Metapopulation dynamics for spatially extended predator–prey interactions

Traditional metapopulation theory classifies a metapopulation as a spatially homogeneous population that persists on neighboring habitat patches. The fate of each population on a habitat patch is a function of a balance between births and deaths via establishment of new populations through migration to neighboring patches. In this study, we expand upon traditional metapopulation models by incorporating spatial heterogeneity into a previously studied two-patch nonlinear ordinary differential equation metapopulation model, in which the growth of a general prey species is logistic and growth of a general predator species displays a Holling type II functional response. The model described in this work assumes that migration by generalist predator and prey populations between habitat patches occurs via a migratory corridor. Thus, persistence of species is a function of local population dynamics and migration between spatially heterogeneous habitat patches. Numerical results generated by our model demonstrate that population densities exhibit periodic plane-wave phenomena, which appear to be functions of differences in migration rates between generalist predator and prey populations. We compare results generated from our model to results generated by similar, but less ecologically realistic work, and to observed population dynamics in natural metapopulations.     

Migration and the genetic covariance between habitat preference and performance

Studies of the genetic covariance between habitat preference and performance have reported conflicting outcomes ranging from no covariance to strong covariance. The causes of this variability remain unclear. Here we show that variation in the magnitude of genetic covariance can result from variability in migration regimes. Using data from walking stick insects and a mathematical model, we find that genetic covariance within populations between host plant preference and a trait affecting performance on different hosts (cryptic color pattern) varies in magnitude predictably among populations according to migration regimes. Specifically, genetic covariance within populations is high in heterogeneous habitats where migration between populations locally adapted to different host plants generates nonrandom associations (i.e., linkage disequilibrium) between alleles at color pattern and host preference loci. Conversely, genetic covariance is low in homogeneous habitats where a single host exists and migration between hosts does not occur. Our results show that habitat structure and patterns of migration can strongly affect the evolution and variability of genetic covariance within populations.     

An expert-based landscape permeability model for assessing the impact of agricultural management on amphibian migration

The persistence of pond-breeding amphibian populations is influenced on different spatial scales ranging from the individual breeding pond to surrounding habitat patches to landscape clusters of breeding pond populations. The connectivity among breeding ponds as well as the availability of suitable terrestrial habitats surrounding the ponds plays a major role in long-term viability of amphibian species. Besides road traffic and urban structures agricultural land activity can disrupt landscape connectivity through the use of pesticides, fertilizers and physical activity such as tillage. We developed an expert-based model to assess the impact of agricultural management measures on the migration area and terrestrial habitat availability for seven amphibian species. The model is based on a Habitat Suitability Index to identify suitable terrestrial habitats and includes a landscape permeability approach. Size of migration areas, connectivity of breeding ponds and number of reachable terrestrial habitats were modeled considering species-specific migration ranges and habitat preferences. We consider how pesticides application might lead to fragmentation and isolation of amphibian breeding pond populations. Therefore the potential disrupting impact of pesticides was simulated by inflating landscape resistance to medium and high migration cost. One amphibian species showed a decrease of migration area by 48.3% and a decrease of reachable terrestrial habitats by 41.5% at high migration costs. Three additional species showed a decrease of their migration areas between 31.5 and 35.7%. At increased migration cost, some of the investigated populations were isolated at breeding pond level or restricted to pond clusters. Our model could be used to prioritize conservation efforts for pond-breeding amphibians with adequate consideration of agricultural land use and its impact on amphibian migration.     

Chaotic dynamics may determine the effect of inter-patch migration on metapopulation survival

A simple, strategic model of a system of habitat fragments connected by conservation corridors is presented. The intrinsic dynamics of the population on each fragment are stochastic. In addition, at each generation there is a probability of a catastrophic event occurring which affects all the habitat fragments by greatly reducing the size of the population on each. Global extinction is considered to occur when all the populations simultaneously fall below a threshold value. If the intrinsic dynamics on each fragment are simple cycles or a stable equilibrium, then the addition of conservation corridors does not reduce the frequency of global extinction. This is because migration between fragments induces their populations to have values which are similar to each other. However, if the intrinsic population dynamics are chaotic then the probability of global extinction is greatly reduced by the introduction of conservation corridors. Although local extinction is likely, the chaos acts to oppose the synchronising effect of migration. Often a subset of the populations survive a catastrophe and can recolonize the other patches.     

Population dynamics in migratory networks

Migratory animals are comprised of a complex series of interconnected breeding and nonbreeding populations. Because individuals in any given population can arrive from a variety of sites the previous season, predicting how different populations will respond to environmental change can be challenging. In this study, we develop a population model composed of a network of breeding and wintering sites to show how habitat loss affects patterns of connectivity and species abundance. When the costs of migration are evenly distributed, habitat loss at a single site can increase the degree of connectivity (mixing) within the entire network, which then acts to buffer global populations from declines. However, the degree to which populations are buffered depends on where habitat loss occurs within the network: a site that has the potential to receive individuals from multiple populations in the opposite season will lead to smaller declines than a site that is more isolated. In other cases when there are equal costs of migration to two or more sites in the opposite season, habitat loss can result in some populations becoming segregated (disconnected) from the rest of the network. The geographic structure of the network can have a significant influence on relative population sizes of sites in the same season and can also affect the overall degree of mixing in the network, even when sites are of equal intrinsic quality. When a migratory network is widely spaced and migration costs are high, an equivalent habitat loss will lead to a larger decline in global population size than will occur in a network where the overall costs of migration are low. Our model provides an important foundation to test predictions related to habitat loss in real-world migratory networks and demonstrates that migratory networks will likely produce different dynamics from traditional metapopulations. Our results provide strong evidence that estimating population connectivity is a prerequisite for successfully predicting changes in migratory populations.     

Carry‐over effects of winter habitat quality on en route timing and condition of a migratory passerine during spring migration

We examined how conditions prior to migration influenced migration performance of two breeding populations of black‐and‐white warblers Mniotilta varia by linking information on the migrant's winter habitat quality, measured via stable carbon isotopes, with information on their breeding destination, measured via stable hydrogen isotopes. The quality of winter habitat strongly influenced the timing of migration when we accounted for differential timing of migration between breeding populations. Among birds migrating to the same breeding destination, males and females arriving early to the stopover site originated from more mesic habitat than later arriving birds, suggesting that the benefits of occupying high‐quality mesic habitat during the winter positively influence the timing of migration. However, male warblers arriving early to the stopover site were not in better migratory condition than later arriving conspecifics that originated from poor‐quality xeric winter habitat, regardless of breeding destination. The two breeding populations stopover at the study site during different time periods, suggesting that the lower migratory condition of early birds is not a function of the time of season, but potentially a migrant's migration strategy. Strong selection pressures to arrive early on the breeding grounds to secure high‐quality breeding territories may drive males from high‐quality winter habitat to minimize time at the expense of energy. This migration strategy would result in a smaller margin of safety to buffer the effects of adverse weather or scarcity of food, increasing the risk of mortality. The migratory condition of females was the same regardless of the timing of migration or breeding destination, suggesting that females adopt a strategy that conserves energy during migration. This study fills an important gap in our understanding of the linkages between winter habitat quality and factors that influence the performance of migration, the phase of the annual cycle thought to be limiting most migratory bird populations.     

Density-dependent migration and stability in a system of linked populations

The effect of adding density-dependent migration between nearest neighbour populations of a single discrete-generation species in a chain of habitat fragments is investigated. The larger the population on a particular habitat fragment, the greater the fraction of inhabitants who migrate before reproducing. It has previously been shown for similar models with density-independent migration that coupling populations in this way has no effect on the stability of these populations. Here, it is demonstrated that this effect is also generally true if migration is density-dependent. However, if the migration rate is large enough and has density dependence of the correct form, then the steady state (with all the populations remaining at the same constant value through time) can be destabilised. The conditions for this to occur are obtained analytically. When this “destabilisation” occurs, the system settles down to an alternative steady state where half of the populations take one constant value which is below that of an equivalent isolated system, and the other populations all share a population value which is greater than the steady state of the isolated populations. Once this configuration is reached, the population size on each patch remains constant over time. hence the change might more properly be described as a decrease in homogeneity rather than in stability.     

Evolution of migration rate in a spatially realistic metapopulation model

We use an individual-based, spatially realistic metapopulation model to study the evolution of migration rate. We first explore the consequences of habitat change in hypothetical patch networks on a regular lattice. If the primary consequence of habitat change is an increase in local extinction risk as a result of decreased local population sizes, migration rate increases. A nonmonotonic response, with migration rate decreasing at high extinction rate, was obtained only by assuming very frequent catastrophes. If the quality of the matrix habitat deteriorates, leading to increased mortality during migration, the evolutionary response is more complex. As long as habitat patch occupancy does not decrease markedly with increased migration mortality, reduced migration rate evolves. However, once mortality becomes so high that empty patches remain uncolonized for a long time, evolution tends to increase migration rate, which may lead to an "evolutionary rescue" in a fragmented landscape. Kin competition has a quantitative effect on the evolution of migration rate in our model, but these patterns in the evolution of migration rate appear to be primarily caused by spatiotemporal variation in fitness and mortality during migration. We apply the model to real habitat patch networks occupied by two checkerspot butterfly (Melitaea) species, for which sufficient data are available to estimate rigorously most of the model parameters. The model-predicted migration rate is not significantly different from the empirically observed one. Regional variation in patch areas and connectivities leads to regional variation in the optimal migration rate, predictions that can be tested empirically.     

Density-dependent dispersal suggests a genetic measure of habitat suitability

Recent research shows that density dependence should result in predictable movements between habitats of different suitability, depending on whether population densities are increasing or decreasing. When population densities are increasing, habitats become filled in order of their suitability, resulting in a net flow from high suitability to low suitability. When populations decrease in density, the reverse can happen. These patterns suggest that genetic information can be used to infer habitat suitability since individual-based genetic assignment tests permit high resolution assessments of migration. We used replicated landscapes to study fishers ( Martes pennanti ) during a population increase and predicted that there should be a net flow of individuals from areas of shallow to deep snow, since snow depth has previously been linked to fisher fitness. A total of 769 fishers were sampled from 35 different landscapes and profiled at 16 microsatellite loci. From assignment tests, we inferred five genetic populations. By assigning each of the 35 landscapes to one of these five populations, we were able to determine the proportion of immigrants to each. Consistent with our prediction, there was a positive relationship between the proportion of immigrants and snow depth. The best model of fisher habitat suitability was one with both snow depth and the proportion of coniferous forest in landscapes. Our findings suggest that where population trend is known, genetic information can be used to measure habitat suitability.     

Single Unpurified Breast Tumor-Initiating Cells from Multiple Mouse Models Efficiently Elicit Tumors in Immune-Competent Hosts

The tumor-initiating cell (TIC) frequency of bulk tumor cell populations is one of the criteria used to distinguish malignancies that follow the cancer stem cell model from those that do not. However, tumor-initiating cell frequencies may be influenced by experimental conditions and the extent to which tumors have progressed, parameters that are not always addressed in studies of these cells. We employed limiting dilution cell transplantation of minimally manipulated tumor cells from mammary tumors of several transgenic mouse models to determine their tumor-initiating cell frequency. We determined whether the tumors that formed following tumor cell transplantation phenocopied the primary tumors from which they were isolated and whether they could be serially transplanted. Finally we investigated whether propagating primary tumor cells in different tissue culture conditions affected their resident tumor-initiating cell frequency. We found that tumor-initiating cells comprised between 15% and 50% of the bulk tumor cell population in multiple independent mammary tumors from three different transgenic mouse models of breast cancer. Culture of primary mammary tumor cells in chemically-defined, serum-free medium as non-adherent tumorspheres preserved TIC frequency to levels similar to that of the primary tumors from which they were established. By contrast, propagating the primary tumor cells in serum-containing medium as adherent populations resulted in a several thousand-fold reduction in their tumor-initiating cell fraction. Our findings suggest that experimental conditions, including the sensitivity of the transplantation assay, can dramatically affect estimates of tumor initiating cell frequency. Moreover, conditional on cell culture conditions, the tumor-initiating cell fraction of bulk mouse mammary tumor cell preparations can either be maintained at high or low frequency in vitro thus permitting comparative studies of tumorigenic and non-tumorigenic cancer cells.     

Limited dispersal by the rare scarce heath butterfly—potential consequences for population persistence

Dispersal between habitat patches may be important for the long-term persistence of populations. We conducted a mark–release–recapture study and analysed the dispersal pattern in the scarce heath butterfly inhabiting a network of suitable habitat patches using stepwise logistic regression (SLR) and the Virtual Migration (VM) model. We also analysed the influence of different types of matrices. We found that the majority of the recaptured butterflies remained within the patch where they were originally caught. However, dispersal between patches did occur and both the SLR analysis and the VM model indicated that the migration pattern was significantly associated with patch area and its level of isolation. The SLR model also showed that there was a positive association between immigration rate and tree density, supporting earlier observations that this species prefers semi-open habitat. We discuss the use of SLR versus the VM model to analyse recapture data in dispersal studies. This system is not at equilibrium, as a number of the most important patches in the network are continuously being lost due to afforestation and a number of populations are facing deterministic extinction. This increases the risk of a chain reaction of local extinctions, which may cause a collapse of the whole system.     

The response of bird populations to habitat loss

Environmental change through altered climate and land use could have a severe impact on bird populations. Predicting the consequences for the size of bird populations is one of the crucial problems for their conservation. We show how a population model based on the behaviour of individuals can be used to predict the consequences of habitat loss. For a wide range of conditions, loss of either wintering or breeding habitat results in population reduction. The approach is then extended to consider the impact of habitat loss in the wintering area on bird species with complex migratory systems. This shows that 'knock-on' effects may occur, so that habitat loss in a wintering area may affect populations which did not initially use that area. The ability to alter migration routes in response to environmental change may be crucial to the future viability of populations. Using a simple model combining genetics and population dynamics, we show that aspects of the biology of a species may affect whether or not its migration strategy is flexible enough to shift in response to habitat change. Some species may be able to adopt new migration routes and avoid the catastrophic consequences of habitat loss in traditional wintering areas; however, other species may lack this flexibility and may suffer severe population declines as a consequence.     

Dispersal, migration, and offspring retention in saturated habitats

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.     

Environmental conditions at arrival to the wintering grounds and during spring migration affect population dynamics of barn swallows Hirundo rustica breeding in Northern Italy

Several populations of long-distance migratory birds are currently suffering steep demographic declines. The identification of the causes of such declines is difficult because population changes may be driven by events occurring in distant geographical areas during different phases of the annual life-cycle of migrants. Furthermore, wintering areas and migration routes of populations of small-sized species are still largely unknown, with few exceptions. In this paper we identified the critical phases of the annual life-cycle that most influence the population dynamics of a small passerine, the Barn Swallow Hirundo rustica. We used information on temporal dynamics of a population breeding in Northern Italy, whose wintering range and timing of migration have been recently described by miniaturised tracking dataloggers. Our results indicated that primary productivity in the wintering grounds in the month when most individuals arrive from autumn migration and primary productivity in an area that is probably a stopover site during spring migration, influenced population dynamics more than habitat conditions at the breeding grounds. By using annual variation in primary productivity at the wintering grounds and stopover sites as predictors, we replicated the observed interannual population changes with great accuracy. However, the steep decline recently suffered by the population could be replicated only by including a constant annual decline in the model, suggesting that changes in primary productivity only predicted the interannual variation around the long-term trend. Our study therefore suggests the existence of critical periods during wintering and migration that may have large impact on population fluctuations of migrant birds.     

Metapopulation structure and movements in five species of checkerspot butterflies

We studied the patterns and rates of migration among habitat patches for five species of checkerspot butterflies (Lepidoptera: Melitaeini) in Finland: Euphydryas aurinia, E. maturna, Melitaea cinxia, M. diamina and M. athalia. We applied the virtual migration (VM) model to mark-release-recapture data collected from multiple populations. The model includes parameters describing migration and survival rates and how they depend on the areas and connectivities of habitat patches. The number of individuals captured varied from 73 to 1,123, depending on species and sex, and the daily recapture probabilities varied between 0.09-0.52. The VM model fitted the data quite well. The results show that the five species are broadly similar in their movement rates and patterns, though, e.g. E. maturna tends to move shorter distances than the other species. There is no indication of any phylogenetic component in the parameter values. The parameter values estimated for each species suggest that a large percentage (80-90%) of migration events were successful in the landscapes that were studied. The area of the habitat patch had a substantial effect on emigration and immigration rates, such that butterflies were more likely to leave small than large patches and large patches were more likely than small patches to receive immigrants.     

Three ways of assessing metapopulation structure in the butterfly Plebejus argus

1. Three independent methods were used to investigate population structure in the butterfly Plebejus argus . First, migration and dispersal ability were measured by mark–release–recapture in seven adjacent habitat patches, and by release of butterflies in unoccupied habitat. Secondly, colonization of newly created habitat was observed over 7 years. Finally, genetic differentiation of local populations within a metapopulation was investigated. Sampled local populations included parts of the mark–release–recapture study area.
2. Plebejus argus is relatively sedentary: the maximum movement detected was 395 m, and only 2% of individuals moved further than 100 m between recaptures on different days. None the less, adjacent local populations in the mark–release–recapture study area were linked by occasional migration, with ≈ 1.4% of individuals moving between patches separated by 13–200 m.
3. Despite low mobility, observed colonizations occurred rapidly over distances of 1 km. Because P. argus occurs at high population densities, 1.4% migration can generate enough migrants to colonize newly suitable habitat quickly at this spatial scale.
4. Mark–release–recapture data were used to predict that there would be limited genetic differentiation through drift between local populations at this spatial scale. The prediction was supported by allele frequency data for the same local populations.
5. Genetic differentiation often indicates higher levels of migration than are revealed by the movements of marked individuals. This study shows that when experimental releases and extensive marking are undertaken in areas that are large relative to most movements, indirect measures of gene flow and direct measures of dispersal can concur.
6. Evidence from the three different approaches was complementary, indicating that P. argus occurs as metapopulations within the study area.     

The migration game in habitat network: the case of tuna

Long-distance migration is a widespread process evolved independently in several animal groups in terrestrial and marine ecosystems. Many factors contribute to the migration process and of primary importance are intra-specific competition and seasonality in the resource distribution. Adaptive migration in direction of increasing fitness should lead to the ideal free distribution (IFD) which is the evolutionary stable strategy of the habitat selection game. We introduce a migration game which focuses on migrating dynamics leading to the IFD for age-structured populations and in time varying habitats, where dispersal is costly. The model predicts migration dynamics between these habitats and the corresponding population distribution. When applied to Atlantic bluefin tunas, it predicts their migration routes and their seasonal distribution. The largest biomass is located in the spawning areas which have also the largest diversity in the age-structure. Distant feeding areas are occupied on a seasonal base and often by larger individuals, in agreement with empirical observations. Moreover, we show that only a selected number of migratory routes emerge as those effectively used by tunas.     

Habitat barriers limit gene flow and illuminate historical events in a wide-ranging carnivore, the American puma

We examined the effects of habitat discontinuities on gene flow among puma (Puma concolor) populations across the southwestern USA. Using 16 microsatellite loci, we genotyped 540 pumas sampled throughout the states of Utah, Colorado, Arizona, and New Mexico, where a high degree of habitat heterogeneity provides for a wide range of connective habitat configurations between subpopulations. We investigated genetic structuring using complementary individual- and population-based analyses, the latter employing a novel technique to geographically cluster individuals without introducing investigator bias. The analyses revealed genetic structuring at two distinct scales. First, strikingly strong differentiation between northern and southern regions within the study area suggests little migration between them. Second, within each region, gene flow appears to be strongly limited by distance, particularly in the presence of habitat barriers such as open desert and grasslands. Northern pumas showed both reduced genetic diversity and greater divergence from a hypothetical ancestral population based on Bayesian clustering analyses, possibly reflecting a post-Pleistocene range expansion. Bayesian clustering results were sensitive to sampling density, which may complicate inference of numbers of populations when using this method. The results presented here build on those of previous studies, and begin to complete a picture of how different habitat types facilitate or impede gene flow among puma populations.     

三江源地区雪豹保护优先区规划

为了将有限资源合理投放到关键区域, 实现物种保护成效的最大化, 找出质量最好的栖息地及它们之间的迁徙通道是制定保护规划的第一步。本研究以三江源的雪豹(Panthera uncia)栖息地为对象, 基于野外调查数据和高分辨率卫星遥感数据, 利用物种分布模型、保护规划模型和连通度分析工具, 找出了三江源地区雪豹的核心栖息地分布和潜在迁徙通道位置, 分析了目前保护中的潜在威胁, 并提出了针对三江源西、中、东三块区域的不同保护对策。结果表明: (1)三江源西部核心栖息地比较小而破碎, 但迁徙通道较多且没有明显窄点, 未来应关注青藏线的潜在阻碍作用, 同时应防范道路沿线的野生动物盗猎; (2)中部区域横跨玉树-杂多-囊谦的雪豹栖息地是三江源最大的核心雪豹栖息地, 在连通其他种群中也处于中心地位, 应通过种群监测确定其健康稳定, 对开发、偷猎等威胁防微杜渐, 保持其源种群的作用; (3)东部区域人口密度高, 受人类活动的影响最大, 需保证阿尼玛卿、年保玉则两块核心栖息地的质量, 并重点监测甘德县境内的省道处雪豹的迁徙通道是否畅通。三江源地区雪豹栖息地总体质量较好, 建议将维持核心源种群的稳定性, 保持种群间迁徙通道的畅通作为三江源的雪豹景观保护工作的整体目标。未来应充分利用天地一体化监测手段, 开展重要保护物种栖息地状况的评估和预警, 尤其是非保护地区域物种核心栖息地的开发建设活动。