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Joint Space Use in a Parthenogenetic Armenian

Rock Lizard (Darevskia armeniaca) Suggests
Weak Competition among...

Article in Journal of Herpetology March 2012

DOI: 10.1670/11-242

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 Joint Space Use in a Parthenogenetic Armenian Rock Lizard (Darevskia
armeniaca) Suggests Weak Competition among Monoclonal Females
Author(s): Eduard Galoyan
Source: Journal of Herpetology, 47(1):97-104. 2013.
Published By: The Society for the Study of Amphibians and Reptiles
DOI: http://dx.doi.org/10.1670/11-242
URL: http://www.bioone.org/doi/full/10.1670/11-242

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Journal of Herpetology, Vol. 47, No. 1, 97104, 2013
Copyright 2013 Society for the Study of Amphibians and Reptiles

Joint Space Use in a Parthenogenetic Armenian Rock Lizard (Darevskia armeniaca) Suggests
Weak Competition among Monoclonal Females

EDUARD GALOYAN

Zoological Museum of Moscow University, Moscow 125009, Russia; E-mail: saxicola@mail.ru

ABSTRACT.According to kin selection theory, competition among monoclonal animals must be lower than between unrelated individuals.
Thus, we propose that home range and core area overlap in parthenogenetic lizards will be broader than female range overlap in similar
gonochoristic species. To test this hypothesis, we examined home range locations and space use of parthenogenetic Armenian Rock Lizards
(Darevskia armeniaca) and compared them with home range locations and space use of gonochoristic species. We demonstrated that
parthenogenetic Rock Lizards have a total range structure typical of insectivorous lizards, consisting of a sally zone, a home range, and one or
more core areas. Some core areas contained activity centers, associated strongly with key basking sites and shelters. Provisional residents were
found within the same range for 1 or 2 years, whereas wanderers visited study sites for 1 or 2 weeks per season. Settlement structure varied
greatly among years. Home ranges, core areas, and even activity centers and basking sites of parthenogenetic females overlapped extensively,
unlike in females of nonparthenogenic species. Monoclonal origins and high level of relatedness within unisexual species are possible
explanations of the extended range overlap among parthenogenetic females.

Animal space use is connected to energetic losses and gains
and is determined largely by reproductive strategy, a strategy
that is different in males than in females (Waser and Wiley, 1979;
Perry and Garland, 2002). Female reproductive success depends
on resource distribution, and male reproductive success is
limited primarily by access to potential mates (Davies, 1991;
Stamps, 1997b; Baird et al., 2003). Intrasexual home range
overlap is common in males; however, territorial males have
exclusive use of defended space (Brattstrom, 1974; Stamps,
1977a; Perry and Garland, 2002). Although range overlap is
more common in females, even nonterritorial females often
defend small parts of their home ranges, such as basking,
foraging, and nesting sites (Evans, 1951; Boag, 1973; Perry and
Garland, 2002). Thus, intrasexual aggression leading to space
separation is determined by competition for important resourc-
es (Stamps, 1977b; Perry and Garland, 2002).
According to kin selection theory, competition among
genetically similar relatives must be lower than competition
between unrelated individuals (Hamilton, 1964; Wilson, 1975).
This relationship suggests that range overlap among relatives
should be greater than that among more genetically different
individuals. The influence of kinship on association patterns
among females has been documented for a wide range of
mammalian species (Dunbar, 1980; Packer et al., 1991; Hole-
kamp et al., 1997; Archie et al., 2006) and in long-living
viviparous Shingle-back Skinks (Tiliqua rugosa) (Bull and
Baghurst, 1998), a species with motheroffspring recognition
and prolonged association (Main and Bull, 1996). If this
suggestion is true and degree of relatedness plays a significant
role in animal spacing, then monoclonal populations descend-
ing from a unique ancestor should demonstrate a high level of
joint space use. Although populations of this sort are not found
in mammals and are rare in birds (Schut et al., 2008), they are
common in several genera of lizards (Darevsky, 1958; Mittwoch,
1978; Cole and Townsend, 1990). Unisexual lizard species
demonstrate extremely low genetic variability within species
(MacCulloch et al., 1997; Fu et al., 1998; Kearney et al., 2009) or
within populations if the asexual lineages emerged several
times (Murphy et al., 1997; Fu et al., 2000).
Diurnal oviparous heliothermic lizards are suitable subjects
for studies on internal range structure, spatial distribution, and
DOI: 10.1670/11-242
social behavior (Brattstrom, 1974; Stamps, 1977a,b; Regal, 1983;
Stamps and Krishnan, 1998; Fox et al., 2003; Chapple and
Keogh, 2005). Lizards demonstrate a wide variety of sociosexual
relations and show complexity of space use, comparable to that
of birds or mammals (Baird et al., 2003; Chapple and Keogh,
2006; Kerr and Bull, 2006). The behavior of unisexual Whiptail
Lizards (Cnemidophorus spp.) has been studied extensively
(Crews and Fitgerald, 1980; Crews et al., 1986; Crews, 1987,
1998), but nearly nothing is known about spacing patterns of
unisexual lizards. Only one study was partly devoted to this
topic (Trofimov, 1981).
Parthenogenetic lizards have been shown to be less aggres-
sive than females of closely related bisexual species (Bracken,
1978; Trofimov, 1981; Leuck, 1985; Grassman and Crews, 1987).
Population densities of unisexual Rock Lizards (Darevskia spp.)
can be an order of magnitude higher than the densities of
bisexual relatives under the same conditions (Darevsky, 1967;
Danielyan, 1971). The low aggressiveness and high population
density in parthenogenetic species suggest widespread joint
space use. We therefore predict that the home ranges and core
area overlap in unisexual lizards must be greater than that in
females of related gonochoristic species.
To test this hypothesis, we described home range and
settlement structure in the parthenogenetic Armenian Rock
Lizard, Darevskia armeniaca. We also determined the extent of
home range overlap, described space use differences in females
of different ages, and described space use strategies. Female
range structure and settlement social organization have been
well studied in related gonochoristic species of the same genus
living in the similar biotopes (Tsellarius and Tsellarius, 2001,
2005, 2006). Thus. we have an opportunity to compare female
home range structure and settlement organization among
bisexual and unisexual Rock Lizards.
Rock Lizards of the genus Darevskia (Arribas, 1999) include 20
bisexual (caucasica, saxicola, and rudis clades) and seven
parthenogenetic species (Darevsky, 1958; Arribas and Carranza,
2007). These lizards are common in the Russian Caucasus and in
Georgia, Abkhazia, Azerbaijan, Eastern Turkey Armenia,
Nagorno-Karabakh, northwestern Iran, and Turkmenia (Darev-
sky, 1967). All of the parthenogenetic Rock Lizards are the
product of hybridization of some closely related gonochoristic
species and appeared <10,000 yr ago in the conditions caused
by the last period of glaciation (Darevsky, 1966).
98 E. GALOYAN
The Armenian Rock Lizard, D. armeniaca (Meheli, 1909), is a
small lacertid (adult snoutvent length [SVL] = 5070 mm),
diurnal, insectivorous, parthenogenetic lizard that is widely
distributed in the Transcaucasian region (Darevsky, 1967). It
originated from hybridization between D. valentini (rudis clade)
and D. mixta (caucasica clade; Murphy et al., 2000). Armenian
Rock Lizards occupy large rocky formations in alpine meadows
and forests, large cliffs, and buildings situated up to 2,500 m
a.s.l. (Darevsky, 1967). These small, flattened lizards can easily
climb rocks, stones, and trees. Rock Lizards are oviparous. No
postnatal parental care is presented (Darevsky, 1967). Although
some populations of Armenian Rock Lizards contain triploid
males, a result from backcrosses with males of the parental
species (Darevsky and Danielyan, 1968; Darevsky et al., 1978;
Darevsky and Kupryanova, 1982), normally there are no males
in populations of this parthenogenetic lizard.
MATERIALS AND METHODS
Data were collected in May, June, and July 20082010 (mainly
in 2009 and 2010) in northern Armenia. Observation times
included the oviposition period (1016 June 2009 and 22 June1
July 2010). Fieldwork took place in Dilijan National Park on the
southern slope of Ijevan Mountain ridge (40845 0 45N; 44853 0 44E;
1,280 m a.s.l.; datum WGS 84). We chose a study area of 2,500
m2 in oak (Quercus orientalis) and hornbeam (Carpinus orientalis)
forest that also had beech (Fagus orientalis) trees situated on the
lower part of the watershed on the southern exposed slope.
Crowns of oaks and hornbeams were close to each other so the
sunlight falling on the surface of the ground formed basking
spots of light during the day. Dog rose (Rosa spinosissima)
shrubs, blackthorn (Prunus spinosa), and oriental hornbeam
grew in the illuminated parts of the study area. In some areas,
ferns and forest grass also were represented.
The surface of the study area was covered by foliage, growing
up to 1520 cm. A forest brook flowed in the bottom of the
valley between the slopes of two watersheds. We constructed a
map of the area with all the primary landmarks (trees, large
stones, rocky formations, roots, and branches) and then overlaid
a virtual grid (with the length of each square side 0.5 m in the
scale of the map).
Stones with an average size of 0.71.0 m2, large rocky
formations, and branches randomly covered the surface of the
study area. Such elements of the substrate occupied nearly
10.5% of the surface area and contained tree litter (primarily
leaves), the main habitat of insects and other invertebrates. The
distance (mean 6 SD) between such microbiotopic objects was
1.3 6 0.90 m. All of these microbiotopic elements affected the
quality of the biotope and were used by the lizards as basking
sites. Night refuge sites were situated under stones, roots, and
tree cavities as well as between stones and in the cracks of the
stone formations. In some cases, lizards used rodent burrows as
shelters.
Lizards within the study area were caught and marked
intensively at the beginning of the study, but they continued to
be marked during the entire study period from 2008 to 2010.
Overall, 75 Armenian Rock Lizards in total were individually
marked. Most of the lizards were caught during the first week of
each observation season. SVL and tail length (TL) of each
animal were measured to within 0.5 mm. Animals were marked
individually by amputating a portion of the distal finger
phalanx (a permanent mark) and by attaching one or two color
beads (a more temporary mark) to the skin on the lizards back
with nylon thread. Because these lizards squeeze through
narrow rock passages, they sometimes lose bead marks. These
two marking techniques do not influence the behavior or
survival of the lizards (Fisher and Muth, 1989; Tsellarius and
Tsellarius, 2001; Husak and Fox, 2003; Nicholson and Richards,
2011). Postamputated finger bones were later used for the
skeletochronological method of age determination (see below).
Visual observations of individually marked lizards within the
study area were performed daily during the study period when
lizards were active. These observations were made between
1100 and 1700 h unless the weather was rainy or cloudy. The
total observation time was 231 h. Observations were made from
a distance of 510 m from the lizards. During observations,
we used a field glass lens (BPC 830). Regular monitoring of the
entire study area was performed routinely, and all observed
lizards were recorded in the field notebook. Location, move-
ment, and activity type of randomly chosen animals were
recorded and described in a field notebook. Later, the
observations were transmitted into registration points, and
the representative dots were added to the map of the scheme
of the study area (within 0.5 m). We defined two kinds of
registration points (Adams and Davis, 1967; Tsellarius and
Tsellarius, 2005): milling points and moving points. Milling
points were put on the study area scheme if a lizards activity
was within the borders of one 0.5 0.5 m2. Milling points are
associated with basking, resting, or foraging activities. Each
moving point was put on the scheme if the lizard crossed the
borders of one 0.5 0.5-m square. An average moving lizard
crossed one 0.5 0.5-m square once per minute.
We define several components within a range (Fig. 1). The
entire area occupied by a lizard was its total range, consisting
of its home range surrounded by a peripheral area called the
sally zone (Linn and Key, 1996). We defined total range
operationally using the 100% minimum convex polygon (MCP)
(Mohr, 1947). We applied the term home range to the zone that
was intensively used by the animal and where normal activity
(e.g., basking, foraging, social interactions, exploiting move-
ments) took place (Kaufmann, 1962; Jewell, 1966; Schoener and
Schoener, 1982). From time to time, lizards made excursions into
the sally zone. Excursion movements were characterized by
rapid movement, with frequent stops and by attention poses
(stances with straightened forelimbs and a raised head). The
distance of an excursion movement was always much longer
than movements within the borders of the home range.
We defined core areas within the home range using the
algorithm of Samuel et al. (1985). We compared the observed
space-use patterns in all 0.5 0.5 m squares, where registration
points were, with those expected from a uniform distribution.
Using the same technique, we defined activity centers within
core areas. One or several core areas, connected by routes, could
be used by one lizard during the period of observations. To
better understand the activity distribution within different parts
of the range, we analyzed the ratios of milling and moving
points in sally zone, home range, core areas, and activity
centers.
To determine the minimum number of registration points
needed for home range structure estimation, we used cumula-
tive graphs of range area against the number of registration
points (Waser and Wiley, 1979; Harris et al., 1990). We found
that 300 registration points were enough to describe 80% of the
seasonal home range size and structure (Rose, 1982). However,
the number of observation points per lizard for one observation
season ranged from nearly 500 to >1,000. Based on these
 SPACE USE IN PARTHENOGENETIC ARMENIAN ROCK LIZARDS
FIG. 1. Diagram of a typical lizards range (adult female in 2009;
home range area, 182 m2) illustrating the terminology used in this paper.
numbers, the number of estimated seasonal home ranges was
18.
We ultimately determined the age of 15 lizards in the study
group. Accurate age determination of young animals (13 yr
old) was based on SVL measurements. Age determination of
older animals was based on the sceletochronological method
(Smirina, 1974; Castanet, 1994). The effectiveness of the
sceletochronological method was shown for estimating age in
numerous species of Darevskia (Arakelyan and Danielyan, 2000;
Arakelyan, 2002; Tsellarius and Tsellarius, 2009) and in other
lizard species within the Lacertidae family (Kolarov et al., 2010).
Amputated digits for individual marking were used for this
procedure. Each dried finger phalange was washed in distilled
water, and then all associated tissue was removed. The bones
were then decalcified in 5% nitric acid and washed under
running water. Microscopic sections (1015 lm) of digits were
made and colored by Erlichs hematoxylin (Castanet and
Smirina, 1990; Castanet, 1994; Tsellarius and Tsellarius, 2009).
Permanent preparations were then made and analyzed using a
Reichert Wien (circa 1906) microscope. Photographs were taken
using a Nikon D 90 camera. Year rings on the sections of each
phalange bone were counted to estimate age. Young Rock
Lizards appear in the study site in the beginning of autumn
(Darevsky, 1967). A juvenile ring can be seen on the sections
of digits after lizards survive their first winter (Fig. 2). Each new
winter hibernation adds one more ring to the bone structure
(Smirina, 1974; Tsellarius and Tsellarius, 2002). The average life
span of bisexual and parthenogenetic Rock Lizards is approx-
imately 56 yr (Arakelyan and Danielyan, 2000), whereas some
individuals live nearly 1314 yr (Tsellarius and Tsellarius, 2009).
Female Rock Lizards lay their first clutch after their third winter
hibernation, when body length reaches 5060 mm (Darevsky,
1967; Tsellarius and Tsellarius, 2009). We classified young
animals that survived two or three winters as subadults and
classified older lizards as adults.
The study area map, registration points, and home range
structures were analyzed in MapInfo 6.5., ArcView 3.2. We used
Microsoft Excel and Statistica 7.0 for statistical analyses. Adobe
Photoshop 7.0 and Corel Draw 12 were used for graphic works.
99
FIG. 2. Microscopic section of the Armenian Rock Lizard finger
phalange. j = juvenile ring. The numbers with arrows indicate winter
hibernation periods.
To determine the monopoly ratio of core area use, we counted
the number of lizards with overlapping core areas and then
calculated the extent of overlap of the core area of each
individual with other core areas (as a percentage of the core
area). For measuring relative exclusiveness of activity centers,
we determined the monopolizing activity center coefficient
(MACC), calculated as a ratio (all subject milling registration
points in a lizards activity center divided by the milling
registration points of all lizards including the subjects in the
same area). So, this value represents the ratio of the time spent
by the animal in its activity center and the time spent there by
the other animals (Waser and Wiley, 1979). This ratio varies from
1 when the owner of the activity center had exclusive use of this
area to 0 if the subject never used the location, but others did.
The Mann-Whitney U-test was used for comparison of the
MCP ranges between different age cohorts and between females
with different strategies of space use (see below). This test also
was used for comparison of the moving ratios (moving
registration points/(moving + milling registration points). The
ratio was taken for each lizard and then the ratios were
compared. We also used the Mann-Whitney U-test for compar-
ison of monopolizing activity center coefficients. We used v2
tests when comparing frequency distributions of the number of
overlapping home ranges, core areas, and activity centers
between years. All mean numbers are given with the SD. We
used Spearmens rank correlation coefficient (rs) to measure
statistical dependence between two empirical samples. The null
hypothesis of all the statistical tests was accepted at the 0.05
significance level.
RESULTS
We distinguished females with three main space use
strategies: 1) provisional residents (10 subadults and 10 adult
animals), lizards that were observed within the borders of the
study area for not less than one observation year. Home range
structures of provisional residents were stable; 2) wanderers
100 E. GALOYAN
TABLE 1. Seasonal range areas (mean 6 SD).
Female Total range (m2) Home range (m2)
Adult 174.0 6 87.51 104.8 6 35.23
provisional (min = 86.0, max = 364.9) (min = 46.3, max = 145.2)
residents
Subadult 196.0 6 90.43 131.1 6 71.36
provisional (min = 63.5, max = 353.7) (min = 63.5, max = 273.8)
residents
Wanderers 85.5 6 65.64 37.3 6 4.68
(min = 39.0, max = 160.6) (min = 32.1, max = 41.0)
(three subadults and seven adult animals), lizards that were
observed within the study area for 1 or 2 weeks per year and
then left; such animals could appear within the study area each
year and they had proper pattern of space use; and 3) passers-
through, lizards noticed within the study area only once for
several hours. We cannot say anything about their space use
strategy.
Subadult provisional residents often used the same home
ranges in two consecutive years. Adult provisional residents
never used the same home range for more than 1 yr.
Transformation from provisional resident to wanderer and vice
versa occurred in adults lizards.
Seasonal home ranges of provisional residents differed
among individuals, but there was no difference between home
ranges of subadult and adult animals (Mann-Whitney U-test, P
> 0.05) (Table 1). Home range area correlated positively with the
number of core areas (rs = 0.638, P = 0.065). Provisional
residents had an average of 2 6 0.8 core areas (varied from one
to four, n = 18). Activity centers were found near shelters and in
sites that were suitable for basking. Each activity center was
only a small component of the core area (0.30.5 m2). The
number of activity centers of different lizards varied from two to
eight, but the total sum area of these centers within the home
range only differed weakly (Table 1). The mean home range size
of wanderers was smaller than in provisional residents (Table 1).
Comparing the two adjacent years (2009 and 2010), we found
no significant differences in mean total ranges, home ranges,
core areas, or activity center sizes for any of the three classes of
lizards: subadults, adults, or provisional residents (Mann-
Whitney U-tests, P > 0.05). The exact borders of the total range
of a particular provisional resident could be very different from
one year to the next. The configuration of the home range and
core areas also often changed. However, if the provisional
resident stayed within the study area, the main cores and
activity centers remained the same. Mean core areas and activity
center overlaps of the same lizard in two adjacent years were
53.9 6 11.97 and 37.8 6 18.20%, respectively.
We recorded more moving points than milling points both
within the sally zone and inside the home range, and the
difference in moving ratios within these two areas was
significant (Mann-Whitney U-test, U = 66.5, Z = 2.687, P <
0.01). Most movements within the home range involved
relocation from one core area to another. Milling (especially
basking and foraging) prevailed in core areas and in activity
centers (Fig. 3). The difference in the ratio of moving points to
milling points within core areas and activity centers was also
significant (Mann-Whitney U-test, U = 54.0, Z = 3.117, P <
0.01). If a lizard passed through the activity center, it usually
stopped there for several minutes or stayed for tens of minutes.
Home range overlap among provisional residents of different
ages could be very large. In some cases, the whole home range
Core areas (m2) Activity centers (m2) n
5.1 6 2.91 1.7 6 1.27 8
(min = 1.3, max = 9.3) (min = 0.5, max = 4.5)
8.0 6 4.12 2.3 6 1.28 10
(min = 3.3, max = 15.3) (min = 0.8, max = 4.8)
4.2 6 0.90 1.3 6 0.57 3
(min = 3.25, max = 5.0) (min = 1.0, max = 2.0)
of one lizard was within the borders of anothers home range.
Mean overlaps of the core areas and activity centers were >50%
(Table 2). Only one lizard (in 2010) had an exclusive activity
center, located near the border of a study place. We did not find
a significant difference in the areas that overlapped between
two adjacent years (Mann-Whitney U-test, P > 0.05), and we
found no difference in the number of overlapping areas or in the
percentage of area overlap between subadults and adults
(Mann-Whitney U-test, P > 0.05). We also did not find
differences in the number of overlapping home ranges (one),
core areas (two), or activity centers (two) between 2009 and 2010
(v21 = 2.33, v22 = 2.66, v23 = 6.07; P > 0.05), though the number
of provisional residents with overlapping home ranges in 2010
was larger: nine vs. seven in 2009 (Table 2).
Unfortunately, we do not have enough information about
most wanderers; therefore, we cannot include them in the
statistical analyses, but it is clear that they used the same space
with provisional residents. Their temporary home ranges
overlapped with each other, and these lizards formed activity
centers in the same sites as each other and as the provisional
residents (Fig. 4).
MACCs for adult and subadult provisional residents were
nearly the same (Table 3; Mann-Whitney U-test, P > 0.05) and
often <0.5. These coefficient values show that parthenogenetic
Rock Lizards use joint activity centers. We have often seen
animals basking on the same stone or even on one another,
sometimes with more than two animals in proximity. The
degree of activity center monopolization negatively correlated
with the number of overlapping activity centers (rs = -0.70, P <
0.01). Even if a particular activity center did not overlap with
FIG. 3. Moving ratio values (mean 6 SE) in different parts of the
range given for 18 lizards.
 SPACE USE IN PARTHENOGENETIC ARMENIAN ROCK LIZARDS 101

TABLE 2. Number (mean 6 SD) of home ranges and core areas overlapping among provisional residents and percentage overlap (mean 6 SD) of
their home ranges and core areas (20092010).

No. of home Home ranges No. of core Core areas No. of activity Activity centers
Year ranges overlapping percentage overlap areas overlapping percentage overlap centers overlapping percentage overlap n

2009 6 6 0.8 93.7 6 6.14 4 6 0.9 55.1 6 6.46 2 6 1.0 51.9 6 29.16 7
2010 7 6 1.7 96.7 6 3.76 4 6 0.9 60.3 6 11.17 3 6 1.4 57.6 6 27.14 9

the others, the MACC value (mean 6 SD) was 0.64 6 0.05 (min support this observation (Table 3). Wanderers come to the
= 0.26, max = 0.92; n = 18) because wanderers and passers- settlement for a short time and their temporary home ranges
through often visited this area. Wanderers that came to the overlap with the home ranges of provisional residents. They
study site formed their activity center in the same place as also use the same basking sites, shelters, and activity centers
provisional residents, so their core areas overlapped substan- (Fig. 4) and become a part of the settlement (returning there in
tially (Fig. 5). Several wanderers and provisional residents often different years). Though passers-through never stay for long
used the same shelter and basking sites at the same time. periods, they also use joint basking sites and shelters. Core area
and activity center overlap among parthenogenetic Armenian
DISCUSSION Rock Lizards explains the high population densities in this
Because of their high parental investment, female activity species. Nothing similar is known for bisexual and unisexual
depends strongly on the abundance and distribution of Whiptails (Aspidocelis [Cnemidophorus] spp.) (Clark, 1976;
resources and conditions; so, female home range structure is Cuellar, 1979; Bennet and Gorman, 1979; Warner, 2000) or
defined by the placement of foraging sites, shelters, basking Scaly-toed Geckos (Lepidodactylus) (Hanley et al., 1994). Hud-
sites, and places suitable for oviposition (Waser and Wiley, 1979; dling behavior in communal hibernacula of numerous bisexual
Davies, 1991; Baird et al., 2003). This explains why the range species, including Rock Lizards, is noted in the spring, after
structure of the parthenogenetic Armenian Rock Lizard is quite winter hibernation, or in autumn before an inactivity period,
typical for insectivorous lacertid lizards (Diego-Rasilla and but never in summer (Darevsky, 1967; Trofimov, 1981; Woodley
Perez-Mellado, 2003). Their routine activity usually takes place and Moore, 1999). This phenomenon may be explained by the
within small components of home range: core areas and activity
high intrasexual aggression level during the breeding season
centers, covering the most profitable microbiotopes (large
and low aggression level at other times (Ruby, 1978; Woodley
stones, rock piles, rocky formations, and tree roots) (Fig. 6).
and Moore, 1999).
Home range size and range structures of females in gonochor-
istic Rock Lizard species are the same as in parthenogenetic
Armenian Lizard (Trofimov, 1981; Tsellarius and Tsellarius,
2005) (Fig. 1).
Home range structure of provisional residents was stable
during one season, but most adult lizards changed their home
ranges, whereas the ranges of subadult provisional residents
lasted for two or three winters. Even old animals (>45 yr old)
never became permanent residents and used the same range for
>2 yr. Thus, general settlement structure was variable.
Populations in which the home ranges of subadult animals
are more stable than those of adults are uncommon (Schoener
and Schoener, 1982). We suggest that the low level of range
stability in provisional resident individuals is caused by features
of social structure of this population of parthenogenetic species.
Space use patterns in males and females are often determined
by intersexual communications and relations (Evans, 1951;
Panov and Zykova, 1995, 1999; Kerr and Bull, 2006; Tsellarius
and Tsellarius, 2006). Some long-living gonochoristic species
demonstrate low immigration rates, high level of population
stability for many years, and constant home range structure. It
seems that female attachment to the same home range in
bisexual species is often formed by intersexual relations with
males (Panov and Zykova, 1995; Tsellarius and Tsellarius, 2005;
Kerr and Bull, 2006). We propose that unstable range structure
in parthenogenetic lizards is caused by the absence of necessary
and prolonged relationships among females.
Our study demonstrates that the nucleus of Armenian Rock
Lizard settlement structure is formed by provisional resident
home ranges, core areas, and activity centers that overlap
among individuals. Parthenogenetic lizards often use these
areas jointly at the same time. High ratios of monopolizing FIG. 4. Core areas and activity centers convex polygons of five
activity center coefficients among different categories of females provisional residents and two wanderers in 2010.
102
TABLE 3. MACC values (mean 6 SD).
MACC
Adult provisional residents 0.40 6 0.040
Subadult provisional residents 0.33 6 0.044
Wanderers 0.22 6 0.220
Intersexual home range and core area overlap is common
(Stamps, 1977a; Regal, 1983; Christian, 1986; Nicholson and
Richards, 2011), especially in lizards with long-term intersexual
relations (Tinkle et al., 1962; Panov and Zykova, 1999; Chapple,
2003; Chapple and Keogh, 2005; Kerr and Bull, 2006), including
bisexual species of Rock Lizards (Tsellarius and Tsellarius, 2005,
2006). However, intrasexual core area overlap is not typical.
Usually, core areas are used exclusively by one male and one
female (Stamps, 1977a; Regal, 1983). Even if the female does not
FIG. 5. Number of activity centers (n = 75) and MACC. The line
indicates locally weighted scatterplot smoothing (LOWESS).
FIG. 6. Distribution of registration points on the part of study area in
2010.
E. GALOYAN
Limits No. of activity centers
min = 0.040, max = 0.87 41
min = 0.023, max = 0.92 30
min = 0.04, max = 0.68 9
have a defined territory (sensu Burt, 1943), she will often defend
a basking site (Boag, 1973). Spatial separation is usually
explained by the competition for food (in females) and for
mates (in males, rarely in females) (Stamps, 1977b; Jaeger et al.,
1982; Gowaty and Wagner, 1988). Even nonterritorial females
limit intrasexual access to core areas (Evans, 1951; Brattstrom,
1974; Waser and Wiley, 1979). Joint space use in parthenogenetic
Armenian Rock Lizards indirectly demonstrates the absence of
competition that may lead to spatial separation. Unfortunately,
nothing is known about differences in space use of other closely
related species of parthenogenetic and gonochoristic species.
However, relatively low aggressiveness level in unisexual
Whiptail Lizards (Bracken, 1978; Leuck, 1985) also may suggest
possibility of joint space use. Thus, our prediction that
monoclonal origins and high relationship levels within parthe-
nogenetic species may lead to low competition levels can be
supported by the results of this study.
Acknowledgments.I thank A. and E. Tsellarius for collabora-
tion and help in preparing the article. Many thanks to A. Tiunov
and P. Freed for commenting on earlier drafts. I thank A.
Paterson for valuable recommendations and editing of the
manuscript on the last stages. I thank two anonymous reviewers
for valuable comments on the manuscript. Also, I am very
thankful to my friends, who helped me in the fieldwork
organization: H. Ter-Voskanyan, M. Ter-Voskanyan, N.
Asatryan, G. Papov, M. Arakelyan, and T. Hayrapetyan. Many
thanks to the administration and staff of Dilijan National Park,
especially to the director, A. Davtyan, for access and assistance
in the national park. The study was supported by Russian
Foundation for Basic Research, project 12-04-31214.
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Accepted: 20 March 2012.