. 103

Asiatic Herpetological Researchfemales NN 22 and 58 (ages from two toseven years) and subadult male N 105.
Summer residences of these lizards areshown in Fig. 4, home areas of others werelocated 100 - 500 m from the communal
winter den.
Individual Ontogenetic Trajectories

An ontogenetic trajectory is defined as asequence of changes in social status and ofsocial roles of an individual during its life
In rock agamas social(Wiley 1981).

behavior and social status of individuals ofboth sexes appear to be similar during thefirst two years of their lives. But thereafter
ontogenetic trajectories of males and females

and 5 females).

When the home ranges offirst year lizards

were immediately adjacent

their interactions appeared to be agonistic.

some cases between adjacent homeranges a well defined boundary wasInestablished and both neighbors displayedpronounced territorial behavior toward each
In other cases the home ranges
other.

overlapped and a stable rank order wasformed so that one lizard appeared todominate the other in the overlap zone.
Generally, the mature members of thesettlement behave indifferently toward
However, in periods of highyearlings.
sexual activity adults may chase theyearlings short distances.

become progressively divergent.
After the second winter the young
It is

not known if lizards remain in thevicinity of their birth place during their lives.
those juveniles (n=61) that wereAmong

captured on the two plots in the western
Kopetdag at the end of September and the
beginning of October (i.e. at the age of two
to threemonths), only five (8.2%) wereagamas returned from communal den to their
original summer home area where theyknew the topography of the place and theirforaging routes became longer andpioneering of new feeding places and new

Apparent
temporary shelters began.
differences between social behaviors ofobserved near the places of their first capture(within a radius of 25 m) in next year. The
proportion of such recaptures on one of thetwo plots was as high as 25%, while on
another plot none of 41 juveniles caught
during preceding fall was observed later on.
Unfortunately of all those juveniles thatwere captured during the first months of
their lives, it remains unknown whether theyremained near their birthplaces or if theydispersed by the time of capturing(dispersion of juveniles just after hatchinghas been described for Anolis aeneus

young males and females began to emerge

Stamps, 1988).

ages and two immature females. In addition
the same year we observed interactions
between this male and immature female N
161 in the border zone between his territory
and that of male N 6 1;

More definitive results were obtained
from recaptures of those first year lizards
that were initially captured soon after theirfirst winter hibernation, in April and May.
Of those lizards 64.5% (20 of 31) wereobserved regularly within a radius of 10 to
50 m from the place of their first capture, insome cases over a period of several years.
First year animals initially occupy small

home ranges of about 10 m in diameter bothoutside the territories of adult males (5 males
and 5 females in Krasnovodsk populationin 1985) and within such territories (4 males

only after their third winter, at an age ofmore than 30 months.

Male Ontogenetic Trajectories
Males participate in reproduction only
after they have acquired a territory. Males
continue to reproduce until the end of theirlives. For example, on the Krasnovodsk
plot male N 3 in 1989 at an age ofapproximately 12 years had a large territorythat included four adult females of different.

If, on the given territory, the only one
female lives, in the case of her
disappearance a holder of territory becomesa widower. He however, subsequently
does not try to search for females outside his
territory. In Gobustan, such case of
widowhood was observed in about 6 years

A

more young male, evidently,
also
turn
out to be a widower.
may

old male.

Vol. 6, p. 104

Asiatic Herpetological Research

June 1995

TABLE 1. Composition and age structure of family groups of the experimental introduced
population at Krasnovodsk and the natural population at Gobustan.

Krasnovodsk
Year
Individual

1986

1987

1988

1989

Gobustan
1990

1986

1987

1988

Vol. 6, p. 105

Asiatic Herpetological Research

June 1995

has selected a male she begins to co-habit
Where there are several
his shelters.

Bfemales on a territory usually the oldest
female cohabits with the male.

Jj
C

.

overlap broadly, especially if the territory
older females exhibit
large. However, someinthe vicinity of theirterritorial behavior
and foraging areas
sites
shelters, basking
older female.
otherwhen approached by
is

D

J* „

l

The home ranges of adult females

1987 1988

1985 1986 1987 1988 1989 1990

I
mmi

territory of theirforegg laying and tofamily groups only

Females leave the1985 1986 1987

1987 1988 1989

1986 1989

CZI2

EE33

VHZ34

A.

Krasnovodsk.

Gorge(spring).
1.

.Kalaligez.
3.

B.

Cr

Gobustan.

C,.

Parkhai

Parkhai Gorge (fall).

D.

2. Second-year lizards.
Adult females.

Yearlings.

Adult males.

4.

1985 ranged from one to five years (mean =2.5
3 years
subsequently over a period of
the neutral areaofresidentwas(1988-1990)
visited at different time by two adultterritorial males (NN 56 and 36). Female N
74 resided at ages of a little less than 4 years
old male N
temporal home range of 3 year
to
on
moved
in
next
territory of
115
year
who
4
N
3
male
e,f)
(Fig.

became a permanent target for her courtshipdisplays.
It isuseful to describe shortly a peculiar

courtship behavior of females addressed bythem towards males, which we regard as atovery important mechanism contributingofmaintenanceandestablishment
personal
bonds between mating partners. At sight ofa male young female moves to him and atonce tries to climb onto his back. A male,as a rule, during first minutes of contact
tolerates these actions of female who isandcrawling over him in different directions
him.
under
crawl
to
insistent
makes
attempts
After that male behaves as if he is inclined tohim andretire, while a female pursues
a behavior is quite
Such
actions.
herrepeats
characteristic of females younger age
2 years
classes, even of those lesser thanold. The behavior retains in older puberal

females, although, contrary to expectation,
almost never occurs prior to actual sexualinteractions, i.e. copulations. Once a femaleitWe

migrate to communal winterdid not observe emigration or dispersal of
females. Over the five year study of the
Krasnovodsk population the time ofresidence of adult females introduced in
shelters.

FIG. 5. Population structure of study populations.

±0.6, n=6).

Family Groups

The mode of sexual relations in rockaagama settlement may be defined as
as
As
many
territorial facultative polygyny.
four females may establish bonds with a
In the natural Gobustan
territorial male.
1.73 females per malewere
population there
the introducedIn
on territories.
Krasnovodsk population the average

number of females per male was increasing
as population structure matured. Over theduration of our study the average number offemales within the family groups was 1.33
(1988), 1.86 (1989) and 2.43 (1990).

The term "family group" is not quite
enters into such a
precise since each female
females. Any
other
from
unit independently

bonds between female

personal or functional
Only
members appear to be absent.
relations between the territorial male andeach of the females may be regarded as
bonds.
of such breeding units isdetermined primarily by an association its

Thestability

Vol. 6, p. 106

Asiatic Herpetological Research

June 1995

TABLE 3. Survival of rock agamas introduced as adults in the Krasnovodsk experimental plot in
May 1985. The number of lizards alive is presented; the number is parentheses is the per centsurviving since the previous year.

YearJune 1995

Asiatic Herpetological Research

The general picture of changes in therelative proportions of immatures, maturesand sex ratio is shown in Fig. 5. It isnecessary to repeat that in 1985 theoriginal colonizing cohort was composed
primarily of immatures, this may haveresulted from capture sampling error. In
1986 the proportion of immatures to adults
was similar but in 1987 and 1988 the ratioof immatures to adults was nearly equal.
Since 1988 adults have predominated.

Comparison with natural populations (Fig.
5) shows that adults predominate thoughrelative proportions may vary significantly

among areas and years. Sex ratio, males
to females,
Table 5).

isgenerally equal (Fig. 5;

Vol. 6, p. 107

Among these lizards 15 (75%) survived to
two years and 13 (65%) to three years ofage.

Of those surviving to three years

were three males and 10 females.
Since the duration of this study wasapproximately half the life span of a rockagama, survivorship in animals older thansix years is based on indirect evidence.

We

estimated the age, based onstandardized size criteria (Zykova andPanov 1991) of adult lizards captured for
introduction onto the Krasnovodsk

experimental plot. Such age estimates areat best imprecise but the adults in thisgroup ranged from three to 1 1 years with
78% being four to six years old (Table 3).

The maximum rate of disappearance

Recruitment(which we attribute to mortality, notSurveys of juveniles were conductedduring the spring (Fig. 5) since few
lizards were active during the hottest

emigration) occurs in the first year (Table
From the second3) after introduction.
adult maleafter
introduction
year

summer andsurvivorship is nearly constant andcomparable to survivorship during the first
The more
six years of life (Table 2).
variable mortality rates for females may be
due to small sample size.

weatherin

Differences in the

early

fall.

numbers of over

wintering juveniles may be attributable toembryonic mortality or hatchling mortality
in the first months after emergence or overthe first winter.

We combined cohorts of lizards ofPost Hatching Mortality and Survivorship

The survivorship of 25 juveniles
into the Krasnovodsk

introduced

experimental plot are presented in Table 2.
We assumed that those lizards thatdisappeared, that is, those not recaptured,
had died. Of the initial 25 hatchlings, 15(10 females, five males) survived tosexual maturity (three years) and of thiscohort four males (80%) and four females(40%) survived six years, to the end of the
The annual survival rate (p x )study.

four, five and six years of age andcalculated survivorship and estimatedsurvivorship to the 9-11 year age class(Table 4).
Ingeneral,survivorship

in

rock

agamas after the first winter following
hatching is relatively constant until theAfter the eighth year
eighth year.
increases.
The maximum
mortality
estimated age of for males was 12-14
years (n=4) and for females it was 9-10
years (n=2).

ranged from 0.72 to 0.83.

Sex Ratio

We believe the high survivorship oflizards in the

Krasnovodsk experimental

plot during the first three years of life arecomparable to natural populations.
tested this by comparing survivorship in acohort of 20 hatchlings, hatched

in

1988

and

sex ratio amongWe

adults does not differ significantly from1986,

even one (Table 5). This relationship was
found at several localities throughout therange and for all localities taken together.

1987, and 1988, that were first captured in

1987,

Although the rock agama social systemis territorial polygyny, the1989 as yearlings.

Vol. 6, p. 108

Asiatic Herpetological Research

June 1995

location and 100

m from the initial capture

point 3 1 months before.

As the population increases, new areas
be pioneered, mainly by young

will

dispersing lizards. Given the sedentary
habits of these agamas we would expect
such expansion to be relatively slow.
Such an expansion occurred in the

introduced population at Krasnovodsk(Fig. 6). From 1985 to 1990 the initialintroduced population occupying an areaof approximately 300 m 2 dispersed into
adjacent areas of approximately 25 ha. Of
the seven subpopulations formed during
period two (Fig. 6, points 1 and 2)
were founded by introduced mature adult
this

FIG. 6. Expansion of the introduced colony at
Krasnovodsk from 1985 to 1990. 1. Unsuitable
habitat.

2.

Boundary of unsuitable habitat.

3.

Principal ridges. 4. Initial introduction site. 5.
Principal study area under regular observation. 6.

Subpopulation established by an introduced adultmale.
7.
Subpopulation established by an
introduced immature male. 8. Subpopulation
established by offspring of introduced lizards. 9.
Single sighting of rock agama. 10. Boundary of
populated area in 1990.

Movement and Dispersion
Rock agamas display a high level of homearea fidelity. Juveniles that were marked
after their first winter were found toremain within 50 m of that location up to
two years or until sexually mature (at twoto three years of age). Some data on thecapture of juveniles before their firstwinter suggests that home site fidelity may

extend from the age of three or fourmonths to the end of life. Four of six
juveniles (66%) marked in October 1988
in the Parkhai Gorge (Western Kopetdag)
were found in the same place the next
The maximum movement ofspring.
immature lizards was not greater than 300
m. A juvenile male first captured in
October 1985 before his first winter wasrecaptured in 1987, 200 m from the first
capture location. He was recaptured in thespring of 1988 as a mature male with afrom the previous capture
territory 250

mlizards after the initial introduction. Twoother subpopulations (Fig. 6, points 3 and6) were founded by lizards that wereimmature when introduced. Finally, threesubpopulations were established by theoffspring of the original introduced
In 1988 (the fourth year after
lizards.
introduction) there were 33 individuals (13adult males, 14 mature females and 10
immatures) in these seven subpopulations.

Individual lizards were observed toleave their home areas only for collective
winter shelter. Females may leave theirhome areas looking for places appropriate
for egg laying (Danieljan

and Grigorjan

1975; Ananyeva and Danieljan 1987).

Conclusions
As it can be seen, the characteristicfeaturesofRock

organization andstability

of

Agama

socialdemography are highbreeding

individuals'

contingent and low population turnover.
This is evident consequence of sedentary
way of life characteristic for males of all
age classes, longevity of these lizards, andpostponed onset of breeding in early
lifetime of young agamas.

The latter may be especially applied to
Although they are capable tomales.

reproduce already at the age of a little
under three years (after their third
wintering), most of them begin to breed,
actually, only at the age of four or fiveJune 1995

Asiatic Herpetological ResearchHow

it may be seen, a male attains
years.
the status of a breeder only after he hadtaken possession of territory of his own.
So each maturing male faces with obvious
difficulties since many features of species'
social organization of settlement (highdensity together with strong territorialityof males retaining control over his homearea until his death) lead to deficiency ofvacancies which might be used by young
male-recruits. Temporary exclusion ofpart of mature (non-territorial) males fromprocess of reproduction may, in principle,
decrease the whole reproductive potentialof the local population.

Acknowledgments

We are indebted to Drs. Mira E.
I.
Vladislav
Vasiljev(Krasnovodsk State Reserve) and Nikolaj
Andreev (Sunt Khasardag State Reserve)

Gauzer,for their help in organization of field workand for participation in catching lizards.

We also wish to express our sincere
thanks to Prof. Ilya

S. Darevsky, Drs.
Natalia B. Ananjeva (Zoological Institute,

Russian Academy of Sciences, St.
Petersburg) and Valentina F. Orlova

The

eggs per mature female during
breeding season- see Ataev, 1985), onlya small number of new-born agamas die.

[Stellio

Even if these losses (especially the latter
overestimated, the analysis of
demographic structure of all demes understudy shows the numerical preponderanceof mature individuals over immature ones.
This is in good agreement with the general
conclusion about the low rate of
figure)is

growthAnother argument

populationinMuseum,(Zoological

final stage of our researchsupported by the Soros Foundation.

was

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To conclude, it may
Rock Agama give us a good example ofbe stated thatlizard species practicing a typical Kstrategy. It was to be expected providing
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pronounced

Moscow

University) for their assistance and help
during our work with museum collections.

Besides such social regulators of
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