Social organization of woodchucks (Marmota monax)

Summary The social organization of woodchucks (Marmota monax) in southeast Ohio was studied at two sites, at one for two 2 years (1979–1980) and the other for 3 years (1981–1983). Spatial organization was determined by trapping and radio tracking. The home ranges of adult females did not overlap in the early spring but during late spring and summer there was some overlap (<10%) as females expanded their home ranges. Adult females tended to occupy the same home range in consecutive years. Some adult males occupied well-defined home ranges that did not overlap the home ranges of other males but did overlap extensively the home range of one to three adult females. These males tended to occupy the same home range in consecutive years. Infants used the same home range of their dam until about 2–3 months of age when most males and females apparently dispersed. About 35% of the juvenile females did not disperse until their second spring, just before their mother's new litter first emerged from their burrow. The average social group consisted of an adult male with two female kin groups comprising an adult female, an offspring (usually female) of the previous year, and the young of the year. Interactions within the kin group and with the adult male were relatively frequent and generally amicable. Interactions between kin groups both within and between different social groups were relatively rare and agonistic. The social organization of woodchucks in Ohio differs from that described in previous studies of woodchucks elsewhere and from that predicted by current models proposed by others on the evolution of social organization of marmots.     

Spatial organization and mating system of Microtus townsendii

Summary Space use by individual Townsend's voles, Microtus townsendii, was investigated in spring and summer by means of radiotelemetry and intensive live trapping in undisturbed grasslands near Vancouver, British Columbia. Home ranges of males were larger than those of females; females had significantly larger ranges in spring than in summer. Most males and females maintained territories free of individuals of the same sex in spring. Male-female pairs had their exclusive territories closely overlapping each other. The 1:1 operational sex ratio and the spatial association of pairs of males and females suggest that the voles were monogamous in the spring of 1988 and that 50% of the males were monogamous in the spring of 1989. In summer, there was more intrasexual overlap between home ranges of males and females and female ranges were considerably smaller than those of males. Females were more philopatric than males and females thought to be members of the same family group lived adjacent to each other or had overlapping home ranges. Males overlapped with more than one female in summer, but most females still overlapped with only one male, which suggests that the mating system is polygynous in summer. Thirty-five percent of the philopatric females became pregnant for the first time when the male spatially associated with their mother in the spring was still alive and thus could potentially have mated with their fathers. Male and female territoriality in spring is the proximate mechanism for the limitation of breeding density by spacing behaviour.[/p] Offprint requests to: C.J. Krebs     

Dynamics of space use and male vigour amongst wood mice,Apodemus sylvaticus,in the cereal ecosystem

The spatial organisation of male and female wood mice,Apodemus sylvaticus, was investigated in a large-scale radio-tracking study on arable farmland near Oxford, United Kingdom, during the breeding season. Both sexes had significantly larger home ranges in the breeding season than at other times, and the breeding season home ranges of male (X = 1.44 ha) were significantly larger than those of females (X = 0.49 ha). Home range overlap was significantly greater between males, and between males and females, than it was between females. Overlap between males tended to be greatest in heavily utilised areas. Except during sexual consortship, there was minimal evidence of dynamic interaction among individuals. Home range sizes of breeding males varied widely, as did their body weights. There was no relationship between male body weight and home range size or any other movement parameter. However, males with the largest home ranges had the highest scores on all other movement parameters, indicating that they expended more energy in movement. These more vigorous males had access to the home ranges of more females than did males with small home ranges.     

Sexual dimorphism and the mating ecology of polar bears (Ursus maritimus) at Svalbard

We assessed the role of size, mass, and age in mating and non-mating polar bears (Ursus maritimus) at Svalbard, Norway, during the spring breeding season. The ratio of male to female mass, in male-female pairs, ranged from 1.00 to 3.02 ([`(x)] = 1.99 \overline x = 1.99 ) indicating that mating males were larger than mating females but with substantial variation. Paired males were older than unpaired males and male mass was related to age. However, males paired with females were not significantly different in body mass from those males caught alone. Wounds and scars resulting from fights between males began at about 6 years of age and peaked at about 17 and 20 years of age, respectively. The frequency of broken canines in males, presumably due to increased male-male conflicts, increased with age but showed little increase in females. The wide range of male size in male-female pairs and the age-related signs of injury suggest that male polar bears engage in both scramble competition and contest competition for access to breeding females. The mating system of polar bears is variable but is best described as female defense polygyny or serial monogamy.     

Lek promiscuity in a semelparous mammal,Antechinus stuartii (Marsupialia: Dasyuridae)?

Summary All male Antechinus stuartii die following a brief rut towards the end of their first year of life. Outside the rutting period, both males and females forage in clearly-defined invidual home ranges, but neither sex is territorial. As temperatures drop during the winter, both sexes may leave their foraging range each day to spend the latter part of the night and most of the daylight hours in communal nests which may be used by fifteen animals simultaneously, and over forty individuals during one winter. These temporary movements away from the foraging range cause an increase in the daily home range revealed by radiotelemetry. When the mating season commences, males abandon their foraging range and aggregate in a few of the communal nest trees, where they spend most of the night. There is some movement between aggregations, but the movement appears to be rapid and direct. Females continue to use their foraging area, and nest solitarily. However, they make excursions to the male aggregations, and spend time in the nests with the males. By the third week of the mating season, immediately prior to male mortality, males move frequently from one aggregation to another, but are still confined to the aggregations and direct movements between those nests. By this time some females have ceased visiting the aggregations and nest solitarily within their foraging range. These data suggest that the mating system of A. stuartii is lek promiscuity, and support the hypothesis that the abrupt mortality of males is a cost of endocrine changes which facilitate gluconeogenic mobilisation of body protein, allowing the males to sustain their vigil at the leks.     

Spacing of leks in relation to female home ranges,habitat requirements and male attractiveness in the great snipe (Gallinago media)

Summary In an attempt to examine three main hypotheses on the evolution of leks, data on female home ranges, distance between leks and male site fidelity in the great snipe (Gallinago media) were obtained. In a 30 × 40 km area in central Sweden, the locations of 12 leks were identified. Six were within the study area (7 × 11 km) in which probably all leks were known. In the study area, nearest neighbor distances between leks were longer than the predicted drawing area (diameter of female home ranges plus the detection range of leks). Furthermore, in only 1 of 13 radio-tracked females did the estimated home range enclose many leks. These data do not fit the prediction from the hotspot hypothesis on the dispersion of leks, namely, that the distance between leks should be less than the drawing area and hence an average female home range should enclose more than one lek. The observed distance between leks was about the same as predicted by the female preference hypothesis. This hypothesis also predicts females visit mainly one lek; however, we found they sometimes visit two and nest close to a third. We propose that males may first settle according to hotspot rules, but females will resettle according to preferences for certain males and/or larger leks. In this way some hotspot leks are abandoned, and the distance between leks is increased as males become more tightly clumped. However, when all males and females have settled, the leks still existing would be on hotspots. In the female preference hypothesis, males are assumed to aggregate on leks because females prefer clustered males as they can be more easily compared than if they are dispersed. Alternatively, in the attractiveness hypothesis, it is suggested that females prefer certain males, and hence unattractive males surround the attractive are forced to join leks in order to come close to females. Our data on male site fidelity support the attractiveness hypothesis because successful and dominant males return to the same lek and territory, both within and between years, whereas unsuccessful males move to other leks. Attractiveness of certain males may explain why according to hotspot rules in this species males are more clumped than expected. Offprint requests to: J. Höglund     

Ranging patterns of bottlenose dolphins living in oceanic waters: implications for population structure

Very little is known about the ecology of common bottlenose dolphins (Tursiops truncatus) living in oceanic waters. This study investigated the ranging and residence patterns of bottlenose dolphins occurring in the Azores (Portugal), the most isolated archipelago in the North Atlantic. Data were collected during standardized boat-based surveys conducted over a 6-year period in an area of approximately 5,400 km² (main study area). To investigate the extent of movements of individual animals, non-systematic surveys were also conducted outside this area. Only 44 individuals out of 966 identified were frequently sighted within and between years. The remaining individuals were either temporary migrants from within or outside the archipelago, or transients. Resident dolphins showed strong geographic fidelity to the area. Long-distance movements (of almost 300 km), consistent with foraging or exploratory trips, were observed among non-resident dolphins. Home range size was estimated for 31 individuals sighted ≥10 times. Range areas of these dolphins varied in size and location, but considerable overlap was observed in the areas used, suggesting the absence of habitat partitioning between resident and non-resident dolphins. Estimates of home range size of bottlenose dolphins in the Azores were found to be considerably larger than those previously reported for this species. It is hypothesized that dolphins living in the Azores carry out extensive movements and have large home ranges in response to the lower density and patchy distribution of prey compared to other areas. The extensive ranging behaviour and the lack of territoriality provide an opportunity for interbreeding between dolphins associated with different islands, thus preventing genetic differentiation within the population of the Azores.     

Ranging patterns of bottlenose dolphins living in oceanic waters: implications for population structure

Very little is known about the ecology of common bottlenose dolphins (Tursiops truncatus) living in oceanic waters. This study investigated the ranging and residence patterns of bottlenose dolphins occurring in the Azores (Portugal), the most isolated archipelago in the North Atlantic. Data were collected during standardized boat-based surveys conducted over a 6-year period in an area of approximately 5,400 km² (main study area). To investigate the extent of movements of individual animals, non-systematic surveys were also conducted outside this area. Only 44 individuals out of 966 identified were frequently sighted within and between years. The remaining individuals were either temporary migrants from within or outside the archipelago, or transients. Resident dolphins showed strong geographic fidelity to the area. Long-distance movements (of almost 300 km), consistent with foraging or exploratory trips, were observed among non-resident dolphins. Home range size was estimated for 31 individuals sighted ≥10 times. Range areas of these dolphins varied in size and location, but considerable overlap was observed in the areas used, suggesting the absence of habitat partitioning between resident and non-resident dolphins. Estimates of home range size of bottlenose dolphins in the Azores were found to be considerably larger than those previously reported for this species. It is hypothesized that dolphins living in the Azores carry out extensive movements and have large home ranges in response to the lower density and patchy distribution of prey compared to other areas. The extensive ranging behaviour and the lack of territoriality provide an opportunity for interbreeding between dolphins associated with different islands, thus preventing genetic differentiation within the population of the Azores.     

Simultaneous GPS tracking reveals male associations in a solitary carnivore

In hypercarnivorous species, females have large spatial requirements to meet their nutritional needs, and food competition among females is intense. As a result, females are typically solitary and territorial, and solitary males compete for access to dispersed females. Yet, largely anecdotal reports indicate that facultative male sociality may be more common in solitary carnivores than previously thought. We studied spatial interactions among fossas (Cryptoprocta ferox), Madagascar's largest carnivore, using simultaneous GPS tracking of 13 adult individuals to determine patterns of sex-specific spatial distribution and sociality. Male home ranges were larger than those of females, male home ranges overlapped more with those of other males than those of females with other females. Whereas some males were solitary, a subset of adult males was found to have very high home range overlap, high rates of co-location within <50 m, low minimum inter-individual distances, and significantly positive “dynamic interaction”. These associated dyads sometimes, but not always, were close relatives. The fact that solitary and associated males coexist in this population raises interesting questions concerning constraints and flexibility of social tolerance. This study yielded preliminary indications that female distribution appears to be primarily structured by resource competition, whereas male sociality seems to depend on demographic chance events, yet unknown proximate determinants of social tolerance, and it is associated with somatic and reproductive advantages. Male associations among carnivores are therefore more widespread and appear to be based on a wider range of factors than previously thought.     

Sequential polyandry by brood desertion increases female fitness in a bird with obligatory bi-parental care

Parental care may be costly to parents because it decreases resources allocated to self-maintenance and may thus reduce survival and future reproductive success. An inter-sexual conflict may exist in animals with obligatory bi-parental care, such as birds of prey, in which females incubate and brood, whereas males provision food for their families. We analysed 29 years of data (1981–2009) from a study population of Tengmalm’s owls Aegolius funereus in western Finland to examine the occurrence and timing of brood desertion and sequential polyandry, and recorded a total of 1,123 monogamous and 12 polyandrous females. These data were supplemented with the 29-year nationwide Finnish ringing data, which included 11,590 monogamous and 20 polyandrous females. The 12 polyandrous females started egg-laying in their two nests at intervals of 54–68 days (mean 60 days), thus deserting their first broods when the age of oldest young averaged 21 days. Thirty-two polyandrous females re-mated and raised a second brood at a median distance of 4.5 km (range 1–196 km). These females produced 79% more eggs, 93% more hatchlings and 73% more fledglings than did females that laid simultaneously but remained monogamous. Our results show that not only males, but also females of altricial species with bi-parental care can increase their fitness by deserting their first brood when it will be cared for by the males. Earlier studies have shown that male owls can increase their lifetime reproductive success by simultaneous polygyny, and we suggest that an inter-sexual “tug-of-war” over bi-parental care exists in Tengmalm’s owls.     

Spatial ecology and residency patterns of adult great barracuda (<Emphasis Type="Italic">Sphyraena barracuda</Emphasis>) in coastal waters of The Bahamas

Great barracuda (Sphyraena barracuda) were implanted with acoustic telemetry transmitters (n = 42) and monitored within a stationary acoustic receiver array (n = 53 receivers) in The Bahamas to examine residency, seasonal movements, and habitat use. Barracuda were monitored for up to 980 days and remained within the array area ~33% (median value) of the time. Most tagged barracuda were transient and would often disappear from the array for months at a time, particularly in the summer where they were usually last detected on receivers located in deeper shelf habitats, and then return at other times in the year. Habitat use across the footprint of the array differed, with most detections occurring in coastal areas and comparatively fewer in deeper mosaic or shelf habitats. Linear home range estimates revealed that some barracuda moved >12 km within a single day and are capable of migrating >100 km to other islands in the Bahamian Archipelago. Our results provide some of the first telemetry data for this apex marine predatory fish and the first reliable information on the residency and localized seasonal movements of adult great barracuda in the coastal waters of the Western Atlantic.     

Female distribution affects mate searching and sexual selection in male northern water snakes (Nerodia sipedon)

Mating systems and sexual selection are assumed to be affected by the distribution of critical resources. We use observations of 312 mating aggregations to compare mate-searching success of male northern water snakes (Nerodia sipedon) in two marshes in which differences in mating substrate availability resulted in more than fourfold differences in female dispersion. Reproductive males had significantly larger home ranges where females were dispersed than where females were clumped. The number of females encountered by males increased significantly with male home range size where females were dispersed, and decreased significantly where females were clumped. Where females were clumped, males were more likely to encounter other males when they located females. We found no evidence in either population that mate searching was energetically expensive or that males with relatively more energy had larger home ranges. However, males with greater fat reserves at the start of the season participated in more mating aggregations when females were dispersed, suggesting that fat reserves could affect a male’s willingness to attempt mating or to persist in aggregations. When females were dispersed there was weak stabilizing selection acting to maintain male body size (β=–0.14), but strong directional selection favoring larger (β=0.50) and fatter (β=0.37) males. Over 7 years, the intensity of selection favoring larger males varied substantially (β=0.14–1.15), but that variation was not related to variation in the operational sex ratio. We found no evidence of directional selection on either body size (β=0.05) or fat reserves (β=0.10) of males when females were spatially clumped. Overall, the distribution of females had a pronounced effect on male behavior, on the factors that affected male success in locating females, and probably on the extent of sperm competition once females had been located. Received: 23 November 1998 / Received in revised form: 9 August 1999 / Accepted: 18 August 1999     

Relatedness and avoidance of inbreeding in the lizard, Tiliqua rugosa

Two potential mechanisms for reducing the level of inbreeding, sex-biased dispersal and kin avoidance, were examined in the Australian sleepy lizard, Tiliqua rugosa. The home range centres, and the genotypes at four polymorphic microsatellite DNA loci were determined for adult lizards in a 70-ha study area near Mount Mary, South Australia. From estimates of genetic relatedness, females were as closely related to other females as they were to males, both within the whole study area, and within home ranges. Similarly, males were as closely related to other males as they were to females. This suggests that dispersal in the population is not sex-biased. Sleepy lizards form monogamous pairs during the spring. Partners were less closely related to each other than to other potential partners in the home range area. This suggests active choice of unrelated partners. The mechanism for recognising related from unrelated individuals is unknown, but the behaviour could reduce inbreeding. Received: 7 November 1998 / Accepted: 30 May 1999     

Socially induced delayed primiparity in brown bears Ursus arctos

Reproductive suppression through behavioral or physiological means is common in group-living and cooperative breeding mammals, but to our knowledge it has not been shown in wild large carnivores other than those with a clear form of social organization. Brown bear (Ursus arctos) females form matrilinear assemblages with related females using a common and largely exclusive area. Behavioral reproductive suppression might develop due to a hierarchical system among females within a matrilinear assemblage or due to inbreeding avoidance, because male brown bears can overlap with their daughters. We tested whether natal dispersal influenced the age of primiparity. We predicted that emigrant females, geographically removed from maternal or paternal influence, would reproduce earlier than philopatric females. The average age of primiparity was 4.3 years in females that dispersed outside their mother’s home range (n=8) and 5.2 years in philopatric females (n=10). Only the overlap with the mother’s home range, and not body size, body mass, growth, local population density, or overlap with the father’s home range, had a significant influence on the age of primiparity. The ultimate role of reproductive suppression for brown bears is likely to avoid inbreeding or to minimize resource competition. Due to the low risk of inbreeding and frequent exposure of young females to unrelated males, we conclude that resource competition within female hierarchies causes reproductive suppression in young females.     

Determinants of male reproductive success in American black bears

We determined annual male reproductive success in black bears (Ursus americanus) using DNA and field data from two populations in New Mexico. We identified the likely father for 78 of 120 genotyped cubs born during 1994–2000 and calculated reproductive success for 102 males known or presumed present within designated mating regions. Age was a strong determinant of reproductive success. The observed peak in reproductive success occurred at roughly the same intermediate age (10 years) as milder peaks in body size characteristics (11–12 years) and frequency of bear-inflicted wounds (13 years), suggesting body size and fighting might be important for mating. Success was negatively associated with the distance between home range centers of males and mothers. Success of young males (<7 years old) was also negatively associated with mature male (≥7 years old) density, and increasing density shifted the peak age of reproduction higher. The dispersed distribution of females likely limited the capacity of large mature males to dominate reproduction; therefore, success was determined by a complex set of variables.     

Reproductive biology of a wood-boring isopod, Sphaeroma terebrans, with extended parental care

The isopod Sphaeroma terebrans Bate, 1866 burrows in aerial roots of the red mangrove Rhizophora mangle L. The burrows serve as shelter and as a reproductive habitat, and females are known to host their offspring in their burrows. I examined the reproductive biology of S. terebrans in the Indian River Lagoon, a shallow lagoon stretching for ∼200 km along the Atlantic coast of Florida, USA. Reproductive isopods were found throughout the year, but reproductive activity was highest in the fall and during late spring/early summer. During the latter periods, large numbers of subadults established their own burrows in aerial roots. The average numbers of S. terebrans per root were high during the fall, but decreased during the winter and reached lowest levels at the end of the summer. Females reached maturity at a larger size than males, but also grew to larger sizes than the males. The average size of females varied between 8 and 10 mm, the average size of males between 6.5 and 8.5 mm. The number of embryos female⁻¹ was strongly correlated with female body length. No indication for embryo mortality during development was found. Parental females (i.e. with juveniles in their burrows) hosted on average between 5 and 20 juveniles in their burrows (range 1 to 59). Most juveniles found in female burrows were in the manca stage and 2 to 3 mm in body length. Juveniles did not increase in size while in the maternal burrow, and juveniles of similar sizes could also be found in their own burrows. Males did not participate in extended parental care, since most of them left the females after copulation. Many females that were born in the summer produced one brood in the fall and a second during winter/early spring. Females that were born in the fall produced one brood during spring/early summer, but then probably died. Extended parental care in S. terebrans is short compared to other peracarid crustaceans. It is concluded that this reproductive strategy in S. terebrans serves primarily to shelter small juveniles immediately after they emerge from the female body, when their exoskeleton is still hardening and their physiological capabilities are still developing. Thus, in S. terebrans, extended parental care probably aids to protect small juveniles from adverse physical conditions in their subtropical intertidal habitat. Received: 9 December 1998 / Accepted: 24 June 1999     

Home range and habitat use of juvenile Atlantic green turtles (Chelonia mydas L.) on shallow reef habitats in Palm Beach, Florida, USA

Many animals, including sea turtles, alter their movements and home range in relation to the particular type and quality of the habitat occupied. When sufficient resources are available, individuals may develop affinities to specific areas for activities, such as foraging and (or) resting. In the case of green sea turtles (Chelonia mydas L.), after a number of years in the open ocean, juveniles recruit to shallow-water developmental habitats where they occupy distinct home ranges as they feed and grow to maturity. Our goal was to study the habitat use and home range movements of juvenile green turtles along a shallow, worm-rock reef tract in Palm Beach, Florida. Six turtles, measuring from 27.9 to 48.1 cm in straight carapace length and from 7.2 to 12.6 kg in mass, were tracked via ultrasonic telemetry from August to November 2003. Upon capture, each turtle’s esophagus was flushed via lavage to determine recently ingested foods. In addition, four turtles were recaptured and fitted with a time-depth recorder to study dive patterns. Home range areas measured with 100% minimum convex polygon and 95% fixed kernel estimators varied from 0.69 to 5.05 km² (mean=2.38±1.78 km²) and 0.73 to 4.89 km² (mean=2.09±1.80 km²), respectively. Home ranges and core areas of turtles were largely restricted to the reef tract itself, and showed considerable overlap between food and shelter sites. The mean number of dives during daylight hours (0600–1800 hours) was 84±5.0 dives, while the mean during night hours (1800–0600 hours) was 39±3.0 dives. Dives during the day were shallower (mean=3.20±1.26 m) than dives at night (mean=5.59±0.09 m). All six turtles were found to have a mixed diet of similar macroalgae and sponge fragments. Our results reveal that juvenile green turtles occupy stable home ranges along the nearshore worm-rock reefs of Southeast Florida, during the summer and fall. Determining which habitats are used by green turtles will assist conservation managers in their global effort to protect this endangered species.     

Social organization and nest co-occupancy in Peromyscus californicus,a monogamous rodent

Summary Home ranges, social organization, and nest co-occupancy of Peromyscus californicus were studied using radiotelemetry at the Hastings Natural History Reservation, California. Mated pairs were ascertained by the transfer of fluorescent pigments from lactating females to putative fathers. Mated pairs had largely overlapping home ranges that were not statistically distinguishable, whereas adjacent adults had mostly exclusive, statistically distinguishable home ranges. There was no difference in the mean home range of males and females, but mated females tended to have smaller ranges than their mate. Home range size was extremely variable (range: 150–3788 m²) and averaged 1161 m² across all individuals. Male home range size was inversely correlated with population density, suggestive of a social influence on home range. Putative fathers spent comparable amounts of time to females in the nest — presumably caring for the young — which supports previous laboratory reports of paternal care in this species. All data collected in this study are consistent with previous suggestions that P. californicus live in semi-permanent family groups and are monogamous. Offprint requests to: D.O. Ribble     

Genetic relatedness and space use in a behaviorally flexible species of marmot, the woodchuck (Marmota monax)

Solitary species show several patterns of space use and relatedness. Individuals may associate randomly or may live near female or male kin, often as a result of natal philopatry or dispersal patterns. Although usually described as solitary or asocial, woodchucks (Marmota monax) are behaviorally flexible marmots that exhibit greater sociality in some populations than others. I examined relationships between kinship, geographic distance, and home range overlap, as well as dispersal and philopatry, to determine the extent to which kin associated spatially. I used a combination of microsatellite DNA analysis, long-term behavioral observations, and radiotelemetry to test predictions that females, but not males, would associate with kin. Indeed, woodchucks lived closer and shared a greater proportion of their home range with more closely related animals. Overlap of females' and males' home ranges was positively correlated with kinship, and male–female dyads shared more area with closer kin. Most juveniles delayed dispersal beyond their first summer. Females often remained philopatric and settled near their natal range. Although males often dispersed as yearlings, some males also established territories within or immediately adjacent to their natal home ranges. A combination of factors can explain these spatial patterns, including high population density associated with the study site's location within a suburban environment, high dispersal costs, and abundant food. Thus, despite their asocial and solitary reputation, woodchucks displayed spatial patterns seen in other, more social species of ground-dwelling sciurids.     

Home range overlap of mothers and their offspring in the sleepy lizard, Tiliqua rugosa

This paper reports a field investigation of interactions between juveniles and their mothers in the Australian sleepy lizard, Tiliqua rugosa. In their first spring season, juvenile lizards maintain home ranges largely within the home range of their mother. Juvenile home ranges are significantly smaller than those of adult males and females, and juveniles move significantly less often and significantly shorter distances than adults. While siblings were never found together in the spring, they showed a significant tendency to be closer to each other than if they were randomly located in their home ranges. Juveniles and mothers were never found together, nor was there any evidence for any positive (or negative) spatial association. Nevertheless, the extended tolerance of home range overlap represents a greater degree of mother-offspring association than has been previously reported for other lizards. Despite this, the level of parental care can only be described as minimal. Received: 20 June 1997 / Accepted after revision: 29 December 1997