Kinship,reproductive strategies and social dynamics of yellow-bellied marmots

Summary Social behavior of yellow-bellied marmots was observed for three years in colonies where kinship was known and for one year in a high elevation colony where harems were contiguous. Social dynamics of yellow-bellied marmots is dependent on kinship, group composition, and age-sex classes. This pattern is a consequence of the reproductive strategies of males and females. Females behave cohesively toward sisters or daughters, but not with sons and agonistically toward other females. Males generally behave amicably toward females and agonistically toward males, including their sons. Thus, reproductive strategies limit nepotism. This behavior is consistent with a population process in which sons typically disperse as yearlings. At least some of the variation in the expected patterns of social behaviors is attributable to individual differences. Because male and female reproductive strategies differ, a marmot population consists of two social subsystems. The female unit is the closely-related kin group which may also be a burrow group. The male unit is a harem which usually consists of two or more competing female kin groups.     

Dispersal decisions and predispersal behavior in <Emphasis Type="Italic">Polistes</Emphasis> paper wasp ‘workers’

Social insects are popular models for studying the evolution of cooperation. Casteless taxa where individuals have the flexibility to either nest alone or cooperate are particularly valuable for understanding the causes and consequences of cooperative behavior. For example, some ‘workers’ from Polistes paper wasp nests disappear from their natal colony soon after pupal emergence and nest independently. However, little is known about dispersal behavior. In this paper, I compare predispersal behavior of wasps who leave their natal colony soon after emergence with behavior of individuals who remain on the natal colony as true workers. I found that P. dominulus females with short nest tenure behave much like gynes (reproductive-destined offspring produced at the end of the season), as wasps with short nest tenure are behaviorally selfish while on the natal colony. They spend a smaller proportion of their time foraging and a larger proportion of their time resting than workers with long nest tenure. In addition, I assessed the factors that may favor early dispersal. Nest environment strongly influenced dispersal; large colonies had a smaller proportion of females with short nest tenure. Queen turnover also increased dispersal behavior perhaps because queen turnover reduces relatedness between a colony’s current and future offspring, thereby reducing the kin-selected benefits of cooperation. Therefore, casteless social insects exhibit a surprising degree of reproductive flexibility. Individuals may use information about their internal state and nest environment to optimize their behavioral strategies.     

Movements of dispersing flying squirrels in relation to siblings and parents

Knowledge of kin interactions can be informative in explaining the processes underlying dispersal. By dispersing, relatives can avoid kin competition for resources or mates and prevent inbreeding. We investigated sibling movements in relation to each other and parents before and after dispersal in radio-collared juvenile flying squirrels (Pteromys volans L.). Before dispersal, most siblings moved in different areas and were not in contact while exploring the surroundings of natal area. After dispersal, all siblings settled far away from each other, as they dispersed different distances and to random directions compared to each other. No clear effect of litter size or presence of same-sex siblings was observed on dispersal. Time spent in the same nest with the mother did not differ between dispersing sons and daughters. Mother did not force dispersers to leave the natal nest; instead, the mother often moved away from the nest before offspring. Father and offspring were not observed to be in contact before dispersal. Dispersal direction was unaffected by the location of fathers’ home range. We conclude that after dispersal interaction between relatives is limited in flying squirrels, but we did not find any indication that interaction between relatives before dispersal is determining dispersal decision. Siblings’ settlement far away from each other was already indicated by the movement within the natal home range, but, interestingly, the latter could not be used to predict dispersal patterns of sexes.     

Dispersal behaviour and colony structure in a colonial spider

Group living in spiders is characterised by two principle modes, the cooperative social mode and the colonial non-cooperative mode. Kin-relationships due to reduced dispersal determine population genetic structure in social spiders, but the dispersal mechanisms underlying group structure remain poorly understood in colonial spiders. Assuming similar ecological benefits of group living, we address the question whether reduced dispersal shapes population structure in a colonial spider, Cyrtophora citricola (Araneidae). We analysed dispersal by studying settling decisions under semi-natural conditions in experimental trees with and without colonies, and in natural populations, we estimated dispersal and colony structure using population genetic analyses. The propensity to disperse decreased with increasing age in experimental colonies. Adult females did not disperse in the experiment. Sub-adult female spiders preferred trees with a colony to trees without a colony. Dispersal in third instar juveniles was influenced significantly by wind but not by the presence of a colony. Thus, we showed that being in a colony did not inhibit juvenile dispersal, but pre-mating females were philopatric. Genetic differentiation among colonies in natural populations was heterogeneous, colonies being either little or highly differentiated. The heterogeneous structure is likely caused by colony founding by one or a few females followed by dispersal among perennial colonies. Gene flow, however, was slightly male-biased. The experimental and indirect, genetic approaches combined showed that dispersal and the breeding system of C. citricola resemble that of solitary spiders, with juvenile dispersal occurring in both sexes, while the colonial distribution is maintained by female philopatry.     

Is kin cooperation going on undetected in marine bird colonies?

In multitudinous breeding colonies, kin interactions could go unnoticed because we are unaware of the kinship among adults we observe. Evidence of cooperation and competition between close adult kin in a blue-footed booby colony was sought by analyzing patterns of natal dispersal and proximity of nests. Male and female recruits nested closer to their own natal sites than to their parents’ current sites. Males (only) dispersed less far when both parents were present than when no parent or one parent was present, but not selectively close to fathers versus mothers when these were divorced. Neither parental presence nor parental proximity affected breeding success of recruits of either sex. Although distances between the nests of simultaneously recruiting broodmates were unrelated to their sex, males dispersed 13.1 m less when a sister was present than when a brother was present. Neither sex was affected in its dispersal distance by the presence or hatching order/dominance of a broodmate. Neither sex was affected in its breeding success by the presence versus absence of a broodmate, although female success increased with proximity of their brothers. Parents and sisters may actively or passively help males establish their first territories near their natal sites and nearby brothers may help females in their first breeding attempts; otherwise, boobies do not influence each other’s natal dispersal and first breeding success. It appears that boobies do not nest selectively close to or far from their parents, offspring, or broodmates. Why there is apparently so little cooperation and altruism between close adult relatives in booby colonies is puzzling.     

Levels of social behaviors and genetic structure in a population of round-tailed ground squirrels (Xerospermophilus tereticaudus)

Ground-dwelling sciurids exhibit a continuum of sociality and several models predict levels of sociality within this taxon. Models of ground squirrel sociality predict round-tailed ground squirrels (Xerospermophilus tereticaudus) to be solitary; however, previous behavioral studies suggest round-tailed ground squirrels have a matrilineal social structure. To resolve this discrepancy, we combined behavioral observations with genetic analyses of population structure. We assessed levels of agonistic and amicable behaviors combined with fine-scale population genetic structure of round-tailed ground squirrels in a multi-year study in AZ. Only 45 agonistic and 40 amicable interactions were observed between adults in over 137 h of observations. Overall rates of agonistic or amicable interactions between adults were low (≤0.69/h), with no relationship between relatedness of individuals and rates of either amicable or agonistic interactions. Interactions between juvenile littermates were predominantly amicable. Population substructure was not evident with Bayesian analyses, global or pairwise F _ST values; average relatedness among females was not different from males. However, in 2006, the year after a population reduction through targeted animal elimination, a population bottleneck was detected within at least five of seven loci. Contrary to previous behavioral studies, this population of round-tailed ground squirrels, although aggregated spatially, did not exhibit high levels of social behavior nor subpopulation genetic structure. Analyses of the genetic relationships and sociality along a continuum, particularly within aggregates of individuals, may lead to insights into the origin and maintenance of social behaviors by elucidating the mechanisms by which aggregates with intermediate social levels are formed and maintained.     

Studying dispersal at the landscape scale: efficient combination of population surveys and capture-recapture data

Researchers often rely on capture-mark-recapture (CMR) data to study animal dispersal in the wild. Yet their spatial coverage often does not encompass the entire dispersal range of the study individuals, sometimes producing misleading results. Information contained in population surveys and variation in population spatial structure can be used to overcome this issue. We build an integrated model in a multisite context in which CMR data are only collected at a subset of sites, but numbers of breeding pairs are counted at all sites. In a Black-headed Gull Chroicocephalus ridibundus population, the integrated-modeling approach induces an increase in precision for the demographic parameters of interest (variances, on average, were decreased by 20%) and provides a more precise extrapolation of results from the CMR data to the whole population. Patterns of condition-dependent dispersal are therefore made easier to detect, and we obtain evidence for colony-size dependence in recruitment, dispersal, and breeding success. These results suggest that first-time breeders disperse to small colonies in order to recruit earlier. The exchange of experienced breeders between colonies appears as a main determinant of the observed variation in colony sizes.     

Effects of Habitat Fragmentation on Effective Dispersal of Florida Scrub‐Jays

Abstract: Studies comparing dispersal in fragmented versus unfragmented landscapes show that habitat fragmentation alters the dispersal behavior of many species. We used two complementary approaches to explore Florida Scrub‐Jay (Aphelocoma c?rulescens) dispersal in relation to landscape fragmentation. First, we compared dispersal distances of color‐marked individuals in intensively monitored continuous and fragmented landscapes. Second, we estimated effective dispersal relative to the degree of fragmentation (as inferred from two landscape indexes: proportion of study site covered with Florida Scrub‐Jay habitat and mean distance to nearest habitat patch within each study site) by comparing genetic isolation‐by‐distance regressions among 13 study sites having a range of landscape structures. Among color‐banded individuals, dispersal distances were greater in fragmented versus continuous landscapes, a result consistent with other studies. Nevertheless, genetic analyses revealed that effective dispersal decreases as the proportion of habitat in the landscape decreases. These results suggest that although individual Florida Scrub‐Jays may disperse farther as fragmentation increases, those that do so are less successful as breeders than those that disperse short distances. Our study highlights the importance of combining observational data with genetic inferences when evaluating the complex biological and life‐history implications of dispersal.     

Dispersal and Use of Corridors by Birds in Wooded Patches on an Agricultural Landscape

Dispersal behavior has important effects on the persistence and recolonization of populations, but is one of the least understood traits of most organisms. Knowledge of patterns of fledgling, natal, and breeding dispersal of birds in a patchy environment will assist in decisions regarding reserve design and protection or construction of corridors. I present data on movement patterns of three migratory bird species, American Robin ( Turdus migratorius ), Brown Thrasher ( Toxostoma rufum ), and Loggerhead Shrike ( Lanius ludovicianus ). These birds are relatively common breeders in south-central North Dakota (U.S.) in riparian woodlands and in shelterbelts (woodlots planted as windbreaks in the open agricultural environment). Field assistants and I individually marked and monitored the movements of more than 500 adults breeding in a network of shelterbelts across an 8 × 11 km study area. Most movement occurred at relatively short distances within a shelterbelt. Movements by adults between shelterbelt sites, although rare, occurred significantly more frequently between sites connected by a wooded corridor than between unconnected sites. For robins, there were on average 2.5 dispersal events between each pair of connected sites, but only 0.17 dispersal events between each pair of unconnected sites (Mann-Whitney test, significant at p < 0.009). Because unconnected and connected sites were similar in average area (1.7 to 1.9 ha), distance to next wooded habitat, and tree-species composition, this result provides a test of the hypothesis that organisms disperse preferentially along connecting corridors.     

Delayed dispersal as a potential route to cooperative breeding in ambrosia beetles

Xyleborini are a species-rich tribe of ambrosia beetles, which are haplodiploid and typically mate among siblings within their natal brood chamber. Several characteristics of this tribe would predict the evolution of higher levels of sociality: high genetic relatedness within galleries due to inbreeding, high costs of dispersal and the potential benefit of cooperation in brood care within the natal gallery (e.g. by fungus gardening, gallery extension, offspring feeding and cleaning). However, information on the social system of these beetles is very limited. We examined the potential for cooperative breeding in Xyleborinus saxeseni by monitoring dispersal in relation to brood size and composition. Results show that adult female offspring delay dispersal despite dispersal opportunities, and apparently some females never disperse. The females’ decision to stay seems to depend on the presence of eggs and dependent siblings. We found no indication that female offspring reproduce in their natal gallery, as colonies with many mature daughters do not contain more eggs than those with few or no daughters. There is a significant positive relationship between the number of females present and the number of dependent siblings (but not eggs), which suggests that cooperative brood care of female offspring raises colony productivity by improving survival rates of immatures. Our results suggest that cooperative breeding is likely to occur in X. saxeseni and possibly other xyleborine species. We argue that a closer look at sociality within this tribe may yield important information on the factors determining the evolution of cooperative breeding and advanced social organization.     

Kin associations and direct vs indirect fitness benefits in colonial cooperatively breeding sociable weavers Philetairus socius

Indirect fitness benefits are believed to be an important force behind the evolution of cooperative breeding. However, helpers may associate with their relatives as a result of delayed dispersal, hence, kin associations might be a consequence of demographic viscosity rather than active choice. In addition, recent studies showed that helpers may have access to reproduction therefore direct benefits might also play an important role. Here, we investigate the possible roles of direct genetic benefits and kin associations on helping behavior in the sociable weaver Philetairus socius, a colonial and cooperatively breeding passerine. We used a microsatellite-based genotyping method to describe the genetic structure within nests and colonies. Within a colony, we found considerable genetic structure between males but not females. Sociable weaver colonies have several nests that are simultaneously active, giving individuals a choice of associating with a range of first-order kin to unrelated individuals. Helpers were significantly more related to the young in the helped nests than in other nests of the colony, suggesting an active choice for associating with kin. The helpers were generally offspring or first-order relatives of one (50%) or both (43%) breeders, although more infrequently, seemingly unrelated individuals also helped (7%). We found no supporting evidence of extrapair parentage and hence no direct genetic gains from helping in our population. This strong reproductive skew is contrary to theoretical models predicting conflicts over reproduction in stepfamilies. We discuss whether female decisions and/or other direct benefits of remaining in kin associations or helping might explain the high skew observed.     

High social density increases foraging and scouting rates and induces polydomy in Temnothorax ants

Many organisms live in crowded groups where social density affects behavior and fitness. Social insects inhabit nests that contain many individuals where physical interactions facilitate information flow and organize collective behaviors such as foraging, colony defense, and nest emigration. Changes in nest space and intranidal crowding can alter social interactions and affect worker behavior. Here, I examined the effects of social density on foraging, scouting, and polydomy behavior in ant colonies—using the species Temnothorax rugatulus. First, I analyzed field colonies and determined that nest area scaled isometrically with colony mass—this indicates that nest area changes proportionally with colony size and suggests that ants actively control intranidal density. Second, laboratory experiments showed that colonies maintained under crowded conditions had greater foraging and scouting activities compared to the same colonies maintained at a lower density. Moreover, crowded colonies were significantly more likely to become polydomous. Polydomous colonies divided evenly based on mass between two nests but distributed fewer, heavier workers and brood to the new nests. Polydomous colonies also showed different foraging and scouting rates compared to the same colonies under monodomous conditions. Combined, the results indicate that social density is an important colony phenotype that affects individual and collective behavior in ants. I discuss the function of social density in affecting communication and the organization of labor in social insects and hypothesize that the collective management of social density is a group level adaptation in social insects.     

Dispersal, pair formation and social structure in gibbons (Hylobates lar)

We report observations on reproduction, natal dispersal, pair formation, and group structure based on longitudinal observations of several white-handed gibbon (Hylobates lar) groups spanning 18 years. Our observations are at odds with the traditional view that gibbons live in nuclear family groups consisting of a pair of adults and their offspring, and that parents exclude young from the family territory when they reach adult size. In the relatively dense Khao Yai study population, dispersing young usually obtain mates by replacing adults in existing territories, which creates non-nuclear families. Six subadults, five males and one female, matured and dispersed at an average age of 10 years, or about 2 years after reaching adult size. Average natal dispersal distance was 710 m, or between one and two territories away. At least two dispersing males replaced adults in neighboring groups. In one case, forcible displacement of the resident male resulted in a group which included a young juvenile presumably fathered by the previous male, two younger juveniles (probably brothers) from the new male's original group, and (later) offspring of the new pair. Social relations within this heterogeneous group remained harmonious: the adults groomed all the young and play occurred between all preadult members. In only two out of a total of seven cases of dispersal seen did two subadults pair and disperse into new territorial space. Nonreproducing subadults which delay dispersal may be tolerated by the adults provided that they contribute benefits to the adults or their offspring. Possible benefits include behaviors such as grooming, social play with juveniles, and support of the adult male in defending the territory. Delayed dispersal is probably advantageous in a saturated environment where there is no room for floaters, but subadults may also gain indirect fitness benefits by aiding siblings and other relatives. Received: 24 January 1997 / Accepted after revision: 12 January 1998     

Small queens in the ant Ectatomma tuberculatum: a new case of social parasitism

Social parasites exploit the worker force of colonies of other social insects to rear their own young. Social parasitism occurs in several Hymenoptera and is particularly common in several tribes of the ant subfamilies Myrmicinae and Formicinae. Here, we document the occurrence of miniaturized queens (microgynes) in colonies of Ectatomma tuberculatum, an ant belonging to the subfamily Ectatomminae. Behavioral observations and genetic analyses show that microgynes concentrate their reproductive efforts almost exclusively on the production of sexual offspring (microgynes and males), whereas the regular, large queens (macrogynes) produce workers in addition to sexuals. According to mitochondrial and nuclear markers, gene flow between microgynes and macrogynes is extremely limited. Whereas the co-occurrence of microgynes and macrogynes in the related species Ectatomma ruidum constitutes an intraspecific polymorphism associated with alternative dispersal tactics, microgynes found in colonies of E. tuberculatum appear to be a distinct species and to represent the first case of social parasitism in the poneromorph subfamilies of ants.     

Nomadic behavior of army ants in a desert-grassland habitat

Summary Nomadic behavior of the army ant Neivamyrmex nigrescens was studied in a desert-grassland habitat. Six colonies were followed through eight nomadic phases (94 nomadic days) while direction and distance of emigrations, growth of larvae, number of adults and larvae per colony were determined.In all colonies, the nomadic phase began when newly eclosed adults and small larvae were present, and ended when the larvae were fully grown. Average emigration distance was positively related to number of larvae in the colony. These findings support Schneirla's theory that brood stimulation is a proximate cause of the nomadic phase.Failures to emigrate were equally likely at all points in the nomadic phase, and there was no systematic increase in emigration distance as the phase progressed. These findings do not support Schneirla's version of brood-stimulative theory.Number of adults per colony was positively related to the directionality of the nomadic phase; however, both the direction and distance of emigrations varied unpredictably from one nomadic day to the next, in marked contrast to predictions from optimal foraging theory.Schneirla's theory is useful in predicting phase differences in colony behavior, but it does not account for characteristics such as frequency, direction, or distance of emigrations within the nomadic phase. These aspects of nomadic behavior are more closely related to characteristics of the habitat such as prey density and availability of nest sites. Army ant nomadism in this habitat may depart from the optimum because of high prey density, small colony size, or lack of nesting sites.     

Eusociality has evolved independently in two genera of bathyergid mole-rats — but occurs in no other subterranean mammal

Summary Extensive field and laboratory studies show that Damaraland mole-rats, like naked mole-rats, have an extreme form of vertebrate sociality. Colonies usually contain 2 reproductives and up to 39 non-breeding siblings, 90% of whom live a socially-induced lifetime of sterility; they remain in the natal colony, forage for food, defend the colony and care for successive litters. Although there is heightened dispersal following good rainfall, the majority of adult non-reproductives remain in their natal colony and failure to disperse is not directly attributable to habitat saturation or unfavourable soil conditions. A major dispersal event follows the death of a reproductive. Differences in colony cohesion, ethology and the hormonal profiles of non-reproductive animals suggests that eusociality evolved along different pathways in these two phylogenetically divergent genera of the Bathyergidae.     

Foraging patterns of yellow-bellied marmosts: role of kinship and individual variability

Summary Two colonies of yellow-bellied marmots (Marmota flaviventris) at an elevation of 2900 m in Colorado were studied to elucidate the role of various behavioral and ecological factors as determinants of spatial foraging patterns. The locations of known individuals were periodically recorded. These locality data were plotted as three-dimensional block diagrams, the peak heights representing the frequency of observation. Predation risk and vegetation distribution influenced the location of foraging areas; kinship was an important factor in the determination of the amount of foraging area shared between individual marmots. Spatial overlap tended to be greater among close kin, but this was modified by individual behavioral characteristics, reproductive state, the existence of separate burrow systems within a colony, and the age of the animal. Mothers and juveniles, and littermates as young and resident yearlings, had nearly identical foraging areas.     

Dispersal Can Limit Local Plant Distribution

The ability of species to establish new populations at unoccupied sites is a critical feature in the maintenance of biological diversity, and it has taken on new importance as a result of global climate change and expected changes in species distribution. To examine the dispersal potential of plant species, seeds of four annual plant species were experimentally dispersed 40 to 600 m from existing populations in Massachusetts (U.S.A.) to 34 nearby unoccupied but apparently suitable sites. At three of these sites new populations were established that persisted for four generations and expanded slowly in area. At seven sites, a small initial population eventually died out. At the 24 other sites, new populations did not become established, indicating that the sites were in some way unsuitable, that not enough seeds arrived, or that conditions suitable for seed germination do not occur every year. These results suggest that some species may be unable to disperse naturally out of their existing ranges in response to global climate change, particularly if habitat fragmentation creates barriers to dispersal. These species may have to be assisted to reach suitable sites nearby to prevent their extinction in the wild.     

Scale-dependent variation in coral community similarity across sites, islands, and island groups

Community similarity is the proportion of species richness in a region that is shared on average among communities within that region. The slope of local richness (alpha diversity) regressed on regional richness (gamma diversity) can serve as an index of community similarity across regions with different regional richness. We examined community similarity in corals at three spatial scales (among transects at a site, sites on an island, and islands within an island group) across a 10 000-km longitudinal diversity gradient in the west-central Pacific Ocean. When alpha diversity was regressed on gamma diversity, the slopes, and thus community similarity, increased with scale (0.085, 0.261, and 0.407, respectively) because a greater proportion of gamma diversity was subsumed within alpha diversity as scale increased. Using standard randomization methods, we also examined how community similarity differed between observed and randomized assemblages and how this difference was affected by spatial separation of species within habitat types and specialization of species to three habitat types (reef flats, crests, and slopes). If spatial separation within habitat types and/or habitat specialization (i.e., underdispersion) occurs, fewer species are shared among assemblages than the random expectation. When the locations of individual coral colonies were randomized within and among habitat types, community similarity was 46-47% higher than that for observed assemblages at all three scales. We predicted that spatial separation of coral species within habitat types should increase with scale due to dispersal/extinction dynamics in this insular system, but that specialization of species to different habitat types should not change because habitat differences do not change with scale. However, neither habitat specialization nor spatial separation within habitat types differed among scales. At the two larger scales, each accounted for 22-24% of the difference in community similarity between observed and randomized assemblages. At the smallest scale (transect-site), neither spatial separation within habitat types nor habitat specialization had significant effects on community similarity, probably due to the small size of transect samples. The results suggest that coral species can disperse among islands in an island group as easily as they can among sites on an island over time scales that are relevant to their establishment and persistence on reefs.     

Correlations of food distribution and patch size with agonistic interactions in female vervets (Chlorocebus aethiops) and patas monkeys (Erythrocebus patas) living in simple habitats

Food distribution is hypothesized to be important in determining the nature of female relationships within social groups of primates. When food limits female reproductive success, spatially clumped foods are expected to produce strong, linear dominance hierarchies within groups, whereas more spatially dispersed foods are expected to produce weaker or non-existent dominance hierarchies. The association between food distribution and competitive relationships presumably occurs because clumped foods are usurpable but dispersed foods are not. We examined the spatial distribution of food patches (trees) and patch size relative to feeding behavior and agonistic interactions in vervets and patas monkeys, two closely related and sympatric species that nonetheless differ in the strength of the female dominance hierarchy. Food patches of both patas monkeys and vervets were small in size and randomly distributed in Acacia drepanolobium habitat. In contrast, in A. xanthophloea woodland, the habitat type that was exclusively used by vervets, food patches were larger and more spatially clumped. These similarities and differences between and within species were correlated with similarities and differences in the strength and linearity of their dominance hierarchies. Patas monkeys and vervets in A. drepanolobium habitat had dominance hierarchies that were weakly defined because there were relatively few agonistic interactions between females. By contrast, in A. xanthophloea habitat, vervets had a stronger, linear dominance hierarchy characterized by a higher rate of agonistic interactions over food. The covariation of agonistic interactions with patch size is discussed in relation to depletion time, another characteristic that may covary with food distribution, and resource renewal rate, an important determinant of agonistic interactions in insectivorous birds, fishes, insects, and mammals. Received: 18 February 2000 / Revised: 5 September 2000 / Accepted: 26 September 2000