The functional significance of infanticide in a population of California ground squirrels (Spermophilus beecheyi)

Naturally-occurring infanticide was observed in a population of California ground squirrels (Spermophilus beecheyi). In four seasons, 40 infanticides were observed. All victims were post-emergent pups. Of 37 killings in which the killer was sexed 36 were by females. All infanticidal females were mothers at the time they killed, but in no case was a mother seen to kill or harm her own young. The victim was cannibalized in 22 cases and taken immediately into the killer’s burrow in 16 others. In no case did killers gain access to the victimized mother’s burrow or territory and female pups were not killed preferentially over males. In light of evolutionary explanations, infanticide in this population may best fit the resource exploitation hypothesis, in which killers commit infanticide to gain a nutritional benefit. Resource competition is a possible auxiliary explanation, since any time a female kills unrelated young she is eliminating possible competitors to her own offspring. This behavior could confer a fitness advantage on killers or it could be an adaptively neutral, alternative feeding strategy. More data are necessary to distinguish between these hypotheses. Received: 26 January 1995/Accepted after revision: 9 September 1995     

Sibling recognition in Richardson's ground squirrels (Spermophilus richardsonii)

Summary Young from 10 litters of Richardson's ground squirrels were crossfostered within 24 h of birth. Litters, composed then of uterine-sibs and fostered non-sibs, were caged together until 37 days of age (1 week after weaning), at which time the juveniles were separated and caged individually. At a mean age of 110 d squirrels were tested for recognition during 10-min trials in an arena using the following pair-types: sibs reared together (SRT), sibs reared apart (SRA), non-sibs reared together (NSRT), and non-sibs reared apart (NSRA). Juvenile S. richardsonii displayed an ability, determined prenatally or soon after birth, to recognize their biological siblings. Familiarity, based upon association before and at the time of weaning, also affected discrimination between conspecifics, although this effect was less apparent.     

Natal dispersal in Belding's ground squirrels (Spermophilus beldingi)

Summary Dispersal from the natal site was documented in two populations of Belding's ground squirrels (Spermophilus beldingi) living at different altitudes in the Sierra Nevada of North America. Distance dispersed and age at dispersal were monitored by a combination of observation, trapping, radio telemetry, and examination of road kills. Dispersal was sexually dimorphic in both populations (Tables 1 and 2). All surviving males emigrated before they were 55 weeks of age, with most dispersing midway through the juvenile summer (Fig. 1). By contrast, most females remained within the boundaries of their mothers' home range (Fig. 2 and 3). Those very few females that did emigrate moved distances from their natal burrows similar to those travelled by dispersing males (200–450 m; Table 2), but females tended to disperse at a slightly older age. Significant differences between the study populations were found in distances moved by juveniles of both sexes (Fig. 2). Body weights of juvenile male dispersers were significantly greater than were those of juvenile males of equivilent ages that had not yet dispersed (Fig. 4). The results were considered in light of Shields' (1982) discussion of dispersal and inbreeding. I concluded that if dispersal is related to reproductive success, then dispersal distances selected for use in tests of evolutionary hypotheses should be measured just prior to the subject animals' first reproduction. Finally, although not tested in the present study, the evolutionary hypothesis most consistent with my data suggests that dispersal in S. beldingi may function to minimize nuclear family incest.     

Temporal overlap of female kin in Columbian ground squirrels (Spermophilus columbianus)

Summary Columbian ground squirrels (Spermophilus columbianus) were live-trapped from 5 to 7 years at three sites in southwestern Alberta, Canada. The two most common adult matrilineal relatives for 2-year-old females were the mother and a 1 year older non-littermate sister. Co-occurrence of female kin depended firstly on size and sexual composition of litters and secondly on age-specific survival and recruitment rates. Adult matrilineal kin frequently coexisted among breeding females of this species, leading us to predict a social system strorgly influenced by nepotism. However, the life history traits of S. columbianus suggest an adult female kin cluster unlike that found for other species of Spermophilus in which littermate sisters are common. If availability of adult female kin influences kin-differential behaviour, then female S. columbianus should favour mothers, daughters and non-ittermate sisters.     

Development of food preferences: social learning by Belding's ground squirrels Spermophilus beldingi

Summary The mechanisms by which food selection behavior develops may constrain the evolution of optimal foraging, yet these mechanisms have received relatively little attention in recent optimal foraging studies. We used cafeteria-style feeding trials to examine the role of social learning in the development of food preferences by a generalist mammalian herbivore, Spermophilus beldingi. Naive young spent relatively little time feeding and showed no preferences among five plant species offered in their feeding trials. This random pattern persisted for one set of young during two subsequent trials in the absence of their mother. A second group of young squirrels was tested initially alone, once with their mothers, and then once more alone. These squirrels fed more than those in the control group during their final trials, and showed significant preferences among plant species. These paralleled the preferences of their mothers. These results suggest that social learning may be important in the development of feeding behavior in ground squirrels, and provide a possible mechanism for cultural differences in food preferences among populations.     

Temporary alteration of local social structure in a threatened population of Cuban iguanas (Cyclura nubila)

In small, insular populations, behavioral patterns that lead to increased variance in individual reproductive success can accelerate loss of genetic variation. Over a 1-year period, we documented behavior and hormone levels in a breeding group of adult Cuban iguanas (Cyclura nubila) at Guantánamo Bay. Male dominance was associated with body and head size, display behavior, testosterone levels, home-range size, and proximity to females. Based on their success in agonistic encounters, we ranked males in a linear dominance hierarchy. During the subsequent breeding season, we conducted a removal experiment in which the five highest-ranking males were temporarily relocated from the study site. Although we were unable to assess reproductive success directly, previously lower-ranking males assumed control of vacated territories, won more fights, and increased their proximity to females in the absence of the dominant males. When it results in greater mating opportunities for otherwise socially suppressed individuals, temporary alteration of local social structure may help limit erosion of genetic variation in small, insular populations. Electronic Publication     

Sibling recognition in thirteen-lined ground squirrels: effects of genetic relatedness,rearing association,and olfaction

Summary I investigated sibling-sibling recognition in captive thirteen-lined ground squirrels (Spermophilus tridecemlineatus) by cross-fostering lab-born pups shortly after birth. When young reached about 45 days of age, I observed dyadic interactions in a test arena of pairs from four relatedness X rearing groups, and recorded the frequency of exploratory encounters between individuals. Sibs-reared together and nonsibs-reared together exhibited significantly fewer exploratory encounters than either sibs-reared apart or nonsibsreared apart. Young reared together were equally exploratory, regadless of relatedness; similarly, young reared apart, whether they were sibs or nonsibs, showed similar levels of exploration. Thus, the differential treatment of siblings in the lab appears to be based on rearing association and not genetic relatedness per se. I interpret this recognition based on association (rearing familiarity) in the context of the species' social organization and compare my results on S. tridecemlineatus with similar studies on S. beldingi, S. parryii, and S. richardsonii. I also used an olfactory impairment technique (zinc sulfate) and found that differential treatment in thirteen-lined ground squirrels was influenced by olfactory cues.     

Ecological and hormonal correlates of antipredator behavior in adult Belding’s ground squirrels (Spermophilus beldingi)

Predator–prey relationships provide an excellent opportunity to study coevolved adaptations. Decades of theoretical and empirical research have illuminated the various behavioral adaptations exhibited by prey animals to avoid detection and capture, and recent work has begun to characterize physiological adaptations, such as immune reactions, metabolic changes, and hormonal responses to predators or their cues. A 2-year study quantified the activity budgets and antipredator responses of adult Belding’s ground squirrels (Spermophilus beldingi) living in three different California habitats and likely experiencing different predation pressures. At one of these sites, which is visually closed and predators and escape burrows are difficult to see, animals responding to alarm calls remain alert longer and show more exaggerated responses than adults living in two populations that likely experience less intense predation pressure. They also spend more time alert and less time foraging than adults at the other two sites. A 4-year study using noninvasive fecal sampling of cortisol metabolites revealed that S. beldingi living in the closed site also have lower corticoid levels than adults at the other two sites. The lower corticoids likely reflect that predation risk at this closed site is predictable, and might allow animals to mount large acute cortisol responses, facilitating escape from predators and enhanced vigilance while also promoting glucose storage for the approaching hibernation. Collectively, these data demonstrate that local environments and perceived predation risk influence not only foraging, vigilance, and antipredator behaviors, but adrenal functioning as well, which may be especially important for obligate hibernators that face competing demands on glucose storage and mobilization.     

Variation in litter size: a test of hypotheses in Richardson's ground squirrels

We studied litter size variation in a population of Richardson's ground squirrels (Spermophilus richardsonii) in Alberta, Canada, from 1987 to 2004. Litter size at first emergence of juveniles from the natal burrow ranged from 1 to 14; the most common litter sizes, collectively accounting for 41.0% of 999 litters, were 6 and 7. The number of offspring surviving to adulthood (attained on emergence from hibernation as yearlings) increased with increasing litter size, a result that was not predicted by Lack's "optimal litter size" hypothesis, Mountford's "cliff-edge" effect, or the "bad-years" effect. Contrary to the negative effects predicted by the "cost of reproduction" hypothesis, litter size had no significant influence on survival of mothers to the subsequent year or on the size of the subsequent litter. Rather, our results best fit the predictions of the "individual optimization" hypothesis, which suggests that litter size is determined by the body condition and environmental circumstances of each mother. Supporting this hypothesis, survival of individual offspring was not significantly associated with litter size. Additionally, year-to-year changes in maternal body mass at mating were positively associated with concurrent changes in litter size (r = 0.56), suggesting that litter size depends on the body condition of the mother. Because the mean number of recruits to adulthood increased as litter size increased (r2 = 0.96) and litter size increased with maternal condition, offspring productivity was greater for mothers in better body condition.     

Inheritance of nest sites in female Columbian ground squirrels

Summary Home ranges, areas of intensive use, and locations of nest sites were determined for female Columbian ground squirrels and their daughters over seven years from a low-density population in southwestern Alberta. Adult females usually retained similar home ranges from year to year. However, they moved their nest sites more often than expected in years when a yearling daughter was present. Core areas and centres of activity also changed when the nest site was moved. Most daughters settled on their natal areas. The relinquishing of nest sites by females to their daughters is seen as a form of parental investment.     

Mode of larval development and genetic population structure inNodilittorina africana knysnaensis (Prosobranchia: Littorinidae)

We examined electrophoretic variation in the supralittoral gastropod,Nodilittorina africana knysnaensis, along the coast of South Africa in 1989. There was little allozyme variation in nine samples collected over 1 500 km for three highly polymorphic loci,Est, Gpi, andPgm, and Wahlund's effect was absent in the pooled sample. A gene diversity analysis showed that 98.1% of total genic diversity, on average, was contained within populations, and that only 1.9% was due to allozyme frequency differences among populations. Since the mode of larval development has not yet been described for this species, we suggest that the lack of strong allozyme frequency differences over long distances indicates thatN. a. knysnaensis has a planktonic mode of larval development.     

Microgeographic genetic differentiation in a colonial ascidian (Botryllus schlosseri) population

Limited gene flow via the restricted dispersal of larvae and gametes is expected to result in the genetic differentiation of populations of clonal invertebrates on small spatial scales. However, occasional dispersal events over greater distances may generate sufficient gene flow to maintain genetic homogeneity. We applied a spatial autocorrelation approach that does not require a priori definitions of subdivision boundaries to examine genetic differentiation within a continuous population of the colonial ascidian Botryllus schlosseri (Pallas) at two allozyme and five polychromatism loci. Colonies were sampled in July 1992, on a 12 by 18 m grid superimposed on a shallow subtidal (1 to 3 m) population in the Damariscotta River estuary in Maine, USA. Low but significant levels of positive autocorrelation were detected over very small spatial scales (<5 m), with negative autocorrelation occurring on larger scales (>8 m). This pattern indicates significant genetic differentiation over distances of 8 to 21 m, and is consistent with genetic drift and inbreeding creating small scale genetic structure. Received: 18 October 1999 / Accepted: 11 July 2000     

Weight comparisons and litter size manipulation in Columbian ground squirrels (Spermophilus columbianus) show evidence of costs of reproduction

I studied reproductive costs in the female Columbian ground squirrel (Spermophilus columbianus) using individually marked animals. I compared weight changes during the active season and over winter, and mortality for females that did and did not wean young. Females raising young were heavier at emergence in that spring than unsuccessful ones. Females that did not raise young gained more weight during summer, were heavier than successful females at the time of entry into hibernation, and were heavier emerging from hibernation the following spring. Over-winter mortality was higher for females that reared young compared to reproductively unsuccessful females. A food supplementation experiment showed that energy-rich food can accelerate individuals’ weight gain. Interactions between litter size, birth weight, weight at emergence from the natal burrow, survival of young to yearling age, and maternal fitness were also studied. Litter sizes were experimentally manipulated to evaluate how females cope with costs of rearing one additional young. Birth weight of juveniles was positively correlated with survival to emergence from the natal burrow and with survival to yearling age. Partial litter loss was higher in experimentally enlarged litters than in either experimentally decreased or control litters. Total litter loss, survival of adult females or the probability of weaning young the following year were not affected by the litter size manipulation. Females appear to adjust the size of their litter before birth, and to some extent during lactation, to their ability to wean young. Received: 20 January 2000 / Received in revised form: 12 March 2000 / Accepted: 18 March 2000     

The squirrel that cried wolf: reliability detection by juvenile Richardson's ground squirrels (Spermophilus richardsonii)

Where alarm signals function to warn others of the presence of threat, variation is likely to exist in the reliability of alarm signalers. Some signalers, with too low a threshold of excitation, will issue false alarms and should be ignored if potential alarm recipients are to maximize energy gains. We exposed juvenile Richardson's ground squirrels to reliable signalers, whose alarm calls were paired with the presentation of a predator model, and unreliable signalers, whose alarm calls were played when no potential predator was present. Call recipients discriminated among individual alarm callers, and reduced responsiveness to callers that had been unreliable. Thus, like primates, squirrels are capable of forming a concept of reliability by associating an individual's identity with that individual's past performance.     

Climate change and population genetic structure of marine species

Climate change influences populations by reducing or extirpating local populations, by disrupting patterns of migration and by shifting geographical distributions. These events can affect genetic population structure in several ways. Molecular markers have been used in numerous population genetic and phylogeographical studies of marine species and have detected population responses to climate change in the last few decades, such as range expansions, adaptative shifts and declines or increases in abundance. Little is known, however, about the molecular and physiological basis of adaptive responses to climate change in marine Mediterranean species. The Mediterranean Sea ecosystem is a ‘living laboratory’ with native species that are challenged by environmental change and by invasive species and a ‘gene-climate’ approach should be adopted as a way of focusing on the relationship between climate warming and genetic diversity.     

Alternative reproductive tactics in the queen-size-dimorphic ant Leptothorax rugatulus (Emery) and their consequences for genetic population structure

We report the results of a comprehensive investigation of the queen size dimorphism in the North American ant Leptothorax rugatulus. Employing allozymes and microsatellites as genetic markers, we found no evidence that the gene pools of large (macrogynes) and small (microgynes) queens are distinct. Queens in polygynous colonies are related to each other, supporting the hypothesis that colonies with more than one queen commonly arise by the adoption of daughter queens into their natal colonies. The higher fat content of macrogynes, their predominance in monogynous societies and in small founding colonies, and their greater flight activity favor the view that macrogynes predominantly found colonies independently, while microgynes are specialized for dependent colony founding by readoption. When comparing the genetic structure of three different subpopulations, we found that the alternative life histories had no significant effect on population viscosity at the scale investigated.     

Complex social structure of southern flying squirrels is related to spatial proximity but not kinship

Social individuals have organized relationships that affect fitness and so a species' tendency to be social has important implications for its population ecology, gene flow, and its distribution in space and time. We quantitatively examined the social structure of southern flying squirrels (Glaucomys volans) and tested for a role of kinship and prior familiarity in predicting social structure. To quantify social structure, we monitored nest group composition of southern flying squirrels. All squirrels at the study site were marked with passive integrated transponder (PIT) tags and nest cavity entrances were monitored with automated PIT tag recorders for a period of 28 months. Squirrels were genotyped at eight microsatellite loci. Permutation tests of associations suggested that individuals nested with other specific individuals more often than expected by chance. The lagged association rate indicated that relationships were stable and persisted across seasons and years. Multiple summer nest associates came together in winter to form larger nest groups which were likely important for social thermoregulation. A measure of prior familiarity, but not kinship, was related to the proportion of time individuals nested together during winter. We suggest that the evolution of sociality in southern flying squirrels is driven largely by mutually beneficial behaviors related to social thermoregulation although other, as of yet, unidentified mechanisms are needed to explain sociality in the warm season. We hypothesize that minimum group size requirements associated with social thermoregulation could explain the absence of this species in patchy landscapes and aspects of range boundary dynamics near their northern range boundary.