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.     

Male philopatry,extra-pack copulations and inbreeding avoidance in Ethiopian wolves (Canis simensis)

Monogamous pairings have been regarded as the fundamental social unit in all canid species, including those living in packs. In Ethiopian wolves, however, habitat saturation limits dispersal, which raises the question of whether they avoid inbreeding and, if so, by what mechanism. In two study areas Ethiopian wolf packs had stable memberships. Each pack comprised two to eight adult males, one to three adult females, including a clear-cut dominant individual of each sex, together with one to six yearlings and up to six pups (n = 9 packs). Males remained in their natal packs, apparently throughout their lives. Some females also failed to disperse while others dispersed in their second or third year and became floaters. Dominant females monopolized breeding, and were succeeded either by their most dominant daughters (three cases) or by floaters (two cases). In the former case there is potential for incest; however, 70% of 30 copulations observed were between the dominant female of one pack and a male from an adjoining pack. In Ethiopian wolves, under conditions where dispersal is constrained and the potential for inbreeding is high, extra-pack matings (and associated multiple paternity) result in outbreeding. We raise the possibility that extra-pair copulations may be widespread in canid societies and that the monogamy supposedly fundamental to the family may be more sociological than genetic. Received: 15 October 1994/Accepted after revision: 9 December 1995     

Microgeographic heterogeneity in spatial distribution and mtDNA variability of gray mouse lemurs (<Emphasis Type="Italic">Microcebus murinus</Emphasis>, Primates: Cheirogaleidae)

The objective of our study was to investigate the spatial distribution and genetic structure of a solitary primate at the microgeographical scale of adjacent local populations. We obtained spatial data and tissue samples for mtDNA analysis from 205 gray mouse lemurs (Microcebus murinus) captured along transects and within 3 grid systems within a 12.3 km² area in Kirindy Forest, western Madagascar. Our capture data revealed that, even though the forest was continuous, gray mouse lemurs were not evenly distributed, and that daily and maximum dispersal distances were significantly greater in males. The frequency distribution of 22 mtDNA D-loop haplotypes was highly skewed. Nine haplotypes were unique to males, indicating male-mediated gene flow from surrounding areas. The geographic distribution of haplotypes revealed that males were also more dispersed than females. Females with the same haplotype showed a tendency towards spatial aggregation, and the correlation between genetic and geographic distances was higher in females. In several areas of the forest, however, spatially clustered females were not of the same haplotype, and females were not always found in clusters. Hence, in contrast to suggestions from previous studies, matrilineal clustering is not the only way females are socially organized. In addition, our study revealed heterogeneity and patterns in population structure that were not evident at smaller spatial scales, some of which may be relevant for designing conservation strategies.Communicated by C. Nunn     

A dynamic model of human dispersal in a land-based economy

This paper presents a dynamic state variable model that examines human dispersal in a land-based economy. In humans, like other animals, many individuals disperse during their lifetimes, with extensive variability in the timing and likelihood of dispersal by individuals of different states. The model places human and non-human dispersal in a common framework, specifically by utilizing a cost-benefit approach. As with other animals, individual access to resources likely plays an important role in shaping human dispersal, and the model explores the mechanisms underlying this relationship. Over a series of time steps, individuals must decide whether to stay in their natal area or disperse, given their states (wealth, marital, and inheritance status) and the environmental constraints on dispersal. Costs of dispersal include time and an initial drop in wealth, while the benefit is the potential of the dispersal area to increase wealth. The model determines whether dispersing or staying results in the highest fitness, where fitness is a function of individual wealth and years married. The model shows that dispersal is favorable under a wide range of environmental conditions, but that it also varies predictably according to individual states. Men of middle wealth values disperse under a wider variety of circumstances than poorer or wealthier men. Wealthy men, who are particularly sensitive to the time cost and mortality risk associated with dispersing, appear to adopt a more conservative dispersal strategy than poorer men, who are more sensitive to the wealth cost and wealth growth rate associated with dispersing. Dispersal behavior is also contingent on its effects on an individual's probability of marrying or inheriting wealth. Finally, the model fosters an examination of how the interaction of life events, and their directions of causation, can be studied. Received: 6 May 1998 / Received in revised form: 3 February 1999 / Accepted: 7 February 1999     

Genetic population structure and sociogenetic organisation in Formica truncorum (Hymenoptera; Formicidae)

Summary The genetic population structure and the sociogenetic organization of the red wood ant Formica truncorum were compared in two populations with monogynous colonies and two populations with polygynous colonies. The genetic population structure was analysed by measuring allele frequency differences among local subsets of the main study populations. The analysis of sociogenetic organisation included estimates of nestmate queen and nestmate worker relatedness, effective number of queens, effective number of matings per queen, relatedness among male mates of nestmate queens and relatedness between queens and their male mates. The monogynous populations showed no differentiation between subpopulations, whereas there were significant allele frequency differences among the subpopulations in the polygynous population. Workers, queens and males showed the same genetical population structure. The relatedness among nestmate workers and among nestmate queens was identical in the polygynous societies. In three of the four populations there was a significant heterozygote excess among queens. The queens were related to their male mates in the polygynous population analysed, but not in the monogynous ones. The data suggest limited dispersal and partial intranidal mating in the populations with polygynous colonies and outbreeding in the populations having monogynous colonies. Polyandry was common in both population types; about 50% of the females had mated at least twice. The males contributed unequally to the progeny, one male fathering on average 75% of the offspring with double mating and 45–80% with three or more matings. Correspondence to: L. Sundström     

Potential correlation between perfluorinated acids and liver morphology in East Greenland polar bears (Ursus maritimus)

In this study, residual concentrations of chlorpyrifos (CPF) were determined in feed (40) and fodder (25) samples collected from various locations of Tarai region of Uttarakhand. For extracting residues, liquid–liquid partition followed by alumina column clean up was used and the detection and quantification of residues was undertaken with the help of high-performance liquid chromatography using C18 column and diode array detector at 220?nm. Of the total 40 feed samples analyzed, 7 (17.5%) samples were found positive for CPF with the mean residual concentration of 0.058?µg?g^?1; while out of 25 fodder samples, CPF residues were detected in a single (4%) sample with residual concentration of 0.39?µg?g^?1. However, none of the feed or fodder samples contained CPF residues above the prescribed limit.     

Factors influencing length of maternal care in brown bears (<Emphasis Type="Italic">Ursus arctos</Emphasis>) and its effect on offspring

Length of maternal care, i.e. the interval between successfully raised litters, is the most important factor explaining the variation in reproductive rate among brown-bear (Ursus arctos) populations. In this paper, we examine the variation in length of maternal care in radio-marked brown bears and its effect on their offspring in northern Sweden. Young stayed with their mothers for 1.4–1.5 or 2.4–2.5 (in one case 3.5) years and were weaned with body masses varying from 17 to 69 kg. The probability of yearling litters staying with their mother for a 2nd year increased with decreasing yearling body mass, and was higher for litters with two offspring than for litters with one or three to four offspring. Staying with their mothers for a 2nd year had a positive effect on mass gain in yearlings and this effect was more pronounced in litters with two than three to four offspring. Body mass of 2-year-olds was not related to age of weaning, suggesting that keeping offspring for an additional year mainly compensated for low yearling body mass. If large offspring body mass positively affects later offspring survival and reproduction, mothers may be able to optimize the length of maternal care according to the litter size and the size of their yearlings.Communicated by F. Trillmich     

Genetic structure,lack of sex-biased dispersal and behavioral flexibility in the pair-living fat-tailed dwarf lemur, <Emphasis Type="Italic">Cheirogaleus medius</Emphasis>

Mating system and dispersal patterns influence the spatio-genetic structure within and between populations. Among mammals, monogamy is rare, and its socio-genetic consequences have not been studied in detail before. The goal of our study was to investigate population history, demographic structure, and dispersal patterns in a population of pair-living fat-tailed dwarf lemurs, Cheirogaleus medius, a small, nocturnal primate from western Madagascar, and to infer their underlying behavioral mechanisms. Tissue samples for DNA extraction were obtained from a total of 140 individuals that were captured in two subpopulations about 3 km apart. Analyses of mtDNA variability at the population level revealed very low levels of genetic variability combined with high haplotype diversity, which is indicative of a recent population bottleneck. We found no evidence for spatial clustering of same-sexed individuals with identical haplotypes within each of two subpopulations but significant clustering between them. Thus, a high level of local subpopulation differentiation was observed (F _ST = 0.230). The sexes showed equal variances in the number of individuals representing each haplotype, as well as equal levels of aggregation of identical haplotypes. Hence, both sexes disperse from their natal area, one pattern expected in a pair-living mammal. There is a possibility of behavioral and social flexibility in this species, however, because we documented pronounced differences in density and sex ratio between the two subpopulations, suggesting that single study sites or populations may not be representative of a given local population or even species.     

Ecology of<Emphasis Type="Italic"> Leptothorax</Emphasis> ants: impact of food,nest sites,and social parasites

In a long-term field manipulation, we demonstrate strong reactions of Leptothorax longispinosus ant colonies to food- and nest-site supplementation. Demographic and genetic responses varied over small geographic scales, and the two ecological factors interacted with the presence of the social parasite Protomognathus americanus. We conducted a 2×2 experiment in three blocks and found that the blocks, which were less than 100 m apart, reacted very differently to the treatments. Blocks differed in degree of polygyny, intranest relatedness, colony size, productivity, and sexual investment. Furthermore, these differences were associated with the presence of slave-making ants and the local availability of nest sites. Nest-site supplementation had a strong effect only in the site with the highest prevalence of social parasites, influencing there the density and investment patterns of colonies. L. longispinosus ants in the least parasitized area were strongly affected by both food- and nest-site supplementation. There, food supplementation led to a decrease in the number of queens per colony and consequently to an increase in intranest relatedness, while colonies in nest-site-supplemented areas invested fewer resources in males and produced a female-biased allocation ratio. By contrast, in a third block with a very low intracolonial relatedness, food supplementation induced an absolute and relative higher investment in males. We conclude that ecological factors influencing social organization in insect societies cannot be studied in isolation, because the interactions among factors produce far richer responses than any one variable.Communicated by L. Sundström     

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     

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     

三江平原典型小叶章种群地上生物量的空间结构分形特征

2005年5月至10月,应用分形几何学的原理和方法对三江平原典型小叶章种群地上生物量与株高的空间结构分形关系进行了研究。结果表明,典型草甸小叶章和沼泽化草甸小叶章的地上生物量与株高存在良好的静态分形关系,其分形维数(D)是对地上生物量在各器官中积累规律的表征;二者地上生物量与株高的对数值线性相关,相关系数分别介于0.579～0.919和0.655～0.914之间(P<0.01,D值介于1.778～3.414和2.238～3.924之间,各时期地上生物量均以株高的幂函数形式积累;二者地上生物量与株高静态分形关系D值的季节动态基本一致,其变化均符合抛物线模型(P<0.05),而局部波动又与地上生物量的变化有关;生长季内,二者地上生物量与株高动态分形关系的D值分别为2.749和2.738,说明地上生物量的积累具有自相似性,符合幂函数增长的分形生长过程。研究还表明,仅用D值来刻画不同植物地上生物量的空间积累与展布存在一定的局限性,它只能用于测度和描述一种植物体生物量的空间积累及其对空间的占据,而一般不能用于植物间相应指标的类比;导致不同植物D值差异的原因可能与植物生长所处生境、外在因素以及模型参数(C)有关。     

Extended family structure in the ant Formica paralugubris: the role of the breeding system

In populations of various ant species, many queens reproduce in the same nest (polygyny), and colony boundaries appear to be absent with individuals able to move freely between nests (unicoloniality). Such societies depart strongly from a simple family structure and pose a potential challenge to kin selection theory, because high queen number coupled with unrestricted gene flow among nests should result in levels of relatedness among nestmates close to zero. This study investigated the breeding system and genetic structure of a highly polygynous and largely unicolonial population of the wood ant Formica paralugubris. A microsatellite analysis revealed that nestmate workers, reproductive queens and reproductive males (the queens' mates) are all equally related to each other, with relatedness estimates centring around 0.14. This suggests that most of the queens and males reproducing in the study population had mated within or close to their natal nest, and that the queens did not disperse far after mating. We developed a theoretical model to investigate how the breeding system affects the relatedness structure of polygynous colonies. By combining the model and our empirical data, it was estimated that about 99.8% of the reproducing queens and males originated from within the nest, or from a nearby nest. This high rate of local mating and the rarity of long-distance dispersal maintain significant relatedness among nestmates, and contrast with the common view that unicoloniality is coupled with unrestricted gene flow among nests. Received: 8 February 1999 / Received in revised form: 15 June 1999 / Accepted: 19 June 1999     

Do dominants have higher heterozygosity? Social status and genetic variation in brown trout, Salmo trutta

A key question of evolutionary importance is what factors influence who becomes dominant. Individual genetic variation has been found to be associated with several fitness traits, including behaviour. Could it also be a factor influencing social dominance? We investigated the association between social status and the amount of intra-individual genetic variation in juvenile brown trout (Salmo trutta). Genetic variation was estimated using 12 microsatellite loci. Dominant individuals had higher mean heterozygosity than subordinates in populations with the longest hatchery background. Heterozygosity–heterozygosity correlations did not find any evidence of inbreeding; however, single-locus analysis revealed four loci that each individually differed significantly between dominant and subordinate fish, thus giving more support to local than general effect as the mechanism behind the observed association between genetic diversity and a fitness-associated trait. We did not find any significant relation between mean d ² and social status, or internal relatedness and social status. Our results suggest that individual genetic variation can influence dominance relations, but manifestation of this phenomenon may depend on the genetic background of the population.     

Emigration mechanisms in feral house mice – a laboratory investigation of the influence of social structure,population density,and aggression

Emigration in small mammals may be strongly related to social factors, but direct observations of emigrants are rare. Feral house mice (Mus domesticus) were studied using a population cage system that allowed continuous observation of individually marked animals. Mice that left their natal cage and took up residence in cages that could only be reached by crossing a water barrier were defined as emigrants. Six pairs of house mice with their litters were placed in the system, and data on aggressive interactions, body weight, reproduction, mortality and emigration were collected daily. Both sexes emigrated, but males did so twice as often as females. Population density was not correlated with the frequency of aggression, and had no effect on the weight of emigrating individuals. Male emigrants suffered more aggression before emigration than their non-emigrant brothers of the same age; they were aggressively driven out by other males, predominantly by the father. Female emigration depended on the female’s chances of reproduction. The probability of a female reproducing decreased with increasing birth order. Females born in a late litter, who therefore had only a low chance of reproduction, dispersed earlier than those of early litters. Resident males were reproductively suppressed. Male offspring had two different strategies for attaining top rank. They could develop rapidly and reach sexual maturity early on, but face competition with the father, risking being forced to emigrate. Alternatively, they could develop slowly, stay within their family and wait for a chance to take over the dominant position. It is concluded that emigration in male and female feral house mice is caused by intrasexual competition. Received: 13 July 1995/Accepted after revision: 8 June 1996     

Application of a spatial meta-population model with stochastic parameters to the management of the invasive grass Nassella trichotoma in North Canterbury, New Zealand

Optimising the management of invasive plants requires the identification of the population size outcomes for alternative management strategies. Mathematical models can be useful tools for making such management strategy comparisons. In this paper we develop a generic landscape meta-population model and apply it to the weedy grass, Nassella trichotoma, an invasive species occupying approximately 800 land parcels, predominantly pastoral farms, in the Hurunui district, North Canterbury, New Zealand. Empirical evidence reveals that this meta-population is currently stable (at a median density of 6 plants ha⁻¹) under a community strategy requiring manual removal (termed ‘grubbing’) of plants annually from all land parcels. Reduction in population size requires an alternative management strategy. Field data, collected over a 12 year period, were used to provide stochastic parameter values for land parcel size, carrying capacity, rates of local population growth and grubbing.The model reveals that at steady state, the most significant contribution to population growth on a land parcel comes from within the land parcel itself; the expected annual per capita growth on an individual land parcel is 4 orders of magnitude greater than the expected annual contribution from plants arising from other land parcels. This result implies that many of the farms currently supporting N. trichotoma may pose little or no threat to, nor are threatened themselves by, other farms infested by the weed. However, the steady state distribution (of the weed's population density) was sensitive to the spread rate, revealing a need for data on this process. It was also sensitive to how any increase in the grubbing rate is distributed; increasing it via a uniform distribution U(0, 1) where all rates between 0 and 100% year⁻¹ are equally probable did not affect the steady state, whereas increasing the rates via the uniform distribution U(0.25, 0.75) resulted in fewer farms with high population densities. In general the model provides a basis for exploring the effects of a wide range of alternative grubbing strategies on population growth in N. trichotoma.     

The effect of positive interactions on community structure in a multi-species metacommunity model along an environmental gradient

Positive interactions are widely recognized as playing a major role in the organization of community structure and diversity. As such, recent theoretical and empirical works have revealed the significant contribution of positive interactions in shaping species’ geographical distributions, particularly in harsh abiotic conditions. In this report, we explore the joint influence of local dispersal and an environmental gradient on the spatial distribution, structure and function of communities containing positive interactions. While most previous theoretical efforts were limited to modelling the dynamics of single pairs of associated species being mutualist or competitor, here we employ a spatially explicit multi-species metacommunity model covering a rich range of interspecific interactions (mutualism, competition and exploitation) along an environmental gradient. We find that mutualistic interactions dominate in communities with low diversity characterized by limited species dispersal and poor habitat quality. On the other hand, the fraction of mutualistic interactions decreases at the expense of exploitation and competition with the increase in diversity caused by higher dispersal and/or habitat quality. Our multi-species model exemplifies the ubiquitous presence of mutualistic interactions and the role of mutualistic species as facilitators for the further establishment of species during ecosystem assembly. We therefore argue that mutualism is an essential component driving the origination of complex and diverse communities.     

Experimental manipulation of colony genetic diversity had no effect on short-term task efficiency in the Argentine ant Linepithema humile

Genetic diversity might increase the performance of social groups by improving task efficiency or disease resistance, but direct experimental tests of these hypotheses are rare. We manipulated the level of genetic diversity in colonies of the Argentine ant Linepithema humile, and then recorded the short-term task efficiency of these experimental colonies. The efficiency of low and high genetic diversity colonies did not differ significantly for any of the following tasks: exploring a new territory, foraging, moving to a new nest site, or removing corpses. The tests were powerful enough to detect large effects, but may have failed to detect small differences. Indeed, observed effect sizes were generally small, except for the time to create a trail during nest emigration. In addition, genetic diversity had no statistically significant impact on the number of workers, males and females produced by the colony, but these tests had low power. Higher genetic diversity also did not result in lower variance in task efficiency and productivity. In contrast to genetic diversity, colony size was positively correlated with the efficiency at performing most tasks and with colony productivity. Altogether, these results suggest that genetic diversity does not strongly improve short-term task efficiency in L. humile, but that worker number is a key factor determining the success of this invasive species.Communicated by L. Sundström     

A genetic component to size in queens of the ant,<Emphasis Type="Italic"> Formica truncorum</Emphasis>

The genetic basis of morphological traits in social insects remains largely unexplored. This is even true for individual body size, a key life-history trait. In the social insects, the size of reproductive individuals affects dispersal decisions, so that small size in queens is often associated with reduced dispersal, and large size with long-range dispersal and independent colony founding. Worker size is connected to division of labour when workers specialize in certain tasks according to their size. In many species, variation in worker size has been shown to increase colony performance. In this study, we present the first evidence of an additive genetic component to queen size in ants, using maternal half sib analysis. We also compared intra-colony size variation in colonies with high (queen doubly mated) versus low (queen singly mated) genetic variability. We found a high and significant heritability (h²=0.51) for queen size in one of the two study years, but not in the other. Size variation among queens was greater in colonies headed by a doubly mated queen in one of the study years, but not in the other. This indicates that genetic factors can influence queen size, but that environmental factors may override these under some circumstances. The heritability for worker size was low (h²=0.09) and non-significant. Increased genetic diversity did not increase worker size variation in the colonies. Worker size appeared largely environmentally determined, potentially allowing colonies to adjust worker size ratios to current conditions.Communicated by J. Heinze