Behaviour and reproductive fitness of postdispersal in plateau pikas (<Emphasis Type="Italic">Ochotona curzoniae</Emphasis>) on the Tibetan Plateau

Dispersal, which is a crucial process in animal population and evolutionary ecology, is considered personality dependent. The fitness consequences of dispersal are important. However, they are difficult and rarely measured in wild populations. In this study, we investigated the behaviour and reproductive fitness in a plateau pika (Ochotona curzoniae) population for two continuous years through mark–recapture experiment, behavioural tests and microsatellite analysis of parentage. Our results determined that around 53.8% males and 16.7% females dispersed to 40.3 and 28.2 m away from their natal family, respectively. The aggression (attack frequency and duration) of philopatric individuals was significantly higher than that of dispersed ones, although significant differences in boldness did not exist between them. For both males and females, the offspring number of philopatric pikas was significantly higher than that of dispersed ones. This result indicated that inbreeding may occur in plateau pikas. Our findings highlighted the importance of integrating behavioural, ecological and genetic analyses to explore the mechanism of dispersal. The findings also suggested that personality may play an important role in life history.     

Genetic diversity and sex‐bias dispersal of plateau pika in Tibetan plateau

Dispersal is an important aspect in organism's life history which could influence the rate and outcome of evolution of organism. Plateau pika is the keystone species in community of grasslands in Tibetan Plateau. In this study, we combine genetic and field data to character the population genetic pattern and dispersal dynamics in plateau pika (Ochotona curzoniae). Totally, 1,352 individual samples were collected, and 10 microsatellite loci were analyzed. Results revealed that plateau pika possessed high genetic diversity and inbreeding coefficient in a fine‐scale population. Dispersal distance is short and restricted in about 20 m. An effective sex‐biased dispersal strategy is employed by plateau pika: males disperse in breeding period for mating while females do it after reproduction for offspring and resource. Inbreeding avoiding was shown as the common driving force of dispersal, together with the other two factors, environment and resource. In addition, natal dispersal is female biased. More detailed genetic analyzes are needed to confirm the role of inbreeding avoidance and resource competition as ultimate cause of dispersal patterns in plateau pika.     

Dispersal, philopatry, and genetic relatedness in a social carnivore: comparing males and females

A balance must be maintained between the proportion of individuals dispersing and the proportion remaining philopatric such that inbreeding and resource competition are minimized. Yet the relative importance of dispersal and philopatric behaviour is uncertain, especially for species with complex social systems. We examine the influence of dispersal on genetic relationships of a white-nosed coati ( Nasua narica : Procyonidae) population from Panama. Field studies of the coati indicate a social system in which all females are highly philopatric and live in bands while all adult males become solitary at maturity, but do not disperse from the home range of their natal band. Based on analyses of multilocus DNA fingerprints, we confirm that female philopatry is the rule, long-distance dispersal is rare, and that relatedness between most bands is low. However, some new bands result from fission events and these bands retain relatively high relatedness to one another for several years. Adult males inhabiting the home range of a band are closely related to band members. In contrast, males and band members whose ranges do not overlap are unrelated or only slightly related. Adult males are also more closely related to other males whose home ranges they overlap extensively than to males whose home ranges they overlap only slightly. These results indicate that males initially disperse from their natal bands to reduce resource competition and not to avoid inbreeding. Inbreeding avoidance, if it occurs, results from more extensive range movements by males during the mating season.     

高原鼠兔间断性移动模式与反捕食对策分析

本文采用直接观测法, 测定了高原鼠兔地面移动频率及每次移动距离。结果表明, 高原鼠兔地面移动频率和每次移动距离依繁殖时期、年龄及性别而存在极显著的差异。繁殖早期, 成年雄体地面活动频率大于成年雌体。繁殖后期, 第1 胎雄性幼体大于同年龄的雌体。繁殖早期, 成年雌体地面活动频率高于繁殖后期, 而幼体与成体之间无显著的差异。繁殖初期雄体每次移动距离大于雌体, 其它时期雌、雄体之间则无明显的差别。雄体每次移动距离逐月降低。雌体每次移动距离无显著的季节性差异。研究结果进一步验证了暴露于风险环境中的植食性小哺乳动物, 主要采取间断性移动模式, 增加反捕食时间投入, 降低风险的假设。     

食物及洞道密度对高原鼠兔冬季死亡率的影响

本研究在野外围栏条件下采用2×2 析因实验设计,通过最小存活数法分别统计冬季前后种群数量,测定附加食物及洞道密度对高原鼠兔冬季死亡率的影响,旨在检验栖息地质量对降低高原鼠兔冬季死亡率的效应。4种野外实验处理2 个重复的研究结果表明,夏季无附加食物与低密度洞道处理组高原鼠兔冬季死亡率为71% ;而附加食物和高密度洞道处理冬季死亡率仅为47% ;添加食物和低密度洞道及无附加食物和高密度洞道处理组种群死亡率分别为60% 和53% 。每一处理组雌性死亡率均显著低于雄性。夏季添加食物和增加洞道密度均能明显地降低高原鼠兔种群冬季死亡率,但无显著的交互作用。     

Philopatry and within-colony movements in Columbian ground squirrels

Philopatry and dispersal result in selection of habitat locations that may differ in resources and social environment and thus should influence fitness components like survival and reproduction. We examined short-distance movements of young and adult females from natal or previous nesting sites within a colony of Columbian ground squirrels (Urocitellus columbianus) in the Rocky Mountains of Alberta, Canada, over a 17-year period. Females of all ages were strongly philopatric, yet a few (10-15%) exhibited movements that took them to new home ranges. We tested three hypotheses to explain the pattern of female natal and breeding movements: (1) that movements of philopatric females promote proximity to close kin; (2) that range shifts favour close kin via bequeathal of territory and (3) that dispersers move to lower density areas where competition for resources is lower. Tests of these three hypotheses revealed that: (1) philopatry and movements of young and older philopatric females led to proximity to mothers and local presence of close kin; (2) breeding dispersal did not result in bequeathal of home range to daughters, but movements of philopatric females suggested that they shared space with close kin and (3) adult females moved to new ranges with lower local densities, though dispersing females also left ranges where local density was significantly lower than for philopatric females. Natal and breeding movements among years produced two opportunities for territorial females: close spatial proximity to close kin via short philopatric movements, and habitats with fewer competitors via longer dispersal movements.     

The mating system and gene dynamics of plateau pikas

Evolutionary theory suggests that mating systems should have substantial effects on gene dynamics of local populations. In polygynous species, local 'breeding groups' may produce significant genetic structure, due to genetic differences among groups, and rate of loss of genetic variation from such populations may be considerably slowed. We examined possible influences of the variable mating system and family group structure on genetic properties of a population of plateau pikas (Ochotona curzoniae). Pika gene dynamics were examined via F-statistics and effective population sizes (N(e)), calculated from genetic correlations within and among individuals and families. Genetic correlations were estimated from mating patterns, population demography, and dispersal patterns. Substantial genetic structure within the population was indicated by a strongly positive F(LS). Genetic influence of natal dispersal out of pika families was indicated by a strongly negative inbreeding statistic (F(IL)=-0.34). Effective size of the population was not greatly different from the census population, whereas a traditional estimate of effective size of the population was much lower, indicating that the family structure of the pikas results in a slowed loss of genetic variation over time. Thus, even though mating patterns of plateau pikas were variable, family structure had a strong influence on pika gene dynamics.     

高原鼠兔血清无机元素含量的分析

测定分析了高原鼠兔血清电解质、无机元素、血清铁和总铁结合力, 并与人和其它实验动物的相应指标进行了比较。得出以下主要结论: 1.高原鼠兔血清电解质浓度不受生活环境和食物条件的影响, 其机体能自身调节血清离子浓度, 以维持电解质在血液中的相对衡定; 2.高原鼠兔血清铁含量与其栖息地海拔相关, 随海拔升高血清铁含量增加; 3.高原鼠兔通过增加运铁蛋白与铁的结合力作为适应高原低氧环境的方式之一。     

草地退化影响高原鼠兔血清总IgG水平、肠道寄生物感染及肝脏和肾脏指数

高原鼠兔(Ochotona curzoniae)作为青藏高原高寒草地生态系统中的关键物种,其种群数量随着过度放牧导致的草地退化而暴发式增长。阐明此过程中高原鼠兔天然免疫能力及肝脏和肾脏器官指数的变化,对深入揭示青藏高原草地退化引起的高原鼠兔种群暴发机理具有重要意义。本研究在青海海北高寒草地生态系统国家野外科学观测研究站地区,调查了轻度、中度、重度退化草地中高原鼠兔洞口数量,并测定了成体高原鼠兔血清总IgG含量、肠道寄生物感染状况及肝脏和肾脏指数。结果发现：随着草地退化,高原鼠兔洞口数量和血清总IgG水平显著增加;绦虫和线虫的感染率和感染强度以及肝脏和肾脏指数均显著降低。雄性对绦虫的感染率和肝脏指数显著低于雌性;雌性在重度退化草地中对绦虫和线虫的感染率及感染强度显著低于轻度退化草地。以上结果表明,过度放牧介导的草地退化增强了高原鼠兔的天然免疫功能,缓解了毒素对肝脏和肾脏的损伤,进而可能促进其种群暴发式增长。     

达乌尔鼠兔扩散过程中的生境选择

通过调控牧压,研究了达乌尔鼠兔扩散过程中的生境选择。结果表明,放牧活动强了草原植被的斑块性,改变了植被为鼠兔提供的资源条件,导致鼠兔斑块间的扩散和扩散鼠兔对不同植被斑块的生境选择。     

Dispersal Patterns Among Olive Colobus in Taï National Park

In Primates, females are more likely to be philopatric than males. However, in some species like Procolobus verus, females or individuals of both sexes disperse. In Taï National Park, Ivory Coast, olive colobus groups are small, with one or two adult males and 6 females. Dispersal is common for juveniles and adults of both sexes. Adult male dispersal is less common than adult female dispersal. Adult females immigrated especially into small, one-male groups indicating that food competition played a role. Furthermore, unknown sexually receptive females visited resident groups and mated with the resident males for a few days before disappearing again. Adult males dispersed when this improved their mating opportunities. All juveniles left their natal groups. The dispersal of juveniles may be a strategy to prevent inbreeding with their parents. Dispersal by juvenile males furthermore seemed to be the result of mate competition. The high dispersal rates, visits by receptive females, and dispersal of all individuals in the population suggest that moving between groups is a strategy that can be used ad hoc in several situations more easily in the olive colobus than in most other primates. The predation risks related to moving between groups were reduced by dispersing in conspecific or allospecific groups and by dispersing to neighboring groups.     

Social organization and space use in California voles: seasonal,sexual, and age-specific strategies

We intensively monitored space use and movement in Microtus californicus over a 2-year period that included 1 year of high density (maximum 618/ha) and one of low (minimum 5/ha); historically this population has exhibited cycles of 2 or 4 years. Adults of both sexes dispersed at the start of the breeding season, culminating in the establishment of intrasexually exclusive territories. In females, these territories persisted throughout life, except that many young females recruiting during the breeding season established contiguous, overlapping, or adjacent home ranges with their mothers. This female philopatry explains the conclusion of previous workers that females of this species are non-territorial. In the dry (non-breeding) season, females had smaller ranges that often overlapped and were clustered. Adult males moved breeding territories at a modal interval of 6 weeks; this is consistent with their avoidance of inbreeding with philopatric daughters. Ranges overlapped 1–4 adult females at any one time, and a cohort of 7 long-lived males overlapped an average of 16.4 females during their tenure on the grid. The period of maximum overlap with adult females varied among individual males, and did not correlate with the time of maximum body weight. Ranges of males in the dry season overlapped extensively, with persistent associations among some individuals. In the lowdensity year, ranges of some adults failed to overlap intersexually. Juvenile males dispersed gradually between 3 and 13 weeks of age (half before 9 weeks), with some leaving after reaching sexual maturity; a few remained philopatric. Of juvenile females, 47% remained philopatric with the rest disappearing before 9 weeks of age. New understanding of vole social behavior, dispersal, and space use is achieved by focusing on the seasonal dynamics of spatial relationships among individuals with respect to age, sex, and relatedness.     

Natal Dispersal and Philopatry in the Carnivorous Marsupial Phascogale tapoatafa (Dasyuridae)

The proximate and ultimate causes of dispersal in semelparous carnivorous marsupials (Phascogalinae) have previously been hypothesized to be maternal aggression and inbreeding avoidance, respectively. This study tests these hypotheses by exposing 26 litters of Phascogale tapoatafa to a diverse range of social and environmental conditions that potentially affect dispersal (e.g. supplemental feeding, post-weaning desertion by the mother, orphaning, and release of subadults into unoccupied habitat). The mean dispersal age was 162 ± 5.6 d, which is about 3 wk after weaning is complete. Juvenile dispersal was strongly male biased under all conditions, suggesting that extrinsic proximate causes do not adequately account for male emigration. Home range establishment by males was contingent on the presence of females. Half of the monitored daughters were philopatric, and others typically settled adjacent to the natal site, thus possibly enhancing their reproductive potential by occupying an area of known resource quality. Because philopatry increases the risk of incest, females may be selected to preferentially mate with unrelated males (immigrants), when they are available, to avoid inbreeding. If so, the presence of immigrant males would reduce the probability of locally born, related males reproducing at their natal site. Thus inbreeding avoidance by females may create local mate competition among males and select for male dispersal. Emigration also ensured that males avoided inbreeding, but, if they dispersed into unoccupied habitat, male P. tapoatafa often returned to the natal area. This 'boomerang strategy' of returning to mate with related females suggests that, in the absence of conspecifics along the dispersal path of a male, mate competition will be weak at the natal site and female mate choice will not preclude related males. Thus while inbreeding avoidance by either or both sexes is perhaps the most parsimonious explanation of male-biased emigration, dispersal patterns were apparently strongly influenced by additional factors, so that the ultimate causation of the dispersal regime may be more complex.     

Sexual segregation in winter, spring dispersal and habitat use in the pheasant (Phasianus colchicus)

Sexual segregation, spring dispersal and late winter to spring habitat selection were studied in the pheasant Phasianus colchicus L. on two areas for which the proportions of different habitats were similar. On one of the sites (Damerham) hand-reared pheasants were released for shooting. On the other (Fulwell) no releasing or shooting took place.
The sex ratio of birds feeding in fields in autumn favoured males because females fed in groups within woods. Groups of birds were significantly segregated by sex during early to late winter periods, but by late March they were not. Male groups and solitary males in winter were more evenly dispersed than female groups.
At Fulwell, female winter groups consisted of regular group members (mainly adults) and transitory individuals called 'drifters' (mainly immatures).
Females in their first breeding attempt dispersed further from their winter range to their breeding range than did adult females and females dispersed further than males. Males gaining a territory for the first time moved significantly further than old territory owners. Territorial males moved less than non-territorial males between successive spring ranges.
During March June, habitat use by 33 radio-tagged females changed markedly from mixed and deciduous woodland to fields sown with winter and spring corn. The implications for management are outlined.     

Determining natal dispersal patterns in a population of North American pikas (Ochotona princeps) using direct mark-resight and indirect genetic methods

A combination of direct (mark-resight) and indirect geneticmethods were used to investigate natal dispersal patterns andgenetic population structure in a population of North Americanpikas, Ochotona princeps. Pikas are small lagomorphs found intalus habitat of alpine areas throughout western North America.Adult pikas are individually territorial and rarely disperse.I used multilocus DNA fingerprinting to identify the parentsof juvenile animals. The settlement pattern of marked juvenilesand the pattern of relatedness of pikas across the study sitewas then examined within the study area. Although juvenilesborn at the study site exhibited a philopatric settlement pattern,an isolation-by-distance analysis did not reveal clusters ofhighly related individuals within the population. The F_ST estimatesuggests little genetic differentiation between populations2 km apart, and average DNA fingerprinting band-sharing amongadults was similar to values reported for outbred species. Anaverage of 34% of the adult population was replaced each winterby immigrants. DNA fingerprinting band-sharing analysis suggeststhat these immigrants had dispersed short, intermediate, andlong distances. These findings differ from earlier studies whichused observations of marked animals only to characterize dispersalpatterns. Direct observations of marked juveniles had documenteda philopatric settlement pattern, little or no dispersal outof natal populations, and no direct evidence of long distancemovement. Of the three major hypotheses proposed to explainthe evolution of dispersal in birds and mammals, competitionfor resources, competition for mates, and inbreeding avoidance,the results of this study support a competition for resourceshypothesis, where the key resource is territory     

Keystone status of plateau pikas (Ochotona curzoniae): effect of control on biodiversity of native birds

The plateau pika (Ochotona curzoniae) of the Qinghai–Xizang (Tibetan) plateau, People's Republic of China, has been considered a pest because it putatively competes with native livestock for forage and contributes to rangeland degradation. As a result the plateau pika has been poisoned across vast areas of the high alpine meadows of the plateau. The plateau pika has also been considered a keystone species for biodiversity on the plateau. As one test of the keystone species hypothesis, we investigated the effects of poisoning plateau pikas on avian species richness and abundance. We conducted standardized censuses of birds on a number of sites across the alpine grassland of Qinghai province on which pikas either had or had not been poisoned. Avian species richness and abundance were higher on non-poisoned sites, in particular for species that nest in pika burrows such as Hume's groundpecker (Pseudopodoces humilis) and six species of snowfinch (Montifringilla spp., Pyrgilauda spp.), and species that prey on pikas (upland buzzard, Buteo hemilasius; black-eared kite, Milvus lineatus). The plateau pika thus appears to be both an allogenic engineer and a keystone species. Poisoning pikas reduces biodiversity of native species on the Qinghai–Xizang plateau, therefore management decisions concerning plateau pikas should reflect caution and careful assessment.     

Sex-biased dispersal in eastern chipmunks, Tamias striatus

I investigated the effect of male mate competition and inbreeding avoidance on natal dispersal of chipmunks by longitudinally monitoring known individuals from 1986 to 1990. Natal males exhibited greater absolute and effective dispersal distances but dispersed at the same proportion as natal females. Recruitment of juvenile males was negatively affected by density of resident males, but there was no evidence of local mate competition among male kin. Analysis of the spatial distribution of neighbors showed that natal males settled farther from their mothers than did their female siblings and farther than unrelated juvenile males. In addition, mothers apparently tolerated daughters as close neighbors and occasionally shared den sites with grandprogeny. Sexually mature males were never neighbors of their mothers and were never observed at maternal mating bouts. Males may disperse to improve reproductive opportunities by avoiding competition with resident males, and by increasing access to unrelated females. Maternal tolerance of daughters but not sons may result in the close affiliation between mothers and daughters, and indirectly contribute to dispersal of natal males. Hence male-biased dispersal could be a consequence of mate competition and maternal avoidance of incestuous matings. This revised version was published online in July 2006 with corrections to the Cover Date.     

Behavioral mechanisms of male sterilization on plateau pika in the Qinghai-Tibet plateau

Fertility control is an alternative non-lethal method in the management of rodents. Previous modeling suggests that the efficacy of male sterilization depends on mating systems of animals, but behavioral mechanisms of male sterilization have not been investigated. Here we investigated the behavioral mechanism of the sterilant quinestrol in reducing the fertility of plateau pika (Ochotona curzoniae) inhabiting the Qinghai-Tibet plateau. Male pikas treated with quinestrol showed reduced aggression compared to control males, but they showed significantly higher levels of territorial behavior such as long-calls and long-chases. Levels of long-call and long-chase were negatively correlated with the number of newborn pikas in the family. Single-baiting of quinestrol effectively sterilized male pikas and reduced the pregnancy rate of female pikas; this was likely achieved by increased territorial behavior of sterilized pikas which resulted in unsuccessful invasions by fertile adult male pikas. Our study reveals a novel behavioral mechanism, increased territoriality in sterilized males, in the fertility control of plateau pikas.     

Strategic promiscuity helps avoid inbreeding at multiple levels in a cooperative breeder where both sexes are philopatric

In cooperative breeders, the tension between the opposing forces of kin selection and kin competition is at its most severe. Although philopatry facilitates kin selection, it also increases the risk of inbreeding. When dispersal is limited, extra-pair paternity might be an important mechanism to avoid inbreeding, but evidence for this is equivocal. The red-winged fairy-wren is part of a genus of cooperative breeders with extreme levels of promiscuity and male philopatry, but is unique in that females are also strongly philopatric. Here, we test the hypothesis that promiscuity is an important inbreeding avoidance mechanism when both sexes are philopatric. Levels of extra-pair paternity were substantial (70% of broods), but did not arise through females mating with their helpers, but via extra-group mating. Offspring were more likely to be sired by extra-pair males when the social pair was closely related, and these extra-pair males were genetically less similar to the female than the social male and thus, inbreeding is avoided through extra-pair mating. Females were consistent in their choice of the extra-pair sire over time and preferred early moulting males. Despite neighbouring males often being close kin, they sired 37% of extra-pair offspring. However, females that gained paternity from neighbours were typically less related to them than females that gained paternity further away. Our study is the first to suggest that mating with both closely related social partners and neighbours is avoided. Such sophistication in inbreeding avoidance strategies is remarkable, as the extreme levels of promiscuity imply that social context may provide little cue to relatedness.     

高寒草甸生态系统中高原鼠兔的繁殖特征

2002年4月至8月,在中国科学院海北高寒草甸生态系统定位站附近,采用标志重捕法对高原鼠免的繁殖特征进行了研究。高原鼠兔的繁殖具有明显的季节性差异,4月下旬和5月为繁殖高峰期,雄性的睾丸重而饱满,雌性的怀孕率最大。雄性成体的体重和睾丸重在繁殖期的不同时段具有显的差异,在4月11日至5月10日、6月11日至7月10日体重和睾丸重之间呈显正相关。雌性怀孕早期胚胎数与临产前胚胎数没有显差异,未发现胚胎吸收现象;同时,胚胎数在繁殖期的不同时段存在显差异。采用种群统计学中同生群的划分方法,将5～8月份出生的幼体依次记为L1、L2、L3、L4。在出生后20天内L1、L2幼体存活率明显高于L3、L4;在从出生50天至80天期间L4的存活率显高于L2。当年出生的雌雄幼体在发育上存在不同步现象,当年出生的雌性幼体性成熟早,有的可以直接参加繁殖,但当年出生的雄性幼体却无此现象。结果表明：在海北地区,经过综合治理后,高原鼠兔的生境发生改变,其繁殖策略也随之发生改变,即,减少每次繁殖活动的投入,增加繁殖次数,延长繁殖时间。