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1.
We examine the evolutionary stability of year-round residency in territorial populations, where breeding sites are a limiting resource. The model links individual life histories to the population-wide competition for territories and includes spatial variation in habitat quality as well as a potential parent-offspring conflict over territory ownership. The general form of the model makes it applicable to the evolution of dispersal, migration, partial migration, and delayed dispersal (offspring retention). We show that migration can be evolutionarily stable only if year-round residency in a given area would produce a sink population, where mortality exceeds reproduction. If this applies to a fraction of the breeding habitat only, partial migration is expected to evolve. In the context of delayed dispersal, habitat saturation has been argued to form an ecological constraint on independent breeding, which favors offspring retention and cooperative breeding. We show that habitat saturation must be considered as a dynamic outcome of birth, death, and dispersal rates in the population, rather than an externally determined constraint. Although delayed dispersal often associates with intense competition for territories, life-history traits have direct effects on stable dispersal strategies, which can often override the effect of habitat saturation. As an example, high survival of floaters selects against delayed dispersal, even though it increases the number of competitors for each breeding vacancy (the "habitat saturation factor"). High survival of territory owners, by contrast, generally favors natal philopatry. We also conclude that spatial variation in habitat quality only rarely selects for delayed dispersal. Within a population, however, offspring retention is more likely in high-quality territories.  相似文献   

2.
American dipper Cinclus mexicanus populations are frequently composed of resident individuals that occupy permanent territories year round and migratory individuals that overwinter with residents but migrate to breeding territories on higher elevation creeks each spring. Between 1999 and 2004 we examined how migratory strategy (resident/migratory) and sex differences influence breeding territory fidelity of American dippers occupying the Chilliwack River watershed, British Columbia, Canada. Counter to expectation we found that the migratory strategy of American dippers did not influence whether birds breeding in one year were found on their former breeding territory in the next. Migratory strategy also did not affect the probability that known surviving dippers occupied the same breeding territory in the following year. Males and females were equally likely to be found on their former territory in the following year (females 43%, males 41%) and known survivors had similar levels of breeding territory fidelity (females 74%, males 68%). However, breeding territory fidelity of males and females varied in response to different factors. Surviving female dippers were more likely to be found on their former breeding territory in the subsequent year following a successful breeding attempt than an unsuccessful breeding attempt. Prior reproductive performance did not influence whether surviving male dippers were found on their former breeding territory. Male dippers were more likely to be found on their former territory and, if they survived, have higher breeding territory fidelity when their mate also returned to that same territory. Mate retention also influenced whether females were found on their former territory in the following year but had no effect on the breeding territory fidelity of known survivors. We argue that sex‐specific dispersal decision rules in American dippers are driven by sex differences in the predictability of breeding performance between years and sex differences in how mate retention influences subsequent reproductive success.  相似文献   

3.
In species in which males defend territories for breeding, males may differ in territorial behavior; alternative behaviors among territorial males are not well understood. In our long‐term study of partially‐migratory song sparrows, we have observed that most territorial males establish territories before females begin nesting and remain site‐faithful both within and between breeding seasons; however, some males establish territories later in the season (late establishers) and/or change territory locations either within or between seasons (movers). Whether late establishment or moving are equally successful strategies for territory defense, or best‐of‐bad‐job options, is not known. Here, we compare the frequencies of these behaviors to demographic variables over a 9‐yr period and compare lifetime tenure and early season nesting success for males who differ in site fidelity and timing of territory establishment. Across years, late establishing was negatively correlated with the return rate of previously territorial males; moving was positively correlated with the number of occupied territories at the start of the breeding season (territory density). While moving was independent of number of years on territory, late territory establishment only occurred in a male’s first year as a territory holder. Of 88 males, 25% established their first territory late, primarily in undefended space; 31% moved. Late and early establishers did not differ in lifetime tenure; movers, however, had longer lifetime tenure than site‐faithful males. Among early establishers, movers and non‐movers did not differ in the number of successful early nests/year or number of young fledged/year; among late establishers, however, movers had significantly higher early nesting success by both measures. Late establishers who moved had higher early season nesting success and higher early season nesting success/year than site‐faithful early establishers. Thus, individual variation in the timing of territory establishment and site fidelity may be facultative alternative territorial strategies.  相似文献   

4.
Many territorial species have a mating system characterized by males establishing home ranges in the breeding grounds prior to females, resulting in males competing for territories and females choosing a mate upon their arrival. It remains unknown, however, how the outcomes of decisions surrounding territory establishment and mate choice are influenced by the spatial configuration of the breeding grounds. We use a spatially explicit, individual-based model to investigate the sex-specific effects of these decisions on reproductive success. In our model, males that arrive earlier obtain higher quality territories and improve their chances for extra-pair copulations. Females can choose their mate to maximize the quality of the male or to attempt to minimize the density of other females near their nesting site to avoid competition. Females therefore face a tradeoff between high-density regions around high-quality males and low-quality males in areas of low competition. Our model predicts a negative correlation between male and female reproductive success under a wide range of conditions when the majority of the territories are on the margins of the breeding area. Most notably, this sexual conflict arises as an edge effect suggesting that fragmentation of breeding habitats could impact the consequences of mate choice in many species with territorial breeding habits.  相似文献   

5.
The population dynamics of tundra-nesting Peregrine Falcons Falco peregrinus tundrius was studied over 7 years on a 450 km2 study area in the Keewatin District of Canada's Northwest Territories. Peregrines showed strong fidelity to nest sites; none of 25 males ringed changed territories, while five of 38 females ringed were recorded changing territories. Such changes usually occurred after nesting failure. Annual turnover of territorial adults was estimated to be 22% (15% for males and 26% for females). Annual mortality of adults was estimated to be 17% (15% for males and 19% for females). If we assumed that territory vacancies, in addition to mate replacements, were indicative of mortality, then maximum annual mortality was estimated at 24% for each sex. Territories were held only by adult Peregrines. The oldest male on territory was at least 7 years old, the youngest was 2. The oldest female on territory was at least 7 years old, the youngest was 3. Territories were held by individuals of each sex for at least 6 years. One pair remained together for at least 4 years. Less than 4% of all young Peregrines produced on the study area in the first 5 years of the study were recruited into the breeding population. More male than female young were recruited despite an even sex ratio among nestlings. Peregrines did poorly in their first breeding attempts. The single young female recruited into the study population dispersed more than three times the median dispersal distance of six recruited males, suggesting that other females probably dispersed beyond the boundaries of the study area.  相似文献   

6.
Breeding dispersal among territorial species is of interest to population biologists because leaving a territory carries fundamental risks to the dispersing individuals, and this may not outweigh the costs of maintaining the territory. Most studies of breeding dispersal have focused on species inhabiting spatially open populations, in which undetected emigration could impart a negative bias to estimates of dispersal. We studied breeding dispersal in an isolated (spatially closed) population of California Spotted Owl Strix occidentalis occidentalis in southern California for 12 years to assess factors that might correlate with breeding dispersal. Twenty‐nine per cent (n = 47) of territorial females and 19% (n = 35) of territorial males dispersed at least once during the study. Annually, 0–13% of the territorial females and 0–12% of the territorial males dispersed. Among a set of a priori and post hoc models related to breeding dispersal, the top a priori model indicated that birds having higher reproductive output relative to the population average were less likely to disperse. A post hoc model based on an index of territory quality was ranked higher than the top a priori model and indicated that birds occupying higher quality territories were less likely to disperse. These two models were correlated and represented short‐ and long‐term reproductive performance, respectively. Birds that dispersed also failed to fledge young in the year prior to dispersal, but the failure to fledge young did not, by itself, explain dispersal. Because Spotted Owls are long‐lived, they may ultimately improve their reproduction by dispersing given that they would have future opportunities to breed over the long term. Birds whose mates are likely to have died tended to improve their reproductive success, whereas the relationship between reproductive success of birds that divorced was less clear. Substantial variation in breeding dispersal was unexplained by our analysis so it is likely that the motivation to disperse was a complex process in this population.  相似文献   

7.
1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.  相似文献   

8.
R. David  Stone 《Journal of Zoology》1987,212(1):117-129
The social organization, spatial utilization and movement patterns of free-ranging desmans ( Galemys pyrenaicus G.) were studied by radiotracking individuals in the French Pyrenees. During the period of study, May to July, single adult male and female desmans formed a pair and defended their shared territory from neighbouring conspecifics by vigilance and scent-marking behaviour. Other adults did not form pair bonds and remained solitary. In such pairs, the territory of the adult male was always larger than, and completely enclosed, that of the paired female. Neighbouring pairs occupied contiguous territories of similar length. The territories of neighbouring males overlapped to a small extent, but the territory of one male and that of a neighbouring female were never seen to overlap. Juveniles were observed to utilize the territory of resident pairs, prior to their dispersal. Paired adults and juveniles exploited their ranges on a regular daily basis, whilst those of solitary adults, being larger than those of the former, were utilized on a 48–hour basis. Paired males appeared to invest more time in defending the borders of their shared territory than did females which, in contrast, spent most of their time within the centre of the territory. Despite some degree of territorial overlap between neighbouring conspecifics, their mutual avoidance, achieved through a system of temporal range utilization, resulted in few agonistic encounters.  相似文献   

9.
We examined the pattern of territory settlement and its consequences for breeding success in the Northern Wheatear Oenanthe oenanthe on Bardsey Island, Wales, during the breeding seasons of 1991-93. Males returned earlier than females, and older males returned earlier than first-year breeders. Although their boundaries shifted between years, the general location of territories was consistent during the three-year study. There was a high degree of fidelity to area and territory between years for both sexes. The order of territory settlement, from which a territory rank was calculated, was highly consistent for males between years irrespective of individual settlement patterns and territory fidelity. Patterns of territory settlement were less predictable for females, although there was a significant correlation between the mean territory ranks of paired males and females. There was a male-biased sex ratio in each year, and between 5% and 26% of males remained unpaired throughout the breeding season. Male mating status and breeding success were dependent on arrival date, territory rank and breeding density. Early-arriving, usually older, males were able to settle on territories first and were more likely to pair, while later-arriving individuals were more likely to remain unmated. These effects were consistent between years, and consequently territories could be classified as either preferred (accounting for proportionately more breeding attempts) or non-preferred. Territory quality as opposed to individual quality appeared to explain much of the variation in breeding success, and both sexes benefited by breeding on preferred territories through enhanced breeding success and an increased probability that their offspring would be recruited to the population.  相似文献   

10.
1.?Because habitats have profound effects on individual fitness, there is strong selection for improving the choice of breeding habitat. One possible mechanism is for individuals to use public information when prospecting future breeding sites; however, to our knowledge, no study has shown prospecting behaviour to be directly linked to subsequent choice of breeding site and future reproductive success. 2.?We collected long-term data on territory-specific prospecting behaviour and subsequent breeding in the short-lived northern wheatear (Oenanthe oenanthe). Non-breeders established prospecting territories (<2 ha) that overlapped the breeding territories of conspecifics. We tested whether: (i) prospectors used social and environmental cues that predicted territory-specific breeding success in the following year, and (ii) the prospecting territory was tightly linked to the subsequent breeding territory of the prospector, and whether this link would be weakened by intraspecific competition with original territory owners if they also survived. 3.?As expected, prospectors were attracted to a combination of site-specific cues that predicted future breeding success, i.e. short ground vegetation, a successfully breeding focal pair and successful close neighbours. 4.?Prospecting behaviour was directly linked to the choice of the following year's breeding territory: 79% of surviving prospectors established a breeding territory at their prospecting site in the following year, with their breeding success being higher than other individuals of the same age. As predicted, fidelity to the prospected site was strongly dependent on whether the original territory owner of the same sex had died or moved. 5.?Our findings suggest that the use of multiple cues reduces the negative impact of stochasticity on the reliability of social cues at small spatial scales (e.g. territories) and hence increases the probability of breeding success in the next year. Also, the use of conspecific attraction (i.e. the preference for breeding aggregations) is selectively advantageous because individuals are more likely to find a vacancy in an aggregation as compared to a solitary site. By extension, we hypothesize that species life-history traits may influence the spatial scale of prospecting behaviour and habitat selection strategies.  相似文献   

11.
1.?While the reasons for group-living have been studied for decades, little is known about why individuals become solitary. 2.?Several previous experimental studies could demonstrate that group-living can arises as a consequence of ecological constraints. 3.?It has been argued that reproductive competition between group members leads to significant costs of group-living, being a main reason of solitary-living. However, so far, no studies tested experimentally whether reproductive competition can explain solitary-living. 4.?Using a socially flexible species, the African striped mouse (Rhabdomys pumilio), we tested experimentally in the field whether dispersal and solitary-living are more likely to occur when reproductive competition is present. 5.?We investigated ecological constraints, here expressed as a function of population density, by removing groups of striped mice and creating vacant territories. To control for the effect of reproductive competition, which occurs only during the breeding season, we performed experiments during both the breeding and the non-breeding season. This is the first removal experiment performed in a species with communal breeding during the non-breeding season. 6.?During the breeding season, when population density was low, more striped mice from experimental groups moved into the vacant territories and became solitary than striped mice from control groups. This is in support of the ecological constraints hypothesis. 7.?During the non-breeding season, striped mice remained group-living despite the availability of free territories. Significantly, more striped mice became solitary-living during the breeding than during the non-breeding season. This is the first experimental support for the reproductive competition hypothesis explaining solitary-living. 8.?Analysis of the sexual maturity of males showed that males which became solitary had a higher reproductive potential than males that remained group-living. Analysis of the body mass data of females showed that more solitary females reproduced than group-living females. These results indicate that by becoming solitary individuals of both sexes avoided costs of reproductive competition within groups. 9.?Our study provides experimental evidence that reproductive competition within groups can lead to dispersal and solitary-living.  相似文献   

12.
Spatial processes could play an important role in density-dependent population regulation because the disproportionate use of poor quality habitats as population size increases is widespread in animal populations-the so-called buffer effect. While the buffer effect patterns and their demographic consequences have been described in a number of wild populations, much less is known about how dispersal affects distribution patterns and ultimately density dependence. Here, we investigated the role of dispersal in spatial density dependence using an extraordinarily detailed dataset from a reintroduced Mauritius kestrel (Falco punctatus) population with a territorial (despotic) breeding system. We show that recruitment rates varied significantly between territories, and that territory occupancy was related to its recruitment rate, both of which are consistent with the buffer effect theory. However, we also show that restricted dispersal affects the patterns of territory occupancy with the territories close to release sites being occupied sooner and for longer as the population has grown than the territories further away. As a result of these dispersal patterns, the strength of spatial density dependence is significantly reduced. We conclude that restricted dispersal can modify spatial density dependence in the wild, which has implications for the way population dynamics are likely to be impacted by environmental change.  相似文献   

13.
Two spatial tactics are usually distinguished in males of Calopteryx damselflies: territorial and nonterritorial. These tactics are believed to underlie two alternative condition-dependent reproductive tactics in these insects, and territorial males are believed to copulate more often. With age, males become weaker, turn nonterritorial, and only occasionally manage to copulate. However, the details of space use by damselflies are poorly known, which hinders the interpretation of the existing empirical data. We describe the space use by individually marked males of the banded demoiselle C. splendens studied during three field seasons in Vladimir Province, Russia. Each male on each day of observations was characterized as either territorial or non-territorial, and the sites of encounter were mapped. The probability of being territorial declined with the male’s age. The spatial tactics (territorial vs. non-territorial) on a given day strongly influenced the tactics used on the following day. We identified the territorial and non-territorial phases in the life of a male damselfly, which occurred consecutively and had a roughly similar duration. During the territorial phase, the male occupied a certain territory and tried to hold it as long as possible. The male abandoned its territory in two cases: (1) when it was driven onto a different territory as the result of competition with other males, or (2) when it was exhausted and became non-territorial. Thus, the space use by the male changed predictably during its life. Therefore, direct comparison of morphological or other characteristics in territorial vs. non-territorial males, frequently made in the literature, makes little sense. Further progress in studying the so-called “alternative reproductive tactics” in damselflies may be more successfully achieved by comparing individual life trajectories of different males (e.g. duration of territorial and non-territorial periods, the number of consecutively occupied territories, etc.). We performed correlation analysis and found that the above parameters did not depend on the wing and abdomen length of the males.  相似文献   

14.
Selection of breeding location can influence reproductive success and fitness. Breeding dispersal links habitat use and reproduction. This study investigated factors affecting breeding dispersal and its reproductive consequences in grey seals (Halichoerus grypus) on Sable Island, Nova Scotia. Breeding dispersal distance was determined in 692 individually marked, known-age female grey seals observed from 2004 to 2014. We used generalized linear mixed-effects models to test hypotheses concerning environmental and demographic factors influencing breeding dispersal distance and the consequences of dispersal distance on offspring weaning mass. Grey seal females rarely exhibited fidelity to previous breeding sites. Median dispersal distance between years was 5.1 km. Only 2.9% of females returned to a previous breeding site. Breeding dispersal distance was affected by parity and density, but effects were small and are presumably of no biological significance. Variation in dispersal distance among adult females was large. Dispersal distance had no significant influence on offspring weaning mass; however, as previously found, pup sex and maternal age did. Although breeding location was not important, heavier pups were born in habitats with no tidal or storm-surge influence indicating that breeding habitat type did influence offspring size at weaning. The lack of site fidelity in grey seals on Sable Island is associated with an unpredictable and changing landscape (sand dunes) that could make it difficult for females to locate previous breeding locations. Although breeding location within habitat type had small consequences on offspring weaning mass, we detected no evidence that breeding site selection within the habitat had consequences to females.  相似文献   

15.
While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.  相似文献   

16.
17.
In a population of Lapwings Vanellus vanellus studied for 5 years near Bergen, southwester Norway, 23–41% of the males were polygynous, having two and sometimes three mates, whereas 50–77% were monogamous and 0–21% remained unmated. Bachelors held territories in or immediately adjacent to the study area each of the years. As predicted from the Polygyny Threshold model, primary females generally laid eggs earlier than monogamous females, but the overlap in laying dates was substantial between these two classes. The number of females and start of breeding on the territories were analysed in relation to nine variables pertaining to safety from predation and to food. Of these, territory size correlated most consistently with number of females (positively) and the laying of first egg (negatively). Numbers of females observed were not significantly different from numbers expected from territory size in 4 out of 5 years; however, earlier start of breeding on large territories indicated that females did not settle according to the Neutral Mate Choice model. The Female-biased Sex Ratio hypothesis was refuted, unmated territorial males being available throughout the breeding seasons.  相似文献   

18.
Male-male competition over territorial ownership suggests that winning is associated with considerable benefits. In the speckled wood butterfly, Pararge aegeria, males fight over sunspot territories on the forest floor; winners gain sole residency of a sunspot, whereas losers patrol the forest in search of females. It is currently not known whether residents experience greater mating success than non-residents, or whether mating success is contingent on environmental conditions. Here we performed an experiment in which virgin females of P. aegeria were allowed to choose between a resident and a non-resident male in a large enclosure containing one territorial sunspot. Resident males achieved approximately twice as many matings as non-residents, primarily because matings were most often preceded by a female being discovered when flying through a sunspot. There was no evidence that territorial residents were more attractive per se, with females seen to reject them as often as non-residents. Furthermore, in the cases where females were discovered outside of the sunspot, they were just as likely to mate with non-residents as residents. We hypothesize that the proximate advantage of territory ownership is that light conditions in a large sunspot greatly increase the male's ability to detect and intercept passing receptive females.  相似文献   

19.
Most wood‐warblers (Parulidae) are non‐migratory residents of the Neotropics and subtropics, and the demographic characteristics of these species are poorly known. I examined the annual survival, reproductive output, dispersal, age of first breeding, and other demographic characteristics of a permanently territorial non‐migratory tropical warbler, the Slate‐throated Redstart (Myioborus miniatus), based on a 5‐yr study of a color‐banded population in Monteverde, Costa Rica. Territorial males showed strong site fidelity, but 26% of females engaged in short‐distance between‐year breeding dispersal. Estimated annual survival of territory holders, corrected for undetected female breeding dispersal, was 0.56 for males and 0.43 for females, values lower than expected and comparable to survival estimates for North American migrant warblers. The lower annual survival of females had two demographic consequences; unpaired territorial males were present in 3 of 5 yr, and some 1‐yr‐old males appeared to be floaters. Unpaired females or female floaters, however, were not observed. Mean natal dispersal distance was significantly greater for females (935 m) than males (485 m). Estimated first‐year survival was 0.29, but this is almost certainly an underestimate because of undetected long‐distance, female‐biased natal dispersal. Annual fecundity (fledglings per female) was 1.8, less than that of temperate warblers and attributable to small mean clutch sizes and a low incidence of double brooding. Estimated population growth rate (λ) was <1 for both males and females, suggesting that the study population was a demographic sink, most likely due to lower‐than‐expected adult survival.  相似文献   

20.
Defense of a territory is a primary factor determining the fitness of territorial individuals. Thus, territorial animals use a variety of means to lower the cost of territorial defense. Although landmarks are known to lower the defense costs of territories, little is known about how they could affect fundamental properties of territories, which could ultimately affect the demography, population regulation, individual behavior, and spatial ecology of a territorial species. This field study examined the effects of natural landmarks on territory size, shape, and defensive costs in Amatitlania siquia, a little‐studied member of the convict cichlid group native to Costa Rica and Nicaragua. We investigated the territorial behavior of 103 breeding pairs in a caldera lake in Nicaragua. During 10‐min focal observations, we recorded the locations at which residents chased intruders and then used these locations to outline the territory boundaries. Landmarks affected both size and shape of A. siquia territories. Territory size decreased as the percentage of the boundary at landmarks increased. Territories with <20% of their boundaries at landmarks were approximately circular in shape, while territories with 20–60% of their boundaries at landmarks were on average more elongated. These variations in territory size and shape significantly affected defense costs, with higher numbers of chases per square meter in small and elongated territories. These effects of landmarks on territory size and shape and defense costs are useful to understand the widely seen size and shape variations in naturally occurring territories.  相似文献   

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