Fitness-dependent dispersal in metapopulations and its consequences for persistence and synchrony

1. We present a novel metapopulation model where dispersal is fitness dependent: the strength of migration from a site is dependent on the expected reproductive fitness of individuals there. Furthermore, individuals continue to migrate until they reach a suitable habitat where their expected fitness is above a threshold value.
2. Fitness-dependent dispersal has a very strong stabilizing effect on population dynamics, even when the intrinsic dynamics of populations in the absence of dispersal exhibit complex high-amplitude oscillations. This stabilizing effect is much stronger than that of the density-independent dispersal normally considered in metapopulation models.
3. Even when fitness-dependent dispersal does not stabilize the dynamics in a formal sense, it generally leads to simplification, with complex or even chaotic fluctuations being reduced to simple cycles.
4. This form of dispersal also has a strong tendency to synchronize local population dynamics across the spatial extent of the metapopulation.
5. These conclusions are robust to the addition of strong stochasticity in the migration threshold.     

Dispersal ability and host-plant characteristics influence spatial population structure of monophagous beetles

Abstract.  1. Dispersal plays an integral role in determining spatial population structure and, consequently, the long-term survival of many species. Theoretical studies indicate that dispersal increases with population density and decreasing habitat stability. In the case of monophagous insect herbivores, the stability of host-plant populations may influence their spatial population structure.
2. The tallgrass prairie in Iowa, U.S.A. is highly fragmented and most prairie insects face a landscape with fewer habitat patches and smaller host-plant populations than 150 years ago, potentially making dispersal between patches difficult. Some herbivores, however, use native plant species with weedy characteristics that have increased in abundance because of disturbances.
3. Mark–recapture data and presence–absence surveys were used to examine dispersal and spatial population structure of two monophagous beetles with host plants that exhibit different population stability and have responded differently to fragmentation of tallgrass prairie.
4. Chrysochus auratus Fabricius exhibits a patchy population structure and has relatively large dispersal distances and frequencies. Its host plant is variable locally in time and space, but is more abundant than 150 years ago. The other species, Anomoea laticlavia Forster, exhibits a metapopulation or non-equilibrium population structure and has relatively small dispersal distances and frequencies. Its host-plant populations are stable in time and space.
5. The results indicate that dispersal ability of monophagous beetles reflects the life-history dynamics of their host plants, but the spatial population structure exhibited today is strongly influenced by how the host plants have responded to the fragmentation process over both time and space.     

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.     

Can population growth rates vary with the spatial scale at which they are measured?

1.  The ratio of successive population censuses is often assumed to reflect population growth rates. We identify three simple potential sources of bias in the estimation of population growth rates that relate to either the total number of censused individuals or the spatial areas over which censuses are conducted.
2.  The commonly used method of adding a constant to time series data to avoid problems caused by division by zero can lead to underestimation of growth rates at low densities in increasing populations.
3.  Variances associated with density estimates can lead to positive bias in estimation of growth rates when populations are distributed in ephemeral patches. The spatial variance and spatio-temporal covariance in bank vole census data suggest that this bias could be severe when small trapping grids are used. Use of logged estimators of growth rate avoids this problem.
4.  Using census data from non-randomly placed trapping grids that are smaller than twice the maximum range of natal dispersal to estimate population growth rates can lead to negatively biased estimates, particularly at low population densities.
5.  These three sources of bias are evaluated as explanations for scale-dependent changes in the estimates of growth rates identified in populations of snowshoe hare ( Lepus americanus ), bank voles ( Clethrionomys glareolus ) and lemmings ( Lemmus lemmus ).     

Flight activity of adult stoneflies in relation to weather

Abstract. 1. Dispersal of adult aquatic insects between streams may have important consequences for local and regional population dynamics, but little is known about how dispersal is affected by weather conditions.
2. The influence of meteorological variables on flight activity of adult stoneflies (Plecoptera: Leuctridae, Nemouridae, and Chloroperlidae) was investigated using Malaise traps adjacent to three upland streams in the Plynlimon area of mid Wales, U.K.
3. Numbers of adult stoneflies captured weekly in the traps were related positively to air temperature and related negatively to wind speed. Meteorological conditions during daylight showed stronger relationships with flight activity than did conditions at night.
4. There was inter-site variation in the strength of weather effects on stonefly flight. Wind speed was significant at only one site, which had higher average wind speed than the other sites.
5. Annual variation in weather conditions during adult flight periods may result in varying extent of dispersal between sites, influencing community dynamics over a wide area.     

Population synchrony and stability in environmentally forced metacommunities

A general prediction from simple metapopulation models is that spatially synchronized forcing can spatially synchronize population dynamics and destabilize metapopulations. In contrast, spatially asynchronous forcing is predicted to decrease population synchrony and promote temporal stability and population persistence, especially in the presence of dispersal. Only recently have studies begun to experimentally address these predictions. Moreover, few studies have experimentally examined how such processes operate in the context of competition communities. Stabilizing processes may continue to operate when placed within a metacommunity context with multiple competing consumers but only at low to intermediate levels of dispersal. High dispersal rates can reverse these predictions and lead to destabilization. We tested this under controlled conditions using an experimental aquatic system composed of three competing species of zooplankton. Metacommunities experienced different levels of dispersal and environmental forcing in the form of spatially synchronous or asynchronous pH perturbations. We found support that dispersal can have contrasting effects on population stability depending on the degree to which population dynamics were synchronized in space. Dispersal under synchronous forcing or no forcing had either neutral of positive effects on spatial population synchrony of all three zooplankton species. In these treatments, dispersal reduced population stability at the local and metapopulation levels for two of three species. In contrast, asynchronously varying environments reduced population synchrony relative to unforced systems, regardless of dispersal level. In these treatments, dispersal enhanced temporal stability and persistence of populations not by reducing population synchrony but by enhancing population minima and spatial averaging of abundances. High dispersal rates under asynchronous forcing reduced the abundance of one species, consistent with increasing regional competition and general metacommunity theory. However, no effects on its stability or persistence were observed. Our work highlights the context‐dependent effects of dispersal on population dynamics in varying environments.     

Population structure of a monophagous moth in a patchy landscape

1. The population structure of a monophagous noctuid moth, Abrostola asclepiadis , living on a patchily distributed perennial herb, Vincetoxicum hirundinaria is described. The study took place over 5 years at a landscape scale (about 12 km²).
2. Patch occupancy rates and population densities were studied in relation to patch size, degree of patch isolation, level of sun exposure and distance from the coast. In addition, flight tests in the laboratory were performed to estimate the potential dispersal capacity of the moth.
3. Occupancy rates were high and the likelihood of extinction depended on patch size. Small patches were less likely to be occupied than were large patches (> 10 m²). Sun-exposed patches were occupied for a lower proportion of years than were shaded patches. No distance effects could be discerned at the spatial scale of study, presumably because the insect is a strong flier.
4. Population densities in occupied patches decreased with increasing patch size. Furthermore, insect densities tended to increase with distance from the coast. Density changes in patches were synchronized.
5. The studied insect population can be described as a 'patchy population' sensu Harrison (1991) with spatially correlated population dynamics. These dynamics are superimposed on a landscape gradient.     

Testing the mechanisms by which source-sink dynamics alter competitive outcomes in a model system

Dispersal among sites can affect within-site competitive outcomes via source-sink dynamics. Source-sink dynamics are thought to affect competitive outcomes primarily via spatial subsidies: by redistributing individuals from sources to sinks, source-sink dynamics can alter competitive outcomes in both sources and sinks. However, dispersal also can affect competitive outcomes via demography modification, which occurs when dispersal alters the parameters governing species' per capita demographic rates. For instance, dispersal of exploitative competitors might cause extinction of some of the resources for which competition occurs, thereby altering the competition coefficients. I used protist microcosms as a model system to test whether spatial subsidies alone could explain the effects of source-sink dynamics on competitive outcomes. I examined the long-term outcome of exploitative competition among three bacterivorous ciliate protists in microcosms of high enrichment (sources) and low enrichment (sinks) in both the presence and the absence of dispersal. Dispersal altered competitive outcomes. Fitting mathematical models to the population dynamics revealed that spatial subsidies were insufficient to account for the effects of dispersal. Fitting alternative models strongly suggested that demography modification was an important determinant of competitive outcomes. These results provide the first evidence that dispersal does not simply redistribute competitors but can alter their per capita demographic rates.     

描述Meta-种群动态的耦合映象格子模型

Ｍｅｔａ－种群观点为在局部种群之上的空间尺度上描述种群的生态学过程提供了一种途径。但是由于植物所拥有的若干特殊性质（如种子休眠、有限扩散和局部适应）使得对某些植物种使用原有的Ｍｅｔａ－种群概念和模型存在许多困难。因此,为了能更精确地反映植物Ｍｅｔａ－种群动态,本文将Ｍｅｔａ－种群动态分解为斑块内局部种群动态和斑块间的扩散过程两个分量。用Ｌｏｇｉｓｔｉｃ方程表述每个斑块内的局部种群动态,用扩散系数ε表示斑块间的扩散,建立了描述Ｍｅｔａ－种群动态的耦合映象格子模型。对单峰映象ｘｎ＋１＝１－ａｘ２ｎ给出了耦合映象格子的时空行为,这对Ｍｅｔａ－种群动态的研究可能是非常重要的。     

Measuring dispersal and detecting departures from a random walk model in a grasshopper hybrid zone

Abstract. 1. The grasshopper species Chorthippus brunneus and C. jacobsi form a complex mosaic hybrid zone in northern Spain. Two mark–release–recapture studies were carried out near the centre of the zone in order to make direct estimates of lifetime dispersal.
2. A model framework based on a simple random walk in homogeneous habitat was extended to include the estimation of philopatry and flying propensity. Each model was compared with the real data, correcting for spatial and temporal biases in the data sets.
3. All four data sets (males and females at each site) deviated significantly from a random walk. Three of the data sets showed strong philopatry and three had a long dispersal tail, indicating a low propensity to move further than predicted by the random walk model.
4. Neighbourhood size estimates were 76 and 227 for the two sites. These estimates may underestimate effective population size, which could be increased by the long tail to the dispersal function. The random walk model overestimates lifetime dispersal and hence the minimum spatial scale of adaptation.
5. Best estimates of lifetime dispersal distance of 7–33 m per generation were considerably lower than a previous indirect estimate of 1344 m per generation. This discrepancy could be influenced by prezygotic isolation, an inherent by-product of mosaic hybrid zone structure.     

Gene flow and genetic structure of the aquatic macrophyte Sparganium emersum in a linear unidirectional river

1. River systems offer special environments for the dispersal of aquatic plants because of the unidirectional (downstream) flow and linear arrangement of suitable habitats.
2. To examine the effect of this flow on microevolutionary processes in the unbranched bur-reed ( Sparganium emersum ) we studied the genetic variation within and among nine (sub)populations along a 103 km stretch of the Niers River (Germany–The Netherlands), using amplified fragment length polymorphisms.
3. Genetic diversity in S. emersum populations increased significantly downstream, suggesting an effect of flow on the pattern of intrapopulation genetic diversity.
4. Gene flow in the Niers River is asymmetrically bidirectional, with gene flow being approximately 3.5 times higher in a downstream direction. The observed asymmetry is probably caused by frequent hydrochoric dispersal towards downstream locations on the one hand, and sporadic zoochoric dispersal in an upstream direction on the other. The spread of vegetative propagules (leaf and stem fragments) is probably not an important mode of dispersal for S. emersum , suggesting that gene flow is mainly via seed dispersal. Realized dispersal distances exceeded 60 km, revealing a potential for long-distance dispersal in S. emersum .
5. There was no correlation between geographical and genetic distances among the nine S. emersum populations (i.e. no isolation by distance), which may be due to the occurrence of long-distance dispersal and/or colonization and extinction dynamics in the Niers River.
6. Overall, the genetic population structure and regional dispersal patterns of S. emersum in the Niers River are best explained by a linear metapopulation model. Our study shows that flow can exert a strong influence on population genetic processes of plants inhabiting stream systems.     

The ecology and mechanisms of overflow‐mediated dispersal in a rock‐pool metacommunity

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Metapopulation dynamics of a flightless alpine insect Hemideina maori in a naturally fragmented habitat

Abstract 1. Despite widespread acceptance of metapopulation theory, the effects that inter-patch dispersal and variability in patch size have on metapopulation dynamics in insects are two issues that require further study. In addition, previous studies of metapopulations have tended to focus on organisms with high dispersal capabilities such as some species of butterfly and bird.
2. Mountain stone weta Hemideina maori are a long-lived, flightless orthopteran that live on island rock outcrops or tors in the alpine region of southern New Zealand. A total of 480 adults and 789 juveniles was marked over three seasons on four large and 14 small tors to assess the effects of habitat fragmentation on the population dynamics of H. maori .
3. Only 12 adults (2.5% of marked adults and 4.0% of recaptured adults) and two juveniles (0.3% of marked juveniles and 0.7% of recaptured juveniles) dispersed between tors. The mean dispersal distance was 361 m (range = 36–672 m). Larger tors supported larger populations and had a higher number of emigrants and immigrants while smaller tors had proportionally higher emigration and immigration rates. Although adults on large and small tors had similar mean lifespans, five extinction events and three recolonisation events occurred during the study period, all on small tors.
4. Hemideina maori conform to many of the predictions of metapopulation theory even though they are flightless, show relatively low dispersal rates, and occur at low densities. Extinction and colonisation events are more common on small tors but may be relatively unimportant for the long-term survival of the metapopulation because they occur on the smallest habitat patches, which support the smallest proportion of the overall population.     

A dispersal-induced paradox: synchrony and stability in stochastic metapopulations

Understanding how dispersal influences the dynamics of spatially distributed populations is a major priority of both basic and applied ecologists. Two well-known effects of dispersal are spatial synchrony (positively correlated population dynamics at different points in space) and dispersal-induced stability (the phenomenon whereby populations have simpler or less extinction-prone dynamics when they are linked by dispersal than when they are isolated). Although both these effects of dispersal should occur simultaneously, they have primarily been studied separately. Herein, I summarise evidence from the literature that these effects are expected to interact, and I use a series of models to characterise that interaction. In particular, I explore the observation that although dispersal can promote both synchrony and stability singly, it is widely held that synchrony paradoxically prevents dispersal-induced stability. I show here that in many realistic scenarios, dispersal is expected to promote both synchrony and stability at once despite this apparent destabilising influence of synchrony. This work demonstrates that studying the spatial and temporal impacts of dispersal together will be vital for the conservation and management of the many communities for which human activities are altering natural dispersal rates.     

Spatial dynamics of Microtus vole populations in continuous and fragmented agricultural landscapes

Small mammal populations often exhibit large-scale spatial synchrony, which is purportedly caused by stochastic weather-related environmental perturbations, predation or dispersal. To elucidate the relative synchronizing effects of environmental perturbations from those of dispersal movements of small mammalian prey or their predators, we investigated the spatial dynamics of Microtus vole populations in two differently structured landscapes which experience similar patterns of weather and climatic conditions. Vole and predator abundances were monitored for three years on 28 agricultural field sites arranged into two 120-km-long transect lines in western Finland. Sites on one transect were interconnected by continuous agricultural farmland (continuous landscape), while sites on the other were isolated from one another to a varying degree by mainly forests (fragmented landscape). Vole populations exhibited large-scale (>120 km) spatial synchrony in fluctuations, which did not differ in degree between the landscapes or decline with increasing distance between trapping sites. However, spatial variation in vole population growth rates was higher in the fragmented than in the continuous landscape. Although vole-eating predators were more numerous in the continuous agricultural landscape than in the fragmented, our results suggest that predators do not exert a great influence on the degree of spatial synchrony of vole population fluctuations, but they may contribute to bringing out-of-phase prey patches towards a regional density level. The spatial dynamics of vole populations were similar in both fragmented and continuous landscapes despite inter-landscape differences in both predator abundance and possibilities of vole dispersal. This implies that the primary source of synchronization lies in a common weather-related environment.     

Spatially autocorrelated disturbances and patterns in population synchrony

Spatially synchronous population dynamics have been documented in many taxa. The prevailing view is that the most plausible candidates to explain this pattern are extrinsic disturbances (the Moran effect) and dispersal. In most cases disentangling these factors is difficult. Theoretical studies have shown that dispersal between subpopulations is more likely to produce a negative relationship between population synchrony and distance between the patches than perturbations. As analyses of empirical data frequently show this negative relationship between the level of synchrony and distance between populations, this has emphasized the importance of dispersal as a synchronizing agent. However, several weather patterns show spatial autocorrelation, which could potentially produce patterns in population synchrony similar to those caused by dispersal. By using spatially extended versions of several population dynamic models, we show that this is indeed the case. Our results show that, especially when both factors (spatially autocorrelated perturbations and distance-dependent dispersal) act together, there may exist groups of local populations in synchrony together but fluctuating asynchronously with some other groups of local populations. We also show, by analysing 56 long-term population data sets, that patterns of population synchrony similar to those found in our simulations are found in natural populations as well. This finding highlights the subtlety in the interactions of dispersal and noise in organizing spatial patterns in population fluctuations.     

Viability analyses with habitat-based metapopulation models

Population viability analysis (PVA) models incorporate spatial dynamics in different ways. At one extreme are the occupancy models that are based on the number of occupied populations. The simplest occupancy models ignore the location of populations. At the other extreme are individual-based models, which describe the spatial structure with the location of each individual in the population, or the location of territories or home ranges. In between these are spatially structured metapopulation models that describe the dynamics of each population with structured demographic models and incorporate spatial dynamics by modeling dispersal and temporal correlation among populations. Both dispersal and correlation between each pair of populations depend on the location of the populations, making these models spatially structured. In this article, I describe a method that expands spatially structured metapopulation models by incorporating information about habitat relationships of the species and the characteristics of the landscape in which the metapopulation exists. This method uses a habitat suitability map to determine the spatial structure of the metapopulation, including the number, size, and location of habitat patches in which subpopulations of the metapopulation live. The habitat suitability map can be calculated in a number of different ways, including statistical analyses (such as logistic regression) that find the relationship between the occurrence (or, density) of the species and independent variables which describe its habitat requirements. The habitat suitability map is then used to calculate the spatial structure of the metapopulation, based on species-specific characteristics such as the home range size, dispersal distance, and minimum habitat suitability for reproduction. Received: April 1, 1999 / Accepted: October 29, 1999     

Host tree influences on the dispersal of first instar gypsy moths, Lymantria dispar (L.)

Abstract. 1. In laboratory tests, first instar gypsy moths attempted dispersal more frequency when exposed to less acceptable foliage.
2. First instars from small eggs attempted dispersal less frequently than larvae from large eggs when exposed to foliage from highly acceptable or marginally acceptable hosts. Dispersal rates of larvae from medium sized eggs were intermediate.
3. These results (1–2) confirm and expand upon the findings of Capinera & Barbosa (1976).
4. In the field, data on the relative densities of larvae on different host species support the conclusion that the frequency of dispersal attempts is inversely related to host acceptability.
5. The implications of these findings for the population dynamics of the gypsy moth are discussed.     

Active dispersal is differentially affected by inter‐ and intraspecific competition in closely related nematode species

Competition is one of the main drivers of dispersal, which can be an important mechanism to achieve permanent or temporal coexistence of multiple species. This coexistence can be achieved by a dispersal‐competition tradeoff, spatial store effects or neutral dynamics. Here we test the effect of inter‐ and intraspecific competition on dispersal of four species of the marine nematode species complex Litoditis marina. A previous study in closed microcosms without a possibility for dispersal had demonstrated pronounced interspecific competition, leading to the exclusion of one species. We now investigated whether 1) the dispersal is affected by interspecific interactions, by intraspecific competition (density) or by food availability, 2) the dispersal dynamics influence assemblage composition and can lead to co‐occurrence of the species, and 3) the abiotic environment (here salinity) can affect these dynamics. We show that density is the main driver for dispersal in two of the four species. Dispersal of a third species always started at the same time irrespective of density, whereas in the fourth species interspecific interactions accelerated dispersal. Remarkably, this fourth species was not a strong competitor, suggesting that a dispersal–competition tradeoff does not explain the observed coexistence. Salinity did not alter the timing of dispersal when interspecific interactions were present but did affect assemblage composition. Consequently, spatial store effects may influence coexistence. All four species co‐occurred in fairly stable abundances throughout the present experiment indicating the importance of species specific dispersal strategies for coexistence. Co‐occurrence can be facilitated because competition is postponed or avoided by dispersal. Neutral dynamics also played a role as intra‐ and interspecific competition were of similar importance in three of the four species. We conclude that dispersal is a driver of the coexistence of closely related nematode species, and that population density and interspecific interactions shape these dynamics.