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.     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests     

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     

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 spacing behavior in microtines: an experimental manipulation of female spatial distribution and density

Theory on microtine mating systems predicts that male spacing behavior will be related to female spatial and temporal distribution. However, data from a natural population of field voles, Microtus agrestis, indicated a potential influence of female density on the spacing behavior of males. Therefore, I experimentally investigated the relative importance of female density and female spatial distribution for the spacing behavior of males in M. agrestis. Males were radio-tracked in enclosed natural habitats in which females at different densities were placed in two different spatial arrangements: clumped versus even distribution. Female density was the main factor determining male spacing behavior. At the high female density males had smaller home ranges and moved shorter distances between radio-tracking recordings. Also, home ranges were more exclusive at high female density. However, since there was a significant positive correlation between home range size and range overlap, range exclusiveness seemed to be influenced by female density indirectly through the effect of range size. Female spatial distribution, on the other hand, had no influence on male home range exclusiveness.     

Histology of the gonads and observations on the social behavior of the Caribbean angelfish Holacanthus tricolor

Male and female specimens of Holacanthus tricolor were collected in May 1981 and September 1982 off the Island of St. Croix. U.S. Virgin Islands, and their gonads were examined histologically. All testes examined showed vasa deferentia which were characteristic of secondary males, suggesting protogynous sex change. A transitional individual supported this interpretation. Ovarian histology and size-specific egg counts revealed seasonal differences in spawning frequency and clutch size. Females appeared to spawn every day in May. The timing of final maturation and hydration of the oocytes indicated that spawning occurred in the late afternoon or early evening. Not all females spawned every day in September, and clutch sizes were smaller than those of May. Behavioral observations of H. tricolor in the field revealed that males defended harems of two to four smaller females. Within each harem, females of different sizes had overlapping home ranges, while the home ranges of females of similar size did not overlap. Besides size dimorphism between the sexes, H. tricolor exhibited a previously unreported sexual dichromatism.     

Mating by pregnant water voles (Arvicola terrestris): a strategy to counter infanticide by males?

Summary Pregnant water voles (Arvicola terrestris) shift nest location, with the result that some females may move into the home range of a new male. These females were found to have a vaginal plug while still pregnant, indicating a recent mating with the new male. No vaginal plugs were found in pregnant females that stayed within their original male's home range. No indications of pregnancy termination were observed in females that encountered a new male. This suggests that pregnant female water voles mate with the new male to prevent infanticide by confusing paternity.     

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     

Kin-related spatial structure in brown bears Ursus arctos

Kin-related social structure may influence reproductive success and survival and, hence, the dynamics of populations. It has been documented in many gregarious animal populations, but few solitary species. Using molecular methods and field data we tested: (1) whether kin-related spatial structure exists in the brown bear (Ursus arctos), which is a solitary carnivore, (2) whether home ranges of adult female kin overlap more than those of nonkin, and (3) whether multigenerational matrilinear assemblages, i.e., aggregated related females, are formed. Pairwise genetic relatedness between adult (5 years and older) female dyads declined significantly with geographic distance, whereas this was not the case for male–male dyads or opposite sex dyads. The amount of overlap of multiannual home ranges was positively associated with relatedness among adult females. This structure within matrilines is probably due to kin recognition. Plotting of multiannual home-range centers of adult females revealed formation of two types of matrilines, matrilinear assemblages exclusively using an area and dispersed matrilines spread over larger geographic areas. The variation in matrilinear structure might be due to differences in competitive abilities among females and habitat limitations. The influence of kin-related spatial structure on inclusive fitness needs to be clarified in solitary mammals.     

Home range and social behavior of three species of European Pitymys (Mammalia,Rodentia)

Summary Habits, home range, and social behavior of three species of voles of the genus Pitymys, P. multiplex, P. subterraneus, and P. savii, were studied in Tessin (Switzerland) by radioactive tagging. P. multiplex and P. savii are fossorial and often use the burrow systems of moles (Talpa); P. subterraneus moves around on the surface under dense vegetation. Males and females of P. multiplex were territorial and their home ranges averaged 342 and 229 m² respectively. The mating system appeared to be monogamous. P. savii male and female home ranges averaged 445 and 298 m² respectively. Groups of up to one to three adult females, one or more adult males, and several young, occupied exclusive communal territories. The mating system appeared to be flexible, including monogamy, polygyny, and polyandry. P. subterraneus females occupied exclusive territories (mean 256 m²) overlapped by the home ranges of one or more males (1025 m²). The mating system appeared to be promiscuous. Individuals of the three species occupied one to seven nests. In P. savii all family members utilized three or more communal nests at the same time or alternatively. Co-nesting between males and females of P. multiplex or P. subterraneus were observed less frequently.     

There is a limbo under the moon: what social interactions tell us about the floaters’ underworld

The ultimate and proximate causes of natal dispersal have been extensively investigated, but the behaviour of dispersers in relation to social interactions has been largely neglected. Here, we investigated the social organisation of floating individuals during their dispersal by analysing the behaviour of 40 radio-tagged eagle owls Bubo bubo during the wandering and stop phases of dispersal. Unexpectedly, eagle owl floaters mixed with conspecifics independently of their sex, age, phase of dispersal, birthplace, health status and habitat features, showing an ‘underworld’ of interactions characterised by the absence of obvious social organisation or short-term strategies. Non-breeding owls were not transient floaters that occurred at numerous sites for short periods of time but rather had fairly stable home ranges: they attempted to settle as soon as possible within well-defined home ranges. The spatial distribution pattern of floaters and high rates of home range overlap support the prediction that floating individuals are not spatially segregated, challenging the expectation that dominance by size, age and/or health status may determine the exclusive use of some portions of the dispersal area. Finally, (1) the short distances among conspecifics and the extensive home range overlaps allowed us to discard the possibility that neighbouring floaters represent a real cost during dispersal and (2) floater interactions showed a lack of clear mechanisms for avoidance of kin competition among offspring or inbreeding.     

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.     

Territorial behavior of the black and yellow rockfish and gopher rockfish (Scorpaenidae,Sebastes)

Tagged black-and-yellow rockfish Sebastes chrysomelas and gopher rockfish S. carnatus were observed in situ for 1 yr at 3 study sites off California, USA in order to determine the nature of their home ranges and territories. Three basic patterns of space use were identified: (1) a shelter hole located within a larger area of activity (in which the fish presumably fed, since its size increased with size of fish and decreased with prey density); (2) discontinuous home ranges, in which the shelter holes and feeding areas were separte; and (3) dispersed home ranges with little or no exclusive area. Fish in the first category were regarded as territorial, since their home ranges were defended and relatively exclusive. Because individuals did not patrol borders, however, territories often overlapped at their peripheries. In addition, some territory holders appeared to tolerate the sheltering of other fish within their territories, as long as these intruders left the territory during periods of activity. Some fish with discontinuous home ranges (commuters) may have been territorial, but others may have been forced to occupy suboptimal feeding areas. Fish in the third category (floaters) were regarded as nonterritorial; their activities were apparently limited to areas where they could escape the aggression of territory holders. The varying patterns appeared to develop through defense and accommodation during the turnover of individuals, with each fish securing shelter and feeding areas as these became available.     

Sex differences in the foraging behavior of squirrel monkeys in a seasonal habitat

Summary The effects of sex and seasonal changes in food abundance on foraging behavior was studied in squirrel monkeys (Saimiri oerstedi) in Costa Rica over an eleven-month period. Females searched for and ate food at significantly greater frequencies than did males throughout the study. The frequency of the specific foraging techniques used occasionally differed significantly within seasons, but not across the study period. Few differences were found in the foraging behaviors of nonreproductive sexually mature females compared to females that were pregnant or lactating. The major exception was that during the month following parturition reproductive females foraged for flowers and fruits more frequently than did non-reproductive females. The reduction of time spent by males in foraging activities gives them more time for other activities, especially anti-predator vigilance. Foraging techniques and the proportions of different food types in the diet changed seasonally. Foraging for arthropods was most frequent in the season when arthropod abundance was lowest, resulting in the amount of time spent eating arthropods to vary less across the seasons. Fruits and flowers were not eaten in a direct relationship to availability, but were used more than expected relative to availability when arthropod abundance was reduced. Individuals were more dispersed when foraging compared to other activities. Overall, there was little evidence of any direct foraging benefits for a squirrel monkey from being social.     

Gunnison’s prairie dog (<Emphasis Type="Italic">Cynomys gunnisoni</Emphasis>): testing the resource dispersion hypothesis

Few studies have experimentally tested the resource dispersion hypothesis (RDH). In this study, I tested whether space use and social organization of Gunnison’s prairie dog responded to changes in the dispersion and abundance of resources. Food manipulations were carried out during the reproductive and nonreproductive seasons across 2 years. Gunnison’s prairie dog adults responded to the experiments by decreasing territory size as food became patchier in space and time. Both males and females modified their home ranges, with no detectable difference between sexes, either prior to or during the experiments. As food became patchier in space and time, the spatial overlap of adults increased, whereas it decreased as food became more evenly dispersed. The average size of a group, defined as those individuals occupying the same territory, did not change significantly as a result of the experiments. Where changes in the composition and size of groups did occur, there was no indication that such changes were sex specific. Results from this study support critical components of the RDH and strongly suggest that patterns of space use and social structure in Gunnison’s prairie dogs are the result of individual responses to resource abundance and distribution.     

Costs and benefits of home range shifts among vervet monkeys (Cercopithecus aethiops) in Amboseli National Park,Kenya

Summary A decline in the density of trees and the deterioration of the habitat of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya, have been accompanied by movement of vervet groups into unfamiliar areas. These home range shifts were documented over a period of 26 months. The largest group moved into the home ranges of neighboring groups and acquired more trees, a resource that had become more limited. The groups whose home ranges were encroached upon did not share their home ranges with the intruders, but instead moved into areas that were unoccupied and had a lower density of trees. The largest group was thus more successful than smaller groups in competition for limited resources. For all but one group, movement into new areas was directly associated with the disappearances of female and immature monkeys. Most of the disappearances were attributable to predation. These results suggest that the cost of predation in unfamiliar areas may contribute to the evolution of philopatry in vervets. However, under unstable conditions, resource competition forces these animals to disperse (in groups) despite the high risk of predation. Offprint requests to: L.A. Isbell     

Size and spatial arrangement of home range of checkered snapper <Emphasis Type="Italic">Lutjanus decussatus</Emphasis> (Lutjanidae) in an Okinawan coral reef determined using a portable GPS receiver

Although snappers (Lutjanidae) are commercially important fishery resources in tropical and subtropical waters, their home range size and its spatial arrangement have not been sufficiently clarified. In the present study, the size and spatial arrangement of the home range of the reef-associated checkered snapper Lutjanus decussatus (Lutjanidae) were investigated by use of a portable GPS receiver. In a 120 m × 120 m quadrat established on a section of the fringing reef of Ishigaki Island, 29 individuals of the species were identified by their color pattern and individuals were divided into five arbitrary size classes (class 1 <10 cm TL ≤ class 2 <15 cm TL ≤ class 3 <20 cm TL ≤ class 4 <25 cm TL ≤ class 5). Fish tracking by snorkeling was conducted with the portable GPS receiver. Site fidelity of this species was high. Home range size ranged from 93.0 to 3638.4 m², and there was a significant positive correlation between the home range size and fish total length. Home ranges of the same-sized individuals abutted each other (8.8% area overlap), whereas those of different-sized individuals overlapped (44.0% area overlap). Agonistic behavior (attack and agonistic display) was more frequently found among same-sized individuals (times of agonistic behavior/times of all encounters × 100 = 71.3%), whereas such agonistic behavior was rarely found among different-sized individuals (times of agonistic behavior/times of all encounters × 100 = 6.9%). These results suggest that home ranges of Lutjanus decussatus can be regarded as territories against same-sized individuals, but not different-sized individuals. The usefulness of the fish tracking by snorkeling using a portable GPS receiver for home range size estimation and the function of the overlapping territory of the species are discussed.