Research progress in avian dispersal behavior

Dispersal, defined as a linear spreading movement of individuals away from others of the population is a fundamental characteristic of organisms in nature. Dispersal is a central concept in ecological, behavioral and evolutionary studies, driven by different forces such as avoidance of inbreeding depression, density-dependent competition and the need to change breeding locations. By effective dispersal, organisms can enlarge their geographic range and adjust the dynamic, sex ratio and genetic compositions of a population. Birds are one of the groups that are studied intensively by human beings. Due to their diurnal habits, diverse life history strategies and complex movement, birds are also ideal models for the study of dispersal behaviors. Certain topics of avian dispersal including sex-biased, asymmetric dispersal caused by differences in body conditions, dispersal processes, habitat selection and long distance dispersal are discussed here. Bird-ringing or marking, radio-telemetry and genetic markers are useful tools widely applied in dispersal studies. There are three major challenges regarding theoretical study and methodology research of dispersal: (1) improvement in research methodology is needed, (2) more in-depth theoretical research is necessary, and (3) application of theoretical research into the conservation efforts for threatened birds and the management of their habitats should be carried out immediately. __________ Translated from Acta Ecologica Sinica, 2008, 28(4): 1354–1365 [译自: 生态学报]     

鸟类的扩散行为研究进展

扩散是生物个体之间相互远离的单线性运动,是生物的基本特征之一,对种群的分布、动态及遗传结构等方面均有重要影响.扩散有出生扩散和繁殖扩散等主要形式.动物发生扩散的主要原因包括:避免近亲繁殖、减少竞争、改变繁殖地点等.近年来,扩散已经成为鸟类学研究的前沿领域.评述了鸟类扩散行为的性别差异、体质对于扩散的影响,阐述了扩散的基本过程及栖息地选择、长距离扩散等内容,同时介绍了环志标记、无线电遥测、分子生物学等研究鸟类扩散的主要方法.展望了鸟类扩散研究的发展趋势,认为新技术和新方法的应用将成为扩散生态学家关注的重要问题,未来研究将更加重视对鸟类扩散理论问题的探讨,而对鸟类扩散行为的研究成果也会更广泛地应用于濒危物种及其栖息地的保护工作中.     

Microsatellite data show evidence for male-biased dispersal in the Caribbean lizard Anolis roquet

Dispersal is a key component of an organism's life history and differences in dispersal between sexes appear to be widespread in vertebrates. However, most predictions of sex-biased dispersal have been based on observations of social structure in birds and mammals and more data are needed on other taxa to test whether these predictions apply in other organisms. Caribbean anole lizards are important model organisms in various biological disciplines, including evolutionary biology. However, very little is known about their dispersal strategies despite the importance of dispersal for population structure and dynamics. Here we use nine microsatellite markers to assess signatures of sex-biased dispersal on two spatial sampling scales in Anolis roquet, an anole endemic to the island of Martinique. Significantly higher gene diversity (H(S)) and lower mean assignment value (mAIC) was found in males on the larger spatial sampling scale. Significant heterozygote deficit (F(IS)), lower population differentiation (F(ST)), mAIC and variance of assignment index (vAIC) was found in males on the smaller spatial scale. The observation of male biased dispersal conform with expectations based on the polygynous mating system of Anolis roquet, and contributes to an explanation of the contrasting patterns of genetic structure between maternal and biparental markers that have been reported previously in this, and other anoline, species.     

Dispersal distances of Tree Swallows estimated from continent-wide and limited-area data

ABSTRACT.   Dispersal is a critical link between organismal and population biology, yet, because of their mobility, our understanding of the causes and consequences of long-distance dispersal by birds remains poorly known. Methods used to study dispersal include (1) marking and recapturing individuals in a limited study area to estimate survival and dispersal rates, and (2) relying on volunteers to mark and recapture individuals over larger areas. We compared these two methods for measuring dispersal distances of Tree Swallows ( Tachycineta bicolor ) using recapture data from a limited-area study in New York State (the Swallow Dispersal Study, SDS) and the recapture dataset from the U.S. Bird Banding Laboratory (BBL). Analysis of BBL records revealed a difference in the dispersal distance distributions (DDD) for data reported before and after 1967. In the earlier data, 84% of the 238 records were for birds within 13.6 km of their first banding location, whereas only 22% of the 799 records in the more recent data were reported in this closest distance belt. These differences are almost certainly due to changes in reporting protocols instituted by the BBL in the mid-1960s. We corrected for recapture effort in the SDS, and, using this corrected SDS data for the proportion of birds returning in the closest distance belt and the recent BBL for the proportions of more distant movements, we created what we think is the best composite DDD for Tree Swallows. Even though dispersal distances up to 2367 km have been reported, the composite DDD indicates that fewer than 3% of birds disperse more than 100 km and that 85% disperse less than 15 km between years. Thus, our results suggest that the dispersal behavior of most individuals can be examined effectively at more local spatial scales. Studies of dispersal and mortality would be facilitated if all recaptures of banded birds were reported with accurate spatial coordinates to the Bird Banding Laboratory.     

Dispersal, interpatch movements, and survival in a shrubland breeding bird community

ABSTRACT Dispersal events can affect the distribution, abundance, population structure, and gene flow of animal populations, but little is known about long‐distance movements due to the difficulty of tracking individuals across space. We documented the natal and breeding dispersal of shrubland birds among 13 study sites in a 1000 km² area in southeastern Ohio. In addition, we radio‐marked and tracked 37 adult males of one shrubland specialist, the Yellow‐breasted Chat (Icteria virens). We banded 1925 juveniles and 2112 adults of nine shrubland species from 2002 to 2005. Of these, 33 (1.7%) juveniles were encountered in subsequent years (2003–2006) as adults (natal dispersal) and 442 (20.9%) birds initially banded as breeding adults were re‐encountered in subsequent years (breeding dispersal). Apparent survival of juvenile shrubland birds on their natal patches was 0.024 (95% CI 0.016–0.036). After accounting for the probability of detection, we found that 21% of birds banded as juveniles and recaptured as adults returned to their natal patches, whereas 78% of adult birds showed fidelity to the patch where they were originally captured. Moreover, natal dispersers tended to move farther than breeding dispersers (corrected natal median = 1.7 km ± 0.37; corrected breeding median = 0.23 km ± 0.10). We used our estimates of natal dispersal and annual apparent survival to estimate true survival at 0.11 (95% CI 0.07–0.18) for juveniles in their first year. However, this estimate was only applicable for birds dispersing within 7 km of their natal patches. Interpatch movements of radio‐marked Yellow‐breasted Chats were not uncommon, with 13 of 37 males located in more than one habitat patch. Overall, we observed low natal philopatry, but high adult site fidelity for shrubland birds in our study area. Considering the frequency of short‐distance movements observed (median = 531 m, range = 88–1045 m), clustering of patches within 1 km might facilitate use of shrubland habitat.     

SPATIALLY CORRELATED EXTINCTIONS SELECT FOR LESS EMIGRATION BUT LARGER DISPERSAL DISTANCES IN THE SPIDER MITE TETRANYCHUS URTICAE

Dispersal is a central process to almost all species on earth, as it connects spatially structured populations and thereby increases population persistence. Dispersal is subject to (rapid) evolution and local patch extinctions are an important selective force in this context. In contrast to the randomly distributed local extinctions considered in most theoretical studies, habitat fragmentation or other anthropogenic interventions will lead to spatially correlated extinction patterns. Under such conditions natural selection is thought to lead to more long‐distance dispersal, but this theoretical prediction has not yet been verified empirically. We test this prediction in experimental spatially structured populations of the spider mite Tetranychus urticae and supplement these empirical results with insights from an individual‐based evolutionary model. We demonstrate that the spatial correlation of local extinctions changes the entire distribution of dispersal distances (dispersal kernel) and selects for overall less emigration but more long‐distance dispersal.     

Dispersal and alternative breeding site fidelity strategies in an amphibian

Dispersal (i.e. movement from a natal or breeding site to another breeding site) is a central process in ecology and evolution as it affects the eco‐evolutionary dynamics of spatially structured populations. Dispersal evolution is regulated by the balance between costs and benefits, which is influenced by the individual phenotype (i.e. phenotype‐dependent dispersal) and environmental factors (i.e. condition‐dependent dispersal). Even though these processes have been extensively studied in species with simple life cycles, our knowledge about these mechanisms in organisms displaying complex life cycles remains fragmentary. In fact, little is specifically known about how the interplay between individual and environmental factors may lead to alternative dispersal strategies that, in turn, lead to the coexistence of contrasted site fidelity phenotypes. In this paper, we examined breeding dispersal in a pond‐breeding amphibian, the great crested newt Triturus cristatus, within usual walking distances for a newt. We took advantage of recent developments in multi‐event capture–recapture models and used capture–recapture data (946 newts marked) collected in a spatially structured population occupying a large pond network (73 ponds). We showed a high rate of breeding site infidelity (i.e. pond use) and the coexistence of two dispersal phenotypes, namely, a highly pond faithful phenotype and a dispersing phenotype. Individuals that were site faithful at time t – 1 were therefore more likely to remain site faithful at time t. Our results also demonstrated that the probability that individuals belong to one or the other dispersal phenotypes depended on environmental and individual factors. In particular, we highlighted the existence of a dispersal syndrome implying a covariation pattern among dispersal behavior, body size, and survival. Our work opens new research prospects in the evolution of dispersal in organisms displaying complex life cycles and raises interesting questions about the evolutionary pathways that contribute to the diversification of movement strategies in the wild.     

16. Dispersal of freshwater algae — a review

A survey is given on the various methods for dispersal of freshwater algae. Dispersal factors are either water or air, or organisms — from beetles, dragonflies and mammals to birds, the latter being the most important group. The question of dispersal distances is discussed, in relation to dispersal mechanisms and to the resistance of the algae to transport conditions. Man's recent importance in algal dispersal is emphasized.     

Going against the flow: a case for upstream dispersal and detection of uncommon dispersal events

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Causes and consequences of animal dispersal strategies: relating individual behaviour to spatial dynamics

Knowledge of the ecological and evolutionary causes of dispersal can be crucial in understanding the behaviour of spatially structured populations, and predicting how species respond to environmental change. Despite the focus of much theoretical research, simplistic assumptions regarding the dispersal process are still made. Dispersal is usually regarded as an unconditional process although in many cases fitness gains of dispersal are dependent on environmental factors and individual state. Condition-dependent dispersal strategies will often be superior to unconditional, fixed strategies. In addition, dispersal is often collapsed into a single parameter, despite it being a process composed of three interdependent stages: emigration, inter-patch movement and immigration, each of which may display different condition dependencies. Empirical studies have investigated correlates of these stages, emigration in particular, providing evidence for the prevalence of conditional dispersal strategies. Ill-defined use of the term 'dispersal', for movement across many different spatial scales, further hinders making general conclusions and relating movement correlates to consequences at the population level. Logistical difficulties preclude a detailed study of dispersal for many species, however incorporating unrealistic dispersal assumptions in spatial population models may yield inaccurate and costly predictions. Further studies are necessary to explore the importance of incorporating specific condition-dependent dispersal strategies for evolutionary and population dynamic predictions.     

Population dynamics and dispersal of Lepthyphantes tenuis in an ephemeral habitat

The population dynamics of Lepthyphantes tenuis (Blackwall) were investigated in two fields of winter wheat in Sussex, England, with the aim of attributing the major factors controlling population fluctuations. Continuous measures of changes in density, reproductive rate and dispersal were obtained using a range of methodologies, including laboratory analysis, suction trapping, rotary sampling of the aerial fauna, caging, and detailed density estimations. The resulting data were then used to determine the major causes of the observed population fluctuations and to describe the processes of dispersal and reproduction in this population.Population densities were initially very low following ploughing operations, but increased throughout the season as a result of high reproductive rates. Investigation of the reproductive potential of the spiders showed that the number of eggs per egg-sac did not vary with season, and that egg development was temperature related. There was no evidence that the number of viable eggs per egg-sac varied during the season. There were three peaks of hatchling production per year suggesting two to three generations of spiders. Dispersal was not responsible for the major changes in field density throughout the season although dispersal activity was high for this species. Dispersal activity increased with age and was highest in females. Dispersal activity as a ratio of activity/field density for females, males, sub-adults, immatures and hatchlings was 4:2:3:1:1.Aerial dispersal is only possible under suitable weather conditions. It was found that female dispersal was correlated with weather conditions far more strongly than other population groups. Calculations showed that under suitable weather conditions (e.g., 25% of the time being suitable for dispersal), almost 4% of the adult females could be expected to leave the population daily. The overall effect is therefore to displace individuals from the population but to have little effect on densities. Dispersal in other stages was controlled by the suitability of weather conditions but also by other, undetermined factors. It is postulated that L. tenuis has a life history strategy whereby suitable habitats act as sources of spiders, mainly females, which are continually involved in dispersal. Dispersal in other population groups may be triggered by factors such as the avoidance of adverse conditions. This type of life-history strategy is typical of organisms, such as weeds, that are successful in ephemeral habitats and may indicate that this species originally evolved in unpredictable habitats such as dry river beds or coastal areas.     

Philopatry and natal dispersal in a sedentary population of western marsh harrier

Dispersal is a key life-history trait because it influences population dynamics and population genetic structure. From a behavioural perspective, the study of natal dispersal requires some understanding of the mechanisms that affect individual movements, because movements of an animal form a path that is continuous throughout its life. Our aim was to investigate juvenile dispersal strategies in the western marsh harrier Circus aeruginosus , between fledging and first breeding attempt (from 1 to 4 years later, depending on the sex and individual). Using radio tracking, we monitored dispersing juvenile harriers and their home-range size variations within a sedentary population in central western France from 2001 to 2007. Juvenile dispersal strategy was mainly characterized by a very high natal philopatry (i.e. birds that remained within the study area) and short-distance dispersal. All but one bird (out of 39) remained within the study area between their first winter and their first spring, and 96.0% during their first spring. The distance moved at 2 years of age was significantly larger for males than for females (3800± sd 3593 m and 935± sd 481 m for seven males and six females, respectively), in contrast to most bird species studied so far. Home-range size was not sex biased and significantly decreased with age. In addition, non-breeding birds had larger home ranges (1603± sd 2128 ha) than breeders (349± sd 185 ha). Using data obtained from other populations, juvenile marsh harrier dispersal strategies appeared to be determined by migratory status (migratory birds dispersing farther) and demographic parameters (juvenile survival or fecundity).     

Dispersal,kinship and inbreeding in an island population of the Great Tit

We investigated the kinship structure of an island population of the Great Tit (Parus major). Kinship of birds could be inferred by comparing their family trees. Dispersal was also studied to explain the observed pattern of kinship. On the island of Vlieland the tits breed in several wooded areas. Both males and females preferred to breed in their natal area; males did so more strongly than females. Hence gene flow between the areas is restricted. However, within the largest wooded area females showed random dispersal, while males showed a slight tendency to breed near their natal site. The degree of kinship of neighbouring birds is a suitable control group for the relatedness of partners that takes into account the effects of dispersal. In the largest wooded area, birds were on average equally related to their partner and to their neighbours. Moreover, the mean coefficient of kinship between male and female neighbours was equal to the average kinship in this part of the population. We conclude that mating is random with respect to kinship. There is no evidence for avoidance of inbreeding. It is unlikely that kin recognition plays an important role in the process of mate choice in this population of Great Tits. We suggest that ecological factors are the main causes for the observed patterns of dispersal and mating. On the island more female than male immigrants enter the population each year. Incidental data indicate an exchange of birds between the population studied and surrounding populations. Ancestries of immigrants are not known, and indeed a first analysis of all birds, including immigrants, showed that males were more closely related than females. However, differential immigration could not fully explain the observed difference in kinship. The presence of local adaptation in males is suggested as a possible additional cause.     

What is the required minimum landscape size for dispersal studies?

Among small animals dispersal parameters are mainly obtained by traditional methods using population studies of marked individuals. Dispersal studies may underestimate the rate and distance of dispersal, and be biased because of aggregated habitat patches and a small study area. The probability of observing long distance dispersal events decreases with distance travelled by the organisms. In this study a new approach is presented to solve this methodological problem. An extensive mark-release-recapture programme was performed in an area of 81 km(2) in southern Sweden. To estimate the required size of the study area for adequate dispersal measures we examined the effect of study area size on dispersal distance using empirical data and a repeated subsampling procedure. In 2003 and 2004, two species of diurnal burnet moths (Zygaenidae) were studied to explore dispersal patterns. The longest confirmed dispersal distance was 5600 m and in total 100 dispersal events were found between habitat patches for the two species. The estimated dispersal distance was strongly affected by the size of the study area and the number of marked individuals. For areas less than 10 km(2) most of the dispersal events were undetected. Realistic estimates of dispersal distance require a study area of at least 50 km(2). To obtain adequate measures of dispersal, the marked population should be large, preferably over 500 recaptured individuals. This result was evident for the mean moved distance, mean dispersal distance and maximum dispersal distance. In general, traditional dispersal studies are performed in small study areas and based on few individuals and should therefore be interpreted with care. Adequate dispersal measures for insects obtained by radio-tracking and genetic estimates (gene flow) is still a challenge for the future.     

Incorporating the geometry of dispersal and migration to understand spatial patterns of species distributions

Dispersal and migration can be important drivers of species distributions. Because the paths followed by individuals of many species are curvilinear, spatial statistical models based on rectilinear coordinates systems would fail to predict population connectivity or the ecological consequences of migration or species invasions. I propose that we view migration/dispersal as if organisms were moving along curvilinear geometrical objects called smooth manifolds. In that view, the curvilinear pathways become the ‘shortest realised paths’ arising from the necessity to minimise mortality risks and energy costs. One can then define curvilinear coordinate systems on such manifolds. I describe a procedure to incorporate manifolds and define appropriate coordinate systems, with focus on trajectories (1D manifolds), as parts of mechanistic ecological models. I show how a statistical method, known as ‘manifold learning’, enables one to define the manifold and the appropriate coordinate systems needed to calculate population connectivity or study the effects of migrations (e.g. in aquatic invertebrates, fish, insects and birds). This approach may help in the design of networks of protected areas, in studying the consequences of invasion, range expansions, or transfer of parasites/diseases. Overall, a geometrical view to animal movement gives a novel perspective to the understanding of the ecological role of dispersal and migration.     

Dispersal depends on body condition and predation risk in the semi‐aquatic insect,Notonecta undulata

Dispersal is the movement of organisms across space, which has important implications for ecological and evolutionary processes, including community composition and gene flow. Previous studies have demonstrated that dispersal is influenced by body condition; however, few studies have been able to separate the effects of body condition from correlated variables such as body size. Moreover, the results of these studies have been inconsistent with respect to the direction of the relationship between condition and dispersal. We examined whether body condition influences dispersal in backswimmers (Notonecta undulata). We also tested whether an interaction between body condition and predation risk (another proximate factor that influences dispersal) could contribute to the previously observed inconsistent relationship between condition and dispersal. We imposed diet treatments on backswimmers in the laboratory, and measured the effects of food availability on body condition and dispersal in the field. We found that dispersal was a positive function of body condition, which may have important consequences for population characteristics such as the rate of gene flow and population growth. However, the effects of body condition and predation risk were additive, not interactive, and therefore, our data do not support the hypothesis that the interaction between condition and predation risk contributes to the inconsistency in the results of previous condition‐dependent dispersal studies.     

Personality-dependent dispersal: characterization, ontogeny and consequences for spatially structured populations

Dispersal is one of the most fundamental components of ecology, and affects processes as diverse as population growth, metapopulation dynamics, gene flow and adaptation. Although the act of moving from one habitat to another entails major costs to the disperser, empirical and theoretical studies suggest that these costs can be reduced by having morphological, physiological or behavioural specializations for dispersal. A few recent studies on different systems showed that individuals exhibit personality-dependent dispersal, meaning that dispersal tendency is associated with boldness, sociability or aggressiveness. Indeed, in several species, dispersers not only develop behavioural differences at the onset of dispersal, but display these behavioural characteristics through their life cycle. While personality-dependent dispersal has been demonstrated in only a few species, we believe that it is a widespread phenomenon with important ecological consequences. Here, we review the evidence for behavioural differences between dispersers and residents, to what extent they constitute personalities. We also examine how a link between personality traits and dispersal behaviours can be produced and how personality-dependent dispersal affects the dynamics of metapopulations and biological invasions. Finally, we suggest future research directions for population biologists, behavioural ecologists and conservation biologists such as how the direction and the strength of the relationship between personality traits and dispersal vary with ecological contexts.     

Consequences of large-scale processes for the conservation of bird populations

1.  Detailed studies of population ecology are usually carried out in relatively restricted areas in which emigration and immigration play a role. We used a modelling approach to explore the population consequences of such dispersal and applied ideas from our simulations to the conservation of wild birds.
2.  Our spatial model incorporates empirically derived variation in breeding output between habitats, density dependence and dispersal. The outputs indicate that dispersal can have considerable consequences for population abundance and distribution. The abundance of a species within a patch can be markedly affected by the surrounding habitat matrix.
3.  Dispersal between habitats may result in lower population densities at the edge of good quality habitat blocks and could partially explain why some species are restricted to large habitat fragments.
4.  Habitat deterioration may not only lead to population declines within that habitat but also in adjacent habitats of good quality. This may confound studies attempting to diagnose population declines.
5.  Although mobile species have the advantages of colonizing sites within metapopulations, dispersal into poorer quality territories may markedly reduce total populations.
6.  There are two main approaches to conservation: one is to concentrate on establishing and maintaining protected areas, while the other involves conservation of the wider countryside. If dispersal is an important process then protecting only isolated areas may be insufficient to maintain the populations within them.     

Assessment of individual and conspecific reproductive success as determinants of breeding dispersal of female tree swallows: A capture–recapture approach

Breeding dispersal is a key process of population structure and dynamics and is often triggered by an individual's breeding failure. In both colonial and territorial birds, reproductive success of conspecifics (RSc) can also lead individuals to change breeding sites after a failure on a site. Yet, few studies have simultaneously investigated the independent contribution of individual reproductive success (RSi) and of RSc on dispersal decision. Here, we develop a modeling framework to disentangle the effects of RSi and RSc on demographic parameters, while accounting for imperfect individual detection and other confounding factors such as age or dispersal behavior in the previous year. Using a 10‐year capture–recapture dataset composed of 1,595 banded tree swallows, we assessed the effects of nonmanipulated RSi and RSc on female breeding dispersal in this semicolonial passerine. Dispersal was strongly driven by RSi, but not by RSc. Unsuccessful females were 9.5–2.5 times more likely to disperse than successful ones, depending if they had dispersed or not in the previous year, respectively. Unsuccessful females were also three times less likely to be detected than successful ones. Contrary to theoretical and empirical studies, RSc did not drive the decision to disperse but influenced the selection of the following breeding site once dispersal had been initiated. Because detection of individuals was driven by RSi, which was positively correlated to RSc, assuming a perfect detection as in previous studies may have lead us to conclude that RSc affected dispersal patterns, yet our approach corrected for this bias. Overall, our results suggest that the value and use of RSc as public information to guide dispersal decisions are likely dictated by multiple ecological determinants, such as landscape structure and extent, if this cue is indeed used.     

Variation in dispersal mortality and dispersal propensity among individuals: the effects of age, sex and resource availability

1.  Dispersal of individuals between habitat patches depends on both the propensity to emigrate from a patch and the ability to survive inter-patch movement. Environmental factors and individual characteristics have been shown to influence dispersal rates but separating the effects of emigration and dispersal mortality on dispersal can often be difficult. In this study, we use a soil mite laboratory system to investigate factors affecting emigration and dispersal mortality.
2.  We tested the movement of different age groups in two-patch systems with different inter-patch distances. Differences in immigration among age groups were primarily driven by differences in emigration but dispersal mortality was greater for some groups. Immigration declined with increasing inter-patch distance, which was due to increasing dispersal mortality and decreasing emigration.
3.  In a second experiment, we compared the dispersal of recently matured males and females and tested the impact of food availability during the developmental period on their dispersal. Dispersal was found to be male biased but there was no significant sex bias in dispersal mortality. There was some evidence that food availability could affect emigration and dispersal mortality.
4.  These results demonstrate that both emigration and dispersal mortality can be affected by factors such as individual age and resource availability. Understanding these effects is likely to be important for predicting the fitness costs and population consequences of dispersal.