When is dispersal for dispersal? Unifying marine and terrestrial perspectives

Recent syntheses on the evolutionary causes of dispersal have focused on dispersal as a direct adaptation, but many traits that influence dispersal have other functions, raising the question: when is dispersal ‘for’ dispersal? We review and critically evaluate the ecological causes of selection on traits that give rise to dispersal in marine and terrestrial organisms. In the sea, passive dispersal is relatively easy and specific morphological, behavioural, and physiological adaptations for dispersal are rare. Instead, there may often be selection to limit dispersal. On land, dispersal is relatively difficult without specific adaptations, which are relatively common. Although selection for dispersal is expected in both systems and traits leading to dispersal are often linked to fitness, systems may differ in the extent to which dispersal in nature arises from direct selection for dispersal or as a by‐product of selection on traits with other functions. Our analysis highlights incompleteness of theories that assume a simple and direct relationship between dispersal and fitness, not just insofar as they ignore a vast array of taxa in the marine realm, but also because they may be missing critically important effects of traits influencing dispersal in all realms.     

Can assignment tests measure dispersal?

Individual-based assignment tests are now standard tools in molecular ecology and have several applications, including the study of dispersal. The measurement of natal dispersal is vital to understanding the ecology of many species, yet the accuracy of assignment tests in situations where natal dispersal is common remains untested in the field. We studied a metapopulation of the grand skink, Oligosoma grande, a large territorial lizard from southern New Zealand. Skink populations occur on isolated, regularly spaced rock outcrops and are characterized by frequent interpopulation dispersal. We examined the accuracy of assignment tests at four replicate sites by comparing long-term mark-and-recapture records of natal dispersal with the results of assignment tests based on microsatellite DNA data. Assignment tests correctly identified the natal population of most individuals (65-100%, depending on the method of assignment), even when interpopulation dispersal was common (5-20% dispersers). They also provided similar estimates of the proportions of skinks dispersing to those estimated by the long-term mark-and-recapture data. Fully and partially Bayesian assignment methods were equally accurate but their accuracy depended on the stringency applied, the degree of genetic differentiation between populations, and the number of loci used. In addition, when assignments required high confidence, the method of assignment (fully or partially Bayesian) had a large bearing on the number of individuals that could be assigned. Because assignment tests require significantly less fieldwork than traditional mark-and-recapture approaches (in this study < 3 months vs. > 7 years), they will provide useful dispersal data in many applied and theoretical situations.     

When can dispersal synchronize populations?

While spatial synchrony of oscillating populations has been observed in many ecological systems, the causes of this phenomenon are still not well understood. The most common explanations have been the Moran effect (synchronous external stochastic influences) and the effect of dispersal among populations. Since ecological systems are typically subject to large spatially varying perturbations which destroy synchrony, a plausible mechanism explaining synchrony must produce rapid convergence to synchrony. We analyze the dynamics through time of the synchronizing effects of dispersal and, consequently, determine whether dispersal can be the mechanism which produces synchrony. Specifically, using methods new to ecology, we analyze a two patch predator-prey model, with identical weak dispersal between the patches. We find that a difference in time scales (i.e. one population has dynamics occurring much faster than the other) between the predator and prey species is the most important requirement for fast convergence to synchrony.     

Does polyploidy facilitate long-distance dispersal?

Background and Aims

The ability of plant lineages to reach all continents contributes substantially to their evolutionary success. This is exemplified by the Poaceae, one of the most successful angiosperm families, in which most higher taxa (tribes, subfamilies) have global distributions. Due to the old age of the ocean basins relative to the major angiosperm radiations, this is only possible by means of long-distance dispersal (LDD), yet the attributes of lineages with successful LDD remain obscure. Polyploid species are over-represented in invasive floras and in the previously glaciated Arctic regions, and often have wider ecological tolerances than diploids; thus polyploidy is a candidate attribute of successful LDD.

Methods

The link between polyploidy and LDD was explored in the globally distributed grass subfamily Danthonioideae. An almost completely sampled and well-resolved species-level phylogeny of the danthonioids was used, and the available cytological information was assembled. The cytological evolution in the clade was inferred using maximum likelihood (ML) as implemented in ChromEvol. The biogeographical evolution in the clade was reconstructed using ML and Bayesian approaches.

Key Results

Numerous increases in ploidy level are demonstrated. A Late Miocene–Pliocene cycle of polyploidy is associated with LDD, and in two cases (the Australian Rytidosperma and the American Danthonia) led to secondary polyploidy. While it is demonstrated that successful LDD is more likely in polyploid than in diploid lineages, a link between polyploidization events and LDD is not demonstrated.

Conclusions

The results suggest that polyploids are more successful at LDD than diploids, and that the frequent polyploidy in the grasses might have facilitated the extensive dispersal among continents in the family, thus contributing to their evolutionary success.     

Birthplace‐dependent dispersal: are directions of natal dispersal determined a priori?

Sib–sib or, more generally, family resemblance for dispersal seems a widespread characteristic of vertebrates, and the birthplace has the potential to shape the dynamics and features of animal populations. Dispersal studies have often stressed the fundamental link between the fate of dispersers and population dynamics, but few have focused on the dispersal directions of individuals, despite the profound implications that this may have on population distribution, structure, dynamics and viability. We investigated the directions followed by 72 radio‐tagged dispersers (43 males and 29 females from 14 nest sites) in an eagle owl Bubo bubo population, and assessed their a) inter‐individual distances during dispersal and b) age at dispersal departure. For siblings, as well as potential‐siblings (i.e. individuals born in the same nest in different years), the birthplace influenced inter‐individual distances and dispersal directions, i.e. dispersers from the same nest moved to similar locations during the study; moreover, in each year, individuals from the same birthplace moved across the same areas in a short time period. Finally, siblings and potential‐siblings born in the same nest in different years started dispersal at similar ages. Based on the movement patterns of dispersers we discuss: a) the potential implications of the birthplace‐dependent dispersal on the ideal free distribution theory, as well as in terms of kin competition, inbreeding avoidance and population dynamics; and, more generally, b) the effect of the temporal features of the natal dispersal on the concept of habitat suitability vs density of individuals developed by the ideal free distribution theory.     

Seed dispersal of East Asian coastal dune plants via seawater – short and long distance dispersal

Coastal plants provide precious and irreplaceable services to human and coastal ecosystems, but people are still rather unaware how coastal plants relate with sea tides. We assumed that their seeds may reflect some relationships with tides. Our objective was to understand seed dispersal of plants living in the upper coastal regions, not reached by normal tides, but inundated by storm surges. For this purpose, we observed seed characteristics, especially seed buoyancy, of coastal and non-inundated plants of the Shandong Peninsula, north China. Through field simulation, we studied how buoyancy affected the dispersal during tides, using twig segments as retrievable dummies of seeds, in order to collect evidences with concern, how seed dispersal could occur during storm surges, which are extremes of tides. Coastal plants had predominantly buoyant seeds, while inland-growing plants had largely non-buoyant seeds. Ninety percent of buoyant twig segments (mimicking seeds) were recaptured after tides on experimentally used coastal areas; only 1% of non-buoyant ones of such dummies were recaptured, whereas the rest probably had been swept away by wave action to deposit at the sea bottom. Buoyant twig segments (mimicking seeds) dispersed along the coastlines as tides surged towards the coast. About 10% of these buoyant twigs were swept away with off-shore currents, but they might land elsewhere, similar as it might happen with long-distance dispersal of seeds.     

Estimating fitness consequences of dispersal: a road to 'know-where'? Non-random dispersal and the underestimation of dispersers' fitness

1. Many studies investigating fitness correlates of dispersal in vertebrates report dispersers to have lower fitness than philopatric individuals. However, if dispersers are more likely to produce dispersing young or are more likely to disperse again in the next year(s) than philopatric individuals, there is a risk that fitness estimates based on local adult survival and local recruitment will be underestimated for dispersers. 2. We review the available empirical evidence on parent-offspring resemblance and individual lifelong consistency in dispersal behaviour, and relate these studies to recent studies of fitness correlates of dispersal in vertebrates. 3. Of the 12 studies testing directly for parent-offspring resemblance in dispersal propensity, five report a significant resemblance. The average effect size (r) of parent-offspring resemblance in dispersal was 0.15 [95% confidence interval (CI) = 0.07-0.22], with no difference between the sexes (average weighted effect size of 0.12 (0.08-0.16) and 0.16 (0.11-0.20) for females and males, respectively). Only three studies report data on within-individual consistency in dispersal propensity, of which two suggest dispersers to be more likely to disperse again. 4. To assess the magnitude of fitness underestimation expected for dispersing individuals depending on the heritability of dispersal distance and study area size, we used a simulation approach. Even when study area size is 10 times the mean dispersal distance, local recruitment per breeding event may be underestimated by 4-10%, generating a potential difference of 4-60% in average lifetime production of recruits between dispersing and philopatric individuals, with larger differences in long-lived species. 5. Estimates of both fitness correlates of dispersal and parent-offspring resemblance or within-individual consistency in dispersal behaviour have been reported for 11 species. Although some comparisons suggest genuine differences in fitness components between philopatric and dispersing individuals, others, based on adult and juvenile survival, are open to the alternative explanation of biased fitness estimates. 6. We list three potential ways of reducing the risk of making wrong inferences on biased fitness estimates due to such non-random dispersal behaviour between dispersing and philopatric individuals: (a) diagnosing effects of non-random dispersal, (b) reducing the effects of spatially limited study area and (c) performing controlled experiments.     

How much is a lot? Seed dispersal by white-faced capuchins and implications for disperser-based studies of seed dispersal systems

The quantity of seeds dispersed is considered one of several means to determine the dispersal effectiveness of an animal. However, there is little consistency in the manner in which quantities are measured or presented. Here, we quantify seed dispersal by white-faced capuchin monkeys (Cebus capucinus) in Santa Rosa National Park, Costa Rica by measuring: degree of frugivory, number of plant species consumed, the number of seeds consumed per unit time, the number of seeds dispersed intact per unit time, and the number of seeds dispersed intact per unit space. Forty-nine percent of C. capucinus diet is composed of the fruit of 39 species, 4 of which constitute 82% of the frugivory. Seventy-four percent of consumed fruits contain seeds that pass intact through the capuchin digestive system. Capuchins pass a mean of 15.7 seeds of a mean of 1.3 species per defecation, and defecate 25.4 times per 12-h day. These numbers are compared with extant data for C. capucinus, and possible reasons for discrepancies among results between studies are discussed. We propose a standardization of quantitative measures of seed dispersal so that quantifications of seed dispersal can be compared within species, and eventually across species.     

Do rivers function as corridors for plant dispersal?

Abstract. We evaluated the importance of dispersal for species frequencies and distribution by comparing dispersal properties of vascular plant species with their frequencies along river banks. We assumed that species with long-floating seeds would be more frequent than species with short-floating seeds. We compiled data on frequencies of vascular plants and their dispersal properties from ten rivers in northern Sweden and compared these with boreal forests and grasslands in the same region. In all rivers, but in none of the reference areas, there was a positive relationship between floating capacity and frequency of species. A comparison of floating capacity between species with and without certain dispersal devices showed that seeds of vegetatively dispersed species had higher floating capacities than other seeds. For other dispersal categories (animal and wind dispersal), floating time did not differ from contrast groups. The results indicate that water dispersal has a certain role in structuring the riparian flora, and provide a basis for explaining species distribution patterns from dispersal characteristics. They also suggest that continuous river corridors are important for maintaining regional biodiversity.     

Coevolution of dispersal in a parasitoid–host system

Interspecific interactions and the evolution of dispersal are both of interest when considering the potential impact of habitat fragmentation on community ecology, but the interaction between these processes is not well studied. We address this by considering the coevolution of dispersal strategies in a host–parasitoid system. An individual-based host–parasitoid metapopulation model was constructed for a patchy environment, allowing for evolution in dispersal rates of both species. Highly rarefied environments with few suitable patches selected against dispersal in both species, as did relatively static environments. Provided that parasitoids persist, all the variables studied led to stable equilibria in dispersal rates for both species. There was a tendency toward higher dispersal rates in parasitoids because of the asymmetric relationships of the two species to the patches: vacant patches are most valuable for hosts, but unsuitable for parasitoids, which require an established host population to reproduce. High host dispersal rate was favoured by high host population growth rate, and in the parasitoid by high growth rates in both species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Community-driven dispersal in an individual-based predator–prey model

We present a spatial, individual-based predator–prey model in which dispersal is dependent on the local community. We determine species suitability to the biotic conditions of their local environment through a time and space varying fitness measure. Dispersal of individuals to nearby communities occurs whenever their fitness falls below a predefined tolerance threshold. The spatiotemporal dynamics of the model is described in terms of this threshold. We compare this dynamics with the one obtained through density-independent dispersal and find marked differences. In the community-driven scenario, the spatial correlations in the population density do not vary in a linear fashion as we increase the tolerance threshold. Instead we find the system to cross different dynamical regimes as the threshold is raised. Spatial patterns evolve from disordered, to scale-free complex patterns, to finally becoming well-organized domains. This model therefore predicts that natural populations, the dispersal strategies of which are likely to be influenced by their local environment, might be subject to complex spatiotemporal dynamics.     

Do pools impede drift dispersal by stream insects?

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Can spatial variation alone lead to selection for dispersal?

The stability of models of age-dependent predation in continuous time with predators exhibiting a functional response are analyzed. A number of new features of biological importance emerge that are not present in simpler models. These include limits to the length of juvenile periods (both upper and lower) for stability, and the possibility that increases or decreases in any of the model parameters can be stabilizing or destabilizing. Hence, increased delays are not necessarily destabilizing. The variance in the length of the juvenile period is shown to be an important factor determining stability. Additionally, the relative stability of predation only on juveniles or only on adults is compared.     

At what scales does aggregated dispersal lead to coexistence?

Aggregation during dispersal can allow persistence of weak competitors, by creating conditions where stronger competitors are more likely to interact with conspecifics than with heterospecifics. However, aggregation mechanisms operate over a wide range of spatial scales, and species experience space in very different ways. The net effect of dispersal aggregation on coexistence will depend on how these scales interact. We show that it is possible to approximate the effects of aggregated dispersal on coexistence by considering three empirically measurable parameters: the spatial scale of interaction (how strongly competition drops off with distance), the spatial scale of aggregation (how large propagule packets are), and the temporal scale of aggregation (how frequently packets arrive). We use a novel metacommunity moment closure based on this approximation and stochastic simulations to show that aggregated dispersal allows for coexistence only when the stronger competitor is both aggregated and interacts at the same or a smaller spatial scale than the weaker competitor. When species interact and are aggregated at the same scales, coexistence outcomes are only weakly sensitive to the absolute scales of interaction and aggregation, as long as the scale of interaction is smaller than the scale of aggregation. However, coexistence is sensitive to the time‐scale of aggregation: increasing the frequency of packet arrival substantially reduces the region of fitness inequalities where both species persist. Finally, coexistence is less likely and global extinction of both competitors is more frequent when aggregation is fixed (with a constant number of propagules per packet), compared to density‐dependent (number of propagules increases with adult density).     

Spatial capture–recapture models allowing Markovian transience or dispersal

Spatial capture–recapture (SCR) models are a relatively recent development in quantitative ecology, and they are becoming widely used to model density in studies of animal populations using camera traps, DNA sampling and other methods which produce spatially explicit individual encounter information. One of the core assumptions of SCR models is that individuals possess home ranges that are spatially stationary during the sampling period. For many species, this assumption is unlikely to be met and, even for species that are typically territorial, individuals may disperse or exhibit transience at some life stages. In this paper we first conduct a simulation study to evaluate the robustness of estimators of density under ordinary SCR models when dispersal or transience is present in the population. Then, using both simulated and real data, we demonstrate that such models can easily be described in the BUGS language providing a practical framework for their analysis, which allows us to evaluate movement dynamics of species using capture–recapture data. We find that while estimators of density are extremely robust, even to pathological levels of movement (e.g., complete transience), the estimator of the spatial scale parameter of the encounter probability model is confounded with the dispersal/transience scale parameter. Thus, use of ordinary SCR models to make inferences about density is feasible, but interpretation of SCR model parameters in relation to movement should be avoided. Instead, when movement dynamics are of interest, such dynamics should be parameterized explicitly in the model.     

Are red imported fire ants facilitators of native seed dispersal?

Invasive ants threaten native communities, in part, through their potential to disrupt mutualisms, yet invasive species may also facilitate native species. The red imported fire ant (Solenopsis invicta) is one of the most conspicuous invasive ants in North America and its high densities, combined with its potential to displace native ants, have led to concerns that it may disrupt ant-plant seed dispersal mutualisms. We examined the potential of fire ants to disperse seeds in the longleaf pine ecosystem by comparing the removal of elaiosome-bearing seeds by fire ants versus native ants. A total of 14 ant species were observed removing seeds, with fire ants responsible for more than half of all removals. While fire ants were the dominant seed remover in this system, they did not remove significantly more seeds than would be expected based on their population density (46% of ground-dwelling ants). Moreover, red imported fire ants were similar to native ants with respect to distance of seed movement and frequency of moving seeds back to the nest. Areas of higher fire ant densities were found to have greater rates of seed removal by ants without a subsequent drop in seed dispersal by native ants, suggesting that fire ant-invaded areas may experience overall higher levels of seed dispersal. Thus, fire ants may actually facilitate dispersal of elaiosome-bearing plant species in the longleaf pine ecosystem.     

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.     

Does maternal oviposition site influence offspring dispersal to suitable habitat?

Orientation and dispersal to suitable habitat affects fitness in many animals, but the factors that govern these behaviors are poorly understood. In many turtle species, hatchlings must orient and disperse to suitable aquatic habitat immediately after emergence from subterranean nests. Thus, the location of nest sites relative to aquatic habitats ideally should be associated with the direction of hatchling dispersal. At our study site, painted turtles (Chrysemys picta) nest to the west (on an island) and east (on the mainland) of a wetland, which determines the direction that hatchlings must travel to reach suitable aquatic habitat. To determine if hatchling orientation is intrinsically influenced by the location where their mothers nest, we employed a two-part cross-fostering experiment in the field, whereby half the eggs laid in mainland nests were swapped with half the eggs laid in island nests. Moreover, because C. picta hatchlings overwinter inside their nests, we performed a second cross-fostering experiment to fully decouple the effects of (1) the maternally chosen nest location, (2) the embryonic developmental location, and (3) the overwinter location. We released hatchlings into a circular arena in the field and found that turtles generally dispersed in a westerly direction, regardless of the maternally chosen nest location and independent of the locations of embryonic development and overwintering. Although this westerly direction was towards suitable aquatic habitat, we could not distinguish whether naïve hatchling turtles (i) use environmental cues/stimuli to orient their movement, or (ii) have an intrinsic bias to orient west in the absence of stimuli. Nevertheless, these findings suggest that the orientation behavior of naïve hatchling turtles during terrestrial dispersal is not dependent upon the location of maternally-chosen nest sites.