Age-related multi-year associations in female humpback whales (Megaptera novaeangliae)

Analyses of social structures in baleen whales are rare, and so far, they are thought to consist of mostly short and unstable associations. We investigated the association patterns of individual humpback whales from a summer feeding aggregation in the Gulf of St. Lawrence from 1997 to 2005. Photo-identified animals were sexed using genetic methods and were grouped into five categories: juvenile males/females, mature males and lactating/non-lactating females. We calculated half-weight association indices within and between the groups and found that 45% of the observation showed single animals and another 45% small groups (two to three) consisting mainly of mature animals besides lactating females. Using permutation tests, we found evidence for long-term associations between mature males and non-lactating females as well as among non-lactating females. Standardised lagged association rates revealed that these male–female groups disassociated quickly over about 2 weeks, whereas associations increased again towards the beginning of the breeding season. Non-lactating females of similar age engaged in multi-seasonal stable pairs for up to six consecutive feeding seasons; no mature male–female association was observed in consecutive years. The females with the most stable and long-term associations also had the highest reproductive output. While the risk of predation could not explain these long-term bonds, feeding cooperation seemed the most plausible explanation for group forming behaviour during the summer months.     

Rank-related partner choice in the fission–fusion society of the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>)

When social partners vary in their relative value, individuals should theoretically initiate partnerships with conspecifics of the highest value. Here, we tested this prediction in a wild population of spotted hyenas (Crocuta crocuta). Crocuta live in complex, fission–fusion societies structured by dominance hierarchies in which individuals vary greatly in their value as social companions. Because patterns of association among Crocuta reflect social preferences, we calculated association indices (AIs) to examine how social rank influences intrasexual partner choice among unrelated adults of both sexes. The highest-ranking individuals were generally most gregarious in both sexes. Females associated most often with dominant and adjacent-ranking females. Females joined subgroups based on the presence of particular conspecifics such that subordinates joined focal females at higher rates than did dominants. Dominants benefit from associations with subordinates by enjoying priority of access to resources obtained and defended by multiple group members, but the benefits of these associations to subordinates are unknown. To investigate this, we tested three hypotheses suggesting how subordinates might benefit from rank-related partner choice among unrelated females. We found that subordinates who initiated group formation benefited by gaining social and feeding tolerance from dominants. However, rates at which dominants provided coalitionary support to subordinates did not vary with AIs. Overall, our data resemble those documenting patterns of association among cercopithecine primates. We consider our results in light of optimal reproductive skew theory, Seyfarth’s rank attractiveness model, and biological market theory. Our data are more consistent with the predictions of Seyfarth’s model and of biological market theory than with those of skew theory.     

Network structure and parasite transmission in a group living lizard,the gidgee skink, <Emphasis Type="Italic">Egernia stokesii</Emphasis>

Gidgee skinks (Egernia stokesii) form large social aggregations in rocky outcrops across the Flinders Ranges in South Australia. Group members share refuges (rock crevices), which may promote parasite transmission. We measured connectivity of individuals in networks constructed from patterns of common crevice use and observed patterns of parasitism by three blood parasites (Hemolivia, Schellackia and Plasmodium) and an ectoparasitic tick (Amblyomma vikirri). Data came from a 1-year mark-recapture study of four populations. Transmission networks were constructed to represent possible transmission pathways among lizards. Two lizards that used the same refuge within an estimated transmission period were considered connected in the transmission network. An edge was placed between them, directed towards the individual that occupied the crevice last. Social networks, a sub-set of same-day only associations, were small and highly fragmented compared with transmission networks, suggesting that non-synchronous crevice use leads to more transmission opportunities than direct social association. In transmission networks, lizards infested by ticks were connected to more other tick-infested lizards than uninfected lizards. Lizards infected by ticks and carrying multiple blood parasite infections were in more connected positions in the network than lizards without ticks or with one or no blood parasites. Our findings suggest higher levels of network connectivity may increase the risk of becoming infected or that parasites influence lizard behaviour and consequently their position in the network. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address     

Association patterns and kinship in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) of southeastern Australia

Kinship has been shown to be an important correlate of group membership and associations among many female mammals. In this study, we investigate association patterns in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) inhabiting an embayment in southeastern Australia. We combine the behavioral data with microsatellite DNA and mitochondrial DNA data to test the hypotheses that genetic relatedness and maternal kinship correlate with associations and social clusters. Mean association between females was not significantly different from a random mean, but the standard deviation was significantly higher than a random standard deviation, indicating the presence of nonrandom associates in the dataset. A neighbor-joining tree, based on the distance of associations between females, identified four main social clusters in the area. Mean genetic relatedness between pairs of frequent female associates was significantly higher than that between pairs of infrequent associates. There was also a significant correlation between mtDNA haplotype sharing and the degree of female association. However, the mean genetic relatedness of female pairs within and between social clusters and the proportion of female pairs with the same and different mtDNA haplotypes within and between clusters were not significantly different. This study demonstrates that kinship correlates with associations among female bottlenose dolphins, but that kinship relations are not necessarily a prerequisite for membership in social clusters. We hypothesize that different forces acting on female bottlenose dolphin sociality appear to promote the formation of flexible groups which include both kin and nonkin.     

Kinship and aggression: do house sparrows spare their relatives?

Kin-selection theory predicts that relatedness may reduce the level of aggression among competing group members, leading to indirect fitness benefits for kin-favoring individuals. To test this hypothesis, we investigated whether relatedness affects aggressive behavior during social activities in captive house sparrow (Passer domesticus) flocks. We found that sparrows did not reduce their aggression towards kin, as neither the frequency nor the intensity of fights differed between close kin and unrelated flock-mates. Fighting success was also unrelated to kinship and the presence of relatives in the flock did not influence the birds’ dominance rank. These results suggest that the pay-offs of reduced aggression towards kin may be low in non-breeding flocks of sparrows, e.g. due to competition among relatives as predicted by a recent refinement of kin-selection theory. Our findings indicate that the significance of kin selection may be restricted in some social systems such as winter aggregations of birds. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The influence of refuge sharing on social behaviour in the lizard Tiliqua rugosa

Refuge sharing by otherwise solitary individuals during periods of inactivity is an integral part of social behaviour and has been suggested to be the precursor to more complex social behaviour. We compared social association patterns of active versus inactive sheltering individuals in the social Australian sleepy lizard, Tiliqua rugosa, to empirically test the hypothesis that refuge sharing facilitates social associations while individuals are active. We fitted 18 neighbouring lizards with Global Positioning System (GPS) recorders to continuously monitor social associations among all individuals, based on location records taken every 10 min for 3 months. Based on these spatial data, we constructed three weighted, undirected social networks. Two networks were based on empirical association data (one for active and one for inactive lizards in their refuges), and a third null model network was based on hypothetical random refuge sharing. We found patterns opposite to the predictions of our hypothesis. Most importantly, association strength was higher in active than in inactive sheltering lizards. That is, individual lizards were more likely to associate with other lizards while active than while inactive and in shelters. Thus, refuge sharing did not lead to increased frequencies of social associations while lizards were active, and we did not find any evidence that refuge sharing was a precursor to sleepy lizard social behaviour. Our study of an unusually social reptile provides both quantitative data on the relationship between refuge sharing and social associations during periods of activity and further insights into the evolution of social behaviour in vertebrates.     

Diet of the social groups of long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas</Emphasis>) in the Strait of Gibraltar

The Strait of Gibraltar is inhabited throughout the year by a group of pilot whales (Globicephala melas), but their spatial distribution varies between Summer and Autumn. In this paper, we have used carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) stable isotope signatures to investigate the differences in diet amongst seasons, sex and stable social units. Skin samples were collected from 56 individually photo-identified pilot whales during Autumn 2005 and Summer 2006. These individuals were genetically sexed and their isotopic signature determined. The level of inter-individual association both within and between stable social units were compared to Euclidean distances between individual isotopes signatures. No differences in either δ¹⁵N or δ¹³C were found according to the sex of individuals, but significant seasonal differences were found in δ¹⁵N, although not in the δ¹³C values. This suggests that pilot whales are resident year round in the Strait, a finding supported by independent photo-identification. The variation in δ¹⁵N could reflect a shift in pilot whale diet through the year, with pilot whales feeding at a higher trophic level in Autumn compared to Summer. This could also represent a change in the diet of pilot whale prey species. The δ¹³C values were significantly different amongst the four stable social units sampled and individual δ¹³C values were significantly related to the level of inter-individual association, while no relationship was found for δ¹⁵N. These results suggest that within the same general area (i.e. the Strait of Gibraltar), there is some level of specialisation in habitat or prey choice between pilot whales social units.     

The genetic population structure of the gray mouse lemur (<Emphasis Type="Italic">Microcebus murinus), </Emphasis>a basal primate from Madagascar

The genetic structure of a population is closely connected to fundamental evolutionary processes and aspects of social behavior. Information on genetic structure is therefore instrumental for the interpretation of social behavior and evolutionary reconstructions of social systems. Gray mouse lemurs (Microcebus murinus) are basal primates endemic to Madagascar whose social organization is characterized by solitary foraging at night and communal resting during the day. Conflicting reports about population structure based on behavioral observations led us to examine the genetic structure of one population in detail in order to: (1) identify natural genetic units in this solitary primate, and (2) to test the assumption of current models of primate social evolution that solitary primates are organized in matrilines. DNA was extracted from tissue samples of 85 individuals from Kirindy forest to determine their variability at a 530 bp fragment of the mitochondrial D-loop and at six microsatellite loci. We found that this population was characterized by a great general diversity among mtDNA haplotypes, a pronounced sex difference in mtDNA haplotype diversity and spatial clustering of females with a particular haplotype, but low average relatedness among members of haplotype clusters. Specifically, we identified 13 different haplotypes, which were unevenly distributed among individuals. About 80% of all individuals, most of which were females or juvenile males, shared a single haplotype. Rare haplotypes were almost exclusively represented by single adult males, who apparently migrated into this population. One other haplotype was represented by a small group of females living at one edge of the study area. Microsatellite analysis revealed above-average relatedness among females with overlapping home ranges, as well as no signs of inbreeding, implying that male dispersal results in high levels of gene flow among matrilineal groups. We conclude that gray mouse lemur populations are hierarchically organized in small family units of closely related females that form stable sleeping groups, several of which are connected through a common mtDNA haplotype and form spatially distinct clusters. The presence of such matrilines supports a basic assumption of current models of primate social evolution.     

Sperm design and variation in the New World blackbirds (Icteridae)

Post-copulatory sexual selection (PCSS) is thought to be one of the evolutionary forces responsible for the rapid and divergent evolution of sperm design. However, whereas in some taxa particular sperm traits are positively associated with PCSS, in other taxa, these relationships are negative, and the causes of these different patterns across taxa are poorly understood. In a comparative study using New World blackbirds (Icteridae), we tested whether sperm design was influenced by the level of PCSS and found significant positive associations with the level of PCSS for all sperm components but head length. Additionally, whereas the absolute length of sperm components increased, their variation declined with the intensity of PCSS, indicating stabilising selection around an optimal sperm design. Given the diversity of, and strong selection on, sperm design, it seems likely that sperm phenotype may influence sperm velocity within species. However, in contrast to other recent studies of passerine birds, but consistent with several other studies, we found no significant link between sperm design and velocity, using four different species that vary both in sperm design and PCSS. Potential reasons for this discrepancy between studies are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Do social networks of female northern long-eared bats vary with reproductive period and age?

Social structure, which is a function of the patterns of interactions among individuals, is particularly variable in fission–fusion societies. The underlying factors that drive this variation are poorly understood. Female northern long-eared bats (Myotis septentrionalis) live in fission–fusion societies where females form preferred associations within groups that vary daily in size and composition as individuals switch roosts. The goal of our study was to test the predictions that preferred associations and social networks of female northern long-eared bats vary with reproductive period and age. We also tested the prediction that preferred relationships persist across years despite movements from summer roosts to winter hibernacula. Network analyses revealed that during gestation, females roosted in smaller groups where they roosted more regularly with fewer individuals than during lactation. This variation may reflect different social strategies to mediate higher energetic costs during lactation. Females of all ages roosted more often with younger individuals, which in turn had more direct and indirect associations than all other age classes. Younger individuals may play a role in maintaining connections between individuals, perhaps as a result of younger individuals being more exploratory. Temporal analyses suggested that relationships can persist for years as some pairs roosted together for multiple summers. We suggest that the dynamic nature of fission–fusion societies is associated with individual strategies to increase fitness relative to individual characteristics, in this case reproductive condition and age.     

Sighting characteristics and photo-identification of Cuvier’s beaked whales (<Emphasis Type="Italic">Ziphius cavirostris</Emphasis>) near San Clemente Island,California: a key area for beaked whales and the military?

The relationship between beaked whales and certain anthropogenic sounds remains poorly understood and of great interest. Although Cuvier’s beaked whales (Ziphius cavirostris) are widely distributed, little is known of their behavior and population structure throughout much of their range. We conducted a series of five combined visual-acoustic marine mammal surveys from 2006 to 2008 in the southern San Nicolas Basin, a site of frequent naval activity off the southern California coast, west of San Clemente Island. The study area was defined by a 1,800 km² array of 88 bottom-mounted hydrophones at depths up to 1,850 m. The array was used to vector visual observers toward vocalizing marine mammal species. Thirty-seven groups of Cuvier’s beaked whales were encountered during the study period. The overall encounter rate was one group for every 21.0 h of survey effort, and was as high as one group per 10.2 h of effort during the October 2007 survey. Whales were encountered in the deepest portion of the study area, at a mean bottom depth of 1,580 m (SD 138). The average group size was 3.8 individuals (SD 2.4), which was higher than has been reported from other studies of this species. Twenty-four groups were observed over multiple surfacings (median = 4 surfacings, range 2–15). The mean encounter duration of extended sightings was 104 min (SD 98, range 12–466 min) and the mean distance moved over the course of sightings was 1.66 km (SD 1.56, range 0.08–6.65 km). Temporal surfacing patterns during extended encounters were similar to dive behavior described from Cuvier’s beaked whales carrying time-depth recording tags. Seventy-eight photographic identifications were made of 58 unique individuals, for an overall resighting rate of 0.26. Whales were sighted on up to 4 days, with duration from first to last sighting spanning 2–79 days. For those whales sighted on subsequent days, the mean distance between subsequent sightings was 8.6 km (SD 7.9). Individuals resighted over 2–3 days were usually in association with previous group members. Approximately one-third of groups contained more than one adult male, and many of the repeated associations involved adult males. These observations suggest the basin west of San Clemente Island may be an important region for Cuvier’s beaked whales, and also one which affords an unusual opportunity to collect detailed data on this species. Given its status as an active military range, it can also provide the ability to monitor the behavior of individuals in the presence of naval sonar, a critical step in the management of this and other beaked whale populations worldwide.     

Mating strategies of queens in <Emphasis Type="Italic">Lasius niger</Emphasis> ants—is environment type important?

High relatedness among society members is believed important for the evolution of highly cooperative behaviours, yet queens of many social insects mate with multiple males which reduces nestmate relatedness and imposes also direct costs on queens. While theoretical models have suggested explanations for this puzzling queen behaviour, empirical studies fail to provide consistent answers especially for species with moderate levels of multiple mating. This may result from multiple mating only conferring benefits in some environments, as suggested by recent genetic variance theory and considerations on types of traits, direct costs and benefits. All concur in an expectation of higher levels of multiple mating in more complex or milder environments, and we perform a first, broad test of this idea by comparing mating strategies of queens in Lasius niger ants from northern (harsh, cold stressed) and southern populations (milder, greater bio-complexity). First, we collected new genetic data from Ireland and Southern France and then compared these to data on Swiss and Swedish populations. Queens from northern populations were near exclusively single mated and even at times inbred (in Ireland), whereas southern queens showed high levels of multiple mating, leading to more genetically diverse colonies in the south. Equally, paternity skew was greater in the north, as expected if northern queens only remate when their first mate transfers few sperm. Our findings are consistent with the idea that environment type may affect mating strategies in social insects and calls for an exploration of such effects.     

Spatial and social connectivity of fish-eating “Resident” killer whales (Orcinus orca) in the northern North Pacific

The productive North Pacific waters of the Gulf of Alaska, Aleutian Islands and Bering Sea support a high density of fish-eating “Resident” type killer whales (Orcinus orca), which overlap in distribution with commercial fisheries, producing both direct and indirect interactions. To provide a spatial context for these interactions, we analyzed a 10-year dataset of 3,058 whale photo-identifications from 331 encounters within a large (linear ~4,000 km) coastal study area to investigate the ranging and social patterns of 532 individually identifiable whales photographed in more than one encounter. Although capable of large-scale movements (maximum 1,443 km), we documented ranges generally <200 km, with high site fidelity across summer sampling intervals and also re-sightings during a winter survey. Bayesian analysis of pair-wise associations identified four defined clusters, likely representing groupings of stable matrilines, with distinct ranging patterns, that combined to form a large network of associated whales that ranged across most of the study area. This provides evidence of structure within the Alaska stock of Resident killer whales, important for evaluating ecosystem and fisheries impacts. This network included whales known to depredate groundfish from longline fisheries, and we suggest that such large-scale connectivity has facilitated the spread of depredation.     

The bottlenose dolphin community of Doubtful Sound features a large proportion of long-lasting associations

More than 12 studies of different bottlenose dolphin populations, spanning from tropical to cold temperate waters, have shown that the species typically lives in societies in which relationships among individuals are predominantly fluid. In all cases dolphins lived in small groups characterised by fluid and dynamic interactions and some degree of dispersal from the natal group by both sexes. We describe a small, closed population of bottlenose dolphins living at the southern extreme of the species' range. Individuals live in large, mixed-sex groups in which no permanent emigration/immigration has been observed over the past 7 years. All members within the community are relatively closely associated (average half-weight index>0.4). Both male–male and female–female networks of preferred associates are present, as are long-lasting associations across sexes. The community structure is temporally stable, compared to other bottlenose dolphin populations, and constant companionship seems to be prevalent in the temporal association pattern. Such high degrees of stability are unprecedented in studies of bottlenose dolphins and may be related to the ecological constraints of Doubtful Sound. Fjords are low-productivity systems in which survival may easily require a greater level of co-operation, and hence group stability. These conditions are also present in other cetacean populations forming stable groups. We therefore hypothesise that ecological constraints are important factors shaping social interactions within cetacean societies.Communicated by D. Watts     

Genetic reconstruction of breeding patterns in gundis (Rodentia: Ctenodactylidae)

Current mating system theory predicts that the number of females breeding in a group will depend on the number of females in the group and the accessibility of unrelated males, whereas the number of males breeding in a group will depend on the ability of males to control access to reproductive females. By combining information on group composition with genetic data, I determined whether breeding patterns in a rock-dwelling rodent, Ctenodactylus gundi, were concordant with these expectations. C. gundi breeding patterns varied from facultative monogamy to uni-male polygyny to multi-male polygyny. The number of reproductive individuals of each sex in a group increased with group size. Whereas communal breeding among related females tended to increase female reproductive success, males breeding in the same group were unrelated to other group members and seemed to compete over access to matings. While some males were assigned offspring from neighboring social groups, most group-living males successfully monopolized the reproduction of females within their group. There was no evidence that females had multiply sired litters, although some bred with alternative males in separate breeding attempts. Although numerous individuals were not assigned as parents or offspring, genetic information enabled me to determine that most unassigned individuals were philopatric group members, whereas only a few were unrelated immigrants into their current social group. Together, these results provide evidence that C. gundi social groups represent fairly distinct reproductive units whose breeding patterns are dependent on group size and composition in accordance with theoretical predictions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Relatedness structure and kin-biased foraging in the greater horseshoe bat (Rhinolophus ferrumequinum)

Female greater horseshoe bats (Rhinolophus ferrumequinum) exhibit strong natal philopatry to their maternity roost over many years, leading to the aggregation of matrilineal kin. Maternity colonies may, therefore, be expected to comprise highly related individuals, and, as such, provide conditions suitable for the evolution of kin-selected behaviours. To test these predictions, we examined relatedness and behaviour among matrilineal kin within a colony in south-west Britain. Genetic analysis of 15 matrilines, identified from microsatellite genotyping and long-term ringing surveys, revealed average relatedness levels of 0.17 to 0.64. In contrast, background relatedness among colony females approximated to zero (0.03). These results suggest that inclusive fitness benefits may only be accrued through discriminate cooperation within matrilines, and not at the wider colony level. To examine whether the potential for such benefits is realised through kin- biased cooperation during foraging, females from two matrilines were radio-tracked simultaneously over 3 years. Pairwise home-range overlap correlated significantly with Hamilton's relatedness coefficient. The greatest spatial associations were observed between females and their adult daughters, which shared both foraging grounds and night roosts, sometimes over several years. Tagged females, however, generally foraged and roosted alone, suggesting that kin-biased spatial association probably does not result from either information-transfer or cooperative territorial defence. Such patterns may instead result from a mechanism of maternal inheritance of preferred foraging and roosting sites.     

Insights on common dolphin (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) social organization from genetic analysis of a mass-stranded pod

Compared to terrestrial mammals, little is known of cetacean social systems as they are generally less accessible to behavioral investigations due to their aquatic environment. The present study investigates group structure of the pelagic common dolphin, Delphinus delphis, using genetic markers. Tissue samples from 52 individuals representing a recent live mass-stranding event were compared to 42 single strandings taken from presumably different groups. The mass-stranding event occurred in 2002 on the French coast of the English Channel, whereas the single strandings were collected between 1993 and 2003 along the western coast of France (Bay of Biscay and English Channel). Analysis of mitochondrial DNA control region sequences indicated that genetic variability within the mass-stranded pod was similar to variability observed in single strandings. The mass-stranded group was composed of 41 different mitochondrial haplotypes or matrilines while the single strandings revealed 29 different haplotypes. Analysis of 11 microsatellite loci revealed that average relatedness of the mass-stranded pod was not different from average relatedness among all single strandings suggesting that individuals within the group had no closer kin relationships than animals taken from presumably different groups. These results do not support a matriarchal system and suggest that common dolphins constituting a pod are not necessarily genetically related.     

Association patterns among wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) reflect sex differences in cooperation

Theory predicts that frequent dyadic association should promote cooperation through kin selection or social tolerance. Here we test the hypothesis that sex differences in the strength and stability of association preferences among free-ranging chimpanzees conform to sex differences in cooperative behavior. Using long-term data from the Kanyawara chimpanzee (Pan troglodytes schweinfurthii) community (Kibale National Park, Uganda), we calculated indices of intra-sexual dyadic association over a 10-year period. We found that (1) male–male dyads had significantly stronger association indices than female–female dyads, (2) the pattern of association preferences in both sexes changed little over the entire study period, and (3) when comparing periods with different alpha males, changes in association strength were more frequent among males. These results demonstrate that both the strength and stability of association patterns are important components of social relationships. Male chimpanzees, which are characterized by frequent cooperation, had association preferences that were both strong and stable, suggesting that forming long-term bonds is an important dominance strategy. However, the fact that male association patterns were sensitive to upheaval in the male dominance hierarchy suggests that males also take advantage of a changing social climate when choosing association partners. By contrast, the overall strength of female associations was relatively weak. Female association preferences were equally stable as males’; however, this reflected a dyad’s tendency to be found in the same party rather than to associate closely within that party. Therefore, in this community, female association patterns appear to be more a consequence of individual ranging behavior rather than a correlate of cooperation.     

Local genetic structure and relatedness in a solitary mammal, Neotoma fuscipes

We used DNA microsatellites to investigate temporal and spatial patterns of local genetic differentiation and relatedness in a solitary mammal, the dusky-footed woodrat (Neotoma fuscipes). Patterns of genetic variation were measured relative to spatial clusters, or neighborhoods, of woodrats. We detected significant genetic differentiation among woodrat neighborhoods in two populations spanning multiple habitat types and densities. Estimates of θ _ST among neighborhoods ranged 0.034–0.075 and were comparable to levels reported in social mammals. Genetic differentiation at such a local scale is noteworthy because it occurred in the absence of any physical barriers to gene flow, suggesting that the patterns observed are linked to the nonrandom patterns of mating and dispersal that characterize woodrat social structure. Genetic differentiation and relatedness among neighborhoods were even higher when only resident females were analyzed. These results are consistent with a pattern of female philopatry and male-biased dispersal in woodrats. Geographic distance and relatedness were inversely correlated in adult females at intermediate densities, but not at low densities. Nonetheless, matrilineal genetic structure was apparent even at low woodrat densities based on estimates of θ _ST among neighborhoods of resident females that were significantly greater than zero and consistently greater than estimates including all individuals. In summary, this study demonstrates a matrilineal genetic structure in dusky-footed woodrats. In addition, our results support the idea that intermediate densities may be better at facilitating the formation of spatial kin clusters than either extreme. An erratum to this article can be found at