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     

Rapid Buildup of Genetic Diversity in Founder Populations of the Gynodioecious Plant Species Origanum vulgare after Semi-Natural Grassland Restoration

In most landscapes the success of habitat restoration is largely dependent on spontaneous colonization of plant species. This colonization process, and the outcome of restoration practices, can only be considered successful if the genetic makeup of founding populations is not eroded through founder effects and subsequent genetic drift. Here we used 10 microsatellite markers to investigate the genetic effects of recent colonization of the long-lived gynodioecious species Origanum vulgare in restored semi-natural grassland patches. We compared the genetic diversity and differentiation of fourteen recent populations with that of thirteen old, putative source populations, and we evaluated the effects of spatial configuration of the populations on colonization patterns. We did not observe decreased genetic diversity in recent populations, or inflated genetic differentiation among them. Nevertheless, a significantly higher inbreeding coefficient was observed in recent populations, although this was not associated with negative fitness effects. Overall population genetic differentiation was low (F_ST = 0.040). Individuals of restored populations were assigned to on average 6.1 different source populations (likely following the ‘migrant pool’ model). Gene flow was, however, affected by the spatial configuration of the grasslands, with gene flow into the recent populations mainly originating from nearby source populations. This study demonstrates how spontaneous colonization after habitat restoration can lead to viable populations in a relatively short time, overcoming pronounced founder effects, when several source populations are nearby. Restored populations can therefore rapidly act as stepping stones and sources of genetic diversity, likely increasing overall metapopulation viability of the study species.     

Very Low Population Structure in a Highly Mobile and Wide-Ranging Endangered Bird Species

The loss of biodiversity following fragmentation and degradation of habitat is a major issue in conservation biology. As competition for resources increases following habitat loss and fragmentation, severe population declines may occur even in common, highly mobile species; such demographic decline may cause changes within the population structure of the species. The regent honeyeater, Anthochaera phrygia, is a highly nomadic woodland bird once common in its native southeast Australia. It has experienced a sharp decline in abundance since the late 1970s, following clearing of large areas of its preferred habitat, box-ironbark woodland, within the last 200 years. A captive breeding program has been established as part of efforts to restore this species. This study used genetic data to examine the range-wide population structure of regent honeyeaters, including spatial structure, its change through time, sex differences in philopatry and mobility, and genetic differences between the captive and wild populations. There was low genetic differentiation between birds captured in different geographic areas. Despite the recent demographic decline, low spatial structure appears to have some temporal consistency. Both sexes appear to be highly mobile, and there does not seem to be significant genetic differentiation between the captive and wild populations. We conclude that management efforts for survival of this species, including habitat protection, restoration, and release of captive-bred birds into the wild, can treat the species as effectively a single genetic population.     

Demographic and genetic collapses in spatially structured populations: insights from a long‐term survey in wild fish metapopulations

Unraveling the relationship between demographic declines and genetic changes over time is of critical importance to predict the persistence of at‐risk populations and to propose efficient conservation plans. This is particularly relevant in spatially structured populations (i.e. metapopulations) in which the spatial arrangement of local populations can modulate both demographic and genetic changes. We used ten‐year demo‐genetic monitoring to test 1) whether demographic declines were associated with genetic diversity declines and 2) whether the spatial structure of a metapopulation can weaken or reinforce these demographic and genetic temporal trends. We continuously surveyed, over time and across their entire range, two metapopulations of an endemic freshwater fish species Leuciscus burdigalensis: one metapopulation that had experienced a recent demographic decline and a second metapopulation that was stable over time. In the declining metapopulation, the number of alleles rapidly decreased, the inbreeding coefficient increased, and a genetic bottleneck emerged over time. In contrast, genetic indices were constant over time in the stable metapopulation. We further show that, in the declining metapopulation, demographic and genetic declines were not homogeneously distributed across the metapopulation. We notably identify one local population situated downstream as a ‘reservoir’ of individuals and genetic variability that dampens both the demographic and genetic declines measured at the metapopulation level. We demonstrate the usefulness of long‐term monitoring that combines both genetic and demographic parameters to understand and predict temporal population fluctuations of at‐risk species living in a metapopulation context.     

Simulation modeling reveals the evolutionary role of landscape shape and species dispersal on genetic variation within a metapopulation

Different shapes of landscape boundaries can affect the habitat networks within them and consequently the spatial genetic-patterns of a metapopulation. In this study, we used a mechanistic framework to evaluate the effects of landscape shape, through watershed elongation, on genetic divergence among populations at the metapopulation scale. Empirical genetic data from four, sympatric stream-macroinvertebrates having aerial adults were collected from streams in Japan to determine the roles of species-specific dispersal strategies on metapopulation genetics. Simulation results indicated that watershed elongation allows the formation of river networks with fewer branches and larger topographic constraints. This results in decreased interpopulation connectivity but a lower level of spatial isolation of distal populations (e.g. those found in headwaters) occurring in the landscapes examined. Distal populations had higher genetic divergence when their downstream-biased dispersal (relative to upstream- and/or overland-biased dispersal) was high. This underscores the importance of distal populations influencing genetic divergence at the metapopulation scale for species having downstream-biased dispersal. In turn, lower genetic divergence was observed under watershed elongation when the genetic isolation of distal populations was decreased in such species. This strong association between landscape shape and evolutionary processes highlights the importance of natural, spatial architecture in assessing the effectiveness of conservation and management strategies.     

Wetland Restoration for Amphibians: Should Local Sites Be Designed to Support Metapopulations or Patchy Populations?

Pond-breeding amphibians have been characterized as having metapopulation structure, and a goal of many local restoration projects is to establish viable metapopulations. However, recent studies suggest that metapopulation organization is unlikely at the local level because of high dispersal rates between neighboring ponds. Although many amphibians avoid ovipositing in habitats that pose high predation risk to their offspring, the spatial scale of avoidance is poorly resolved for natural systems and could involve wholesale movements between ponds. To determine the scale of avoidance, we monitored annual habitat use by the Wood frog ( Rana sylvatica ), American toad ( Bufo americanus ), and Spotted salamander ( Ambystoma maculatum ) at a restoration site in western North Carolina, U.S.A. Wood frogs consistently used most fish-free ponds, but rapidly curtailed use following fish invasions. American toads rarely used the same breeding site from year to year, and adults strongly avoided ovipositing in habitats with predatory Wood frog tadpoles. Spotted salamanders exhibited a predator avoidance response to fish that was weaker than the predator avoidance response of anurans. Our data indicate that the spatial scale of predator avoidance by ovipositing amphibians often exceeds that of an individual pond and that the focal species at this site are organized as patchy populations rather than as metapopulations. At local restoration sites, ponds that are placed in spatial arrays to create metapopulations may not accomplish their goal and may limit the extent to which ovipositing adults can express an adaptive antipredator behavior. We discuss an alternative design that is more likely to enhance the long-term persistence of local populations.     

生境破坏的模式对集合种群动态和续存的影响

构建了空间关联的集合种群模型,该模型不但包含了种群的空间结构信息,而且引入了破坏生境的全局密度和局部密度两个指标,它们描述了破坏生境的模式.模型揭示了破坏生境的空间分布格局复杂地影响了集合种群的动态和续存,破坏和未破坏生境斑块的均匀混合不利于集合种群的增长和续存,而生境类型聚集分布可以促进集合种群的快速增长和长期续存;对于两种斑块类型相对均匀混合的生境来说,均匀场假设可能会高估集合种群的续存,对于相对斑块类型高度聚集的生境,均匀场假设可能会低估集合种群的续存;物种的迁移范围也会影响集合种群的续存,迁移范围越大的物种越容易抵御生境的破坏而免遭灭绝.这意味着在生物保护中不能仅仅考虑生境的恢复和斑块质量的改善,生境结构的构建也是很重要的,加强生境斑块之间的连通性也有利于物种的长期续存.     

Towards a population approach for evaluating grassland restoration—a systematic review

Persistence of restored populations depends on growth, reproduction, dispersal, local adaptation, and a suitable landscape pattern to foster metapopulation dynamics. Although the negative effects of habitat fragmentation on plant population dynamics are well understood, particularly in grasslands, the population traits that control grassland restoration are less known. We reviewed the use of population traits for evaluating grassland restoration success based on 141 publications (1986–2015). The results demonstrated that population demography was relatively well‐assessed but detailed studies providing information on key stages of the life cycle were lacking despite their importance in determining population viability. Vegetative and generative performances have been thoroughly investigated, notably the components of plant fitness, such as reproductive output, while genetic and spatial population structures were largely ignored. More work on the population effects of ecological restoration would be welcomed, particularly with a focus on population genetics. Targeted species were principally common and dominant natives, or invasive plants while rare or threatened species were poorly considered. Evaluation of ecological restoration should be conducted at different scales of ecological complexity, but so far, communities and ecosystems are over represented, and more focus should be directed towards a population approach as population traits are essential indicators of restoration success.     

Fish community composition and diversity at restored estuarine habitats in Tampa Bay,Florida, United States

Estuary restoration in Tampa Bay, Florida, United States, is an ongoing focus of natural resource managers because of pressure from an increasing coastal population, historic habitat loss, and restoration's importance to economic development, recreational activities, and fish habitat. A growing population can also limit future large‐scale restorations due to associations with cost and land availability. This limitation might be overcome by applying the habitat mosaic approach to restoration, which creates distinct habitat types at small spatial scales. This approach was applied to create three types of estuarine habitat, reconnected tidal creek, salt marsh, and tidal pond. The objectives of this study were to (1) initiate monitoring of a restored wetland mosaic and (2) determine how fish diversity and community structure vary among restored habitat types. Replicated sampling using a 3‐mm mesh seine was used to characterize the fish communities. Our results indicate that the habitat mosaic approach creates suitable habitat for a variety of fish species where 37% of fish species were captured in just one habitat type. In particular, the recreationally important Centropomus undecimalis (common snook) was more common in the mangrove‐lined creek and the non‐native Sarotherodon melanotheron (blackchin tilapia) was common in the tidal pond. Greater emphasis should be placed on applied restoration research to identify how habitat types within a larger restoration mosaic contribute to local species diversity and recreationally and commercially important fishes, while limiting non‐natives. This emphasis could reveal how restoration approaches can be modified to include habitat mosaics, maximizing their contribution to productive fish habitat.     

Congruent population structure inferred from dispersal behaviour and intensive genetic surveys of the threatened Florida scrub-jay (Aphelocoma coerulescens)

The delimitation of populations, defined as groups of individuals linked by gene flow, is possible by the analysis of genetic markers and also by spatial models based on dispersal probabilities across a landscape. We combined these two complimentary methods to define the spatial pattern of genetic structure among remaining populations of the threatened Florida scrub-jay, a species for which dispersal ability is unusually well-characterized. The range-wide population was intensively censused in the 1990s, and a metapopulation model defined population boundaries based on predicted dispersal-mediated demographic connectivity. We subjected genotypes from more than 1000 individual jays screened at 20 microsatellite loci to two Bayesian clustering methods. We describe a consensus method for identifying common features across many replicated clustering runs. Ten genetically differentiated groups exist across the present-day range of the Florida scrub-jay. These groups are largely consistent with the dispersal-defined metapopulations, which assume very limited dispersal ability. Some genetic groups comprise more than one metapopulation, likely because these genetically similar metapopulations were sundered only recently by habitat alteration. The combined reconstructions of population structure based on genetics and dispersal-mediated demographic connectivity provide a robust depiction of the current genetic and demographic organization of this species, reflecting past and present levels of dispersal among occupied habitat patches. The differentiation of populations into 10 genetic groups adds urgency to management efforts aimed at preserving what remains of genetic variation in this dwindling species, by maintaining viable populations of all genetically differentiated and geographically isolated populations.     

The applicability of metapopulation theory to large mammals

Metapopulation theory has become a common framework in conservation biology and it is sometimes suggested that a metapopulation approach should be used for management of large mammals. However, it has also been suggested that metapopulation theory would not be applicable to species with long generations compared to those with short ones. In this paper, we review how and on what empirical ground metapopulation terminology has been applied to insects, small mammals and large mammals. The review showed that the metapopulation term sometimes was used for population networks which only fulfilled the broadest possible definition of a metapopulation, i.e. they were subpopulations connected by migrating individuals. We argue that the metapopulation concept should be reserved for networks that also show some kind of metapopulation dynamics. Otherwise it applies to almost all populations and loses its substance. We found much empirical support for metapopulation dynamics in both insects and small mammals, but not in large mammals. A possible reason is the methods used to confirm the existence of metapopulation dynamics. For insects and small mammals, the common approach is to study population turnover through patch occupancy data. Such data is difficult to obtain for large mammals, since longer temporal scales need to be covered to record extinctions and colonizations. Still, many populations of large mammals are exposed to habitat fragmentation and the resulting subpopulations sometimes have high risks of extinction. If there is migration between the subpopulations, the metapopulation framework could provide valuable information on their population dynamics. We suggest that a metapopulation approach can be interesting for populations of large mammals, when there are discrete breeding subpopulations and when these subpopulations have different growth rates and demographic fates. Thus, a comparison of the subpopulations’ demographic fates, rather than subpopulation turnover, can be a feasible alternative for studies of metapopulation dynamics in large mammals.     

Scale dependency of metapopulation models used to predict climate change impacts on small mammals

To investigate potential range shifts in a changing climate it is becoming increasingly common to develop models that account for demographic processes. Metapopulation models incorporate the spatial configuration of occupied habitat (i.e. arrangement, size and quality), population demographics, and inter‐patch dispersal making them suitable for investigating potential threats to small mammal range and abundance. However, the spatial scale (resolution) used to represent species–environment dynamics may affect estimates of range shift and population resilience by failing to realistically represent the spatial configuration of suitable habitat, including stepping stones and refugia. We aimed to determine whether relatively fine‐scale environmental information influenced predictions of metapopulation persistence and range shift. Species distribution models were constructed for four small terrestrial mammals from southern Australia using environmental predictors measured at 0.1 × 0.1 km (0.01 km²) or 1.0 × 1.0 km (1 km²) resolution, and combined with demographic information to parameterise coupled niche‐population models. These models were used to simulate population dynamics projected over 40‐yr under a stable and changing climate. Initial estimates of the area of available habitat were similar at both spatial scales. However, at the fine‐scale, habitat configuration comprised a greater number of patches (ca 12 times), that were more irregular in shape (ca 8 times the perimeter:area), and separated by a tenth of the distance than at the coarse‐scale. While small patches were not more prone to extinction, populations generally declined at a higher rate and were associated with a lower expected minimum abundance. Despite increased species vulnerability at the fine‐scale, greater range shifts were measured at the coarse‐scale (for species illustrating a shift at both scales). These results highlight the potential for range shifts and species vulnerability information to be misrepresented if advanced modelling techniques incorporating species demographics and dispersal inadequately represent the scale at which these processes occur.     

Comparison of reintroduction and enhancement effects on metapopulation viability

Metapopulation viability depends upon a balance of extinction and colonization of local habitats by a species. Mechanisms that can affect this balance include physical characteristics related to natural processes (e.g. succession) as well as anthropogenic actions. Plant restorations can help to produce favorable metapopulation dynamics and consequently increase viability; however, to date no studies confirm this is true. Population viability analysis (PVA) allows for the use of empirical data to generate theoretical future projections in the form of median time to extinction and probability of extinction. In turn, PVAs can inform and aid the development of conservation, recovery, and management plans. Pitcher's thistle (Cirsium pitcheri) is a dune endemic that exhibited metapopulation dynamics. We projected viability of three natural and two restored populations with demographic data spanning 15–23 years to determine the degree the addition of reintroduced population affects metapopulation viability. The models were validated by comparing observed and projected abundances and adjusting parameters associated with demographic and environmental stochasticity to improve model performance. Our chosen model correctly predicted yearly population abundance for 60% of the population‐years. Using that model, 50‐year projections showed that the addition of reintroductions increases metapopulation viability. The reintroduction that simulated population performance in early‐successional habitats had the maximum benefit. In situ enhancements of existing populations proved to be equally effective. This study shows that restorations can facilitate and improve metapopulation viability of species dependent on metapopulation dynamics for survival with long‐term persistence of C. pitcheri in Indiana likely to depend on continued active management.     

Plant Pattern Development during Early Post‐Restoration Succession in Grasslands—A Case Study of Arabis nemorensis

The main objective of this study was to analyze whether plant material transfer is a suitable approach for establishing plant populations with spatial patterns and population structures resembling those of remnant populations. We studied pattern formation and population characteristics in three remnant and two restored populations of the biennial plant species Arabis nemorensis in the upper Rhine valley in southwestern Germany over a period of 2 years. We investigated spatial patterns of seedlings, juveniles, and adults as well as the small‐scale horizontal and vertical distribution of seeds in the soil, followed the fate of individual plants and recorded structural habitat parameters such as vegetation and litter cover. Population dynamics differed between the study sites, but there was no pronounced difference between restored and remnant sites. Seedlings, juveniles, and adults as well as seeds in the seed bank showed aggregated spatial patterns on all study sites, with positive autocorrelation on a scale of 20–60 cm. Within sites, patterns remained approximately stable through time. Restored sites experienced rapid seed bank formation as a result of the restoration measures. Our results suggest that the restoration measures were not only successful in transferring the target species but also triggered rapid formation of spatially structured populations that, years after restoration, closely resembled those of remnant sites.     

Patchy population structure in a short-distance migrant: evidence from genetic and demographic data

Species often occur in subdivided populations as a consequence of spatial heterogeneity of the habitat. To describe the spatial organization of subpopulations, existing theory proposes three main population models: patchy population, metapopulation and isolated populations. These models differ in their predicted levels of connectivity among subpopulations, and in the risk that a subpopulation will go extinct. However, spatially discrete subpopulations are commonly considered to be organized as metapopulations, even though explicit tests of metapopulation assumptions are rare. Here, we test predictions of the three models on the basis of demographic and genetic data, a combined approach so far surprisingly little used in mobile organisms. From 2002 to 2005, we studied nine subpopulations of the wetland-restricted reed bunting ( Emberiza schoeniclus ) in the southeastern part of the Canton Zurich (Switzerland), from which local declines of this species have been reported. Here, wetlands are as small as 2.7 ha and separated through intensively used agricultural landscapes. Demographic data consisted of dispersal of colour-banded individuals among subpopulations, immigration rates and extinction-/recolonization dynamics. Genetic data were based on the distribution of genetic variability and gene flow among subpopulations derived from the analysis of nine microsatellite loci. Both demographic and genetic data revealed that the patchy population model best described the spatial organization of reed bunting subpopulations. High levels of dispersal among subpopulations, high immigration into the patchy population, and genetic admixture suggested little risk of extinction of both subpopulations and the entire patchy population. This study exemplifies the idea that spatially discrete subpopulations may be organized in ways other than a metapopulation, and hence has implications for the conservation of subpopulations and species.     

Landscape-scale response to local habitat restoration in the regal fritillary butterfly (<Emphasis Type="Italic">Speyeria idalia</Emphasis>) (Lepidoptera: Nymphalidae)

The butterfly Speyeria idalia was once a widespread species in the midwestern United States, but declined precipitously during the 1980s. By the early 1990s in NW Indiana and adjacent NE Illinois, it was reduced to a single site where it persisted as a precarious three deme metapopulation managed using prescribed fire. In 1996, a large-scale high diversity restoration was initiated at the site to restore ecological connectivity for grassland and oak barrens communities across the site. Among many restoration goals, we included restoring connectivity for habitat restricted insect communities across the highly fragmented site. In support of that goal, we included over 620 local genotype plant species in the restoration. S. idalia responded positively to the restoration and was the most abundant butterfly across the site in 2014. Ironically, we incorrectly considered the primary hostplant at the site, Viola bicolor, to be non-native when we initiated the restoration and did not include it in our seed mixes. This weedy annual aggressively expanded into the restoration without our assistance. We believe that we transformed the precarious metapopulation into a well-connected patchy population across the restoration. In addition, the butterfly occupied previously unoccupied habitats far beyond the restoration. This suggests that we have created a regional mainland—island metapopulation structure, with the restoration serving as a source of immigrants that sustains colonization rates at or above the rate of local deme extinction across a landscape that currently spans almost 60 km.     

Extinction risks in metapopulations of a beetle inhabiting hollow trees predicted from time series

Ancient and dead trees are declining habitats harbouring many threatened species. These habitats are naturally patchy, and inhabiting species might exhibit metapopulation dynamics at a small spatial scale. In this study, the demography and metapopulation dynamics was analysed for Osmoderma eremita , which is an endangered beetle species associated with tree hollows in Europe. Extinction risks of O. eremita populations were predicted using Monte Carlo simulations based on time series of population assessments. Predicted occurrence patterns were consistent with field observations from an area with many small stands in which the populations are believed to have been more or less isolated from each other during the last 150–200 yr. Population growth was found to be density dependent. Carrying capacity was proportional to the volume of wood mould (i.e. loose material of dead wood in the tree hollows), which varied widely between hollow trees. This generates large differences in local extinction risks between hollow trees. The predicted metapopulation extinction risk was much higher if the habitat dynamics (formation, gradual increase and deterioration of tree hollows) were taken into consideration than in predictions yielded by models in which the amount of wood mould was assumed to be constant over time. Thus, this system has features from both mainland-island metapopulations and habitat-tracking metapopulations, and is rather far from a classic metapopulation. For the long-term persistence of the species in hollow trees, the habitat dynamics seem to be more important than demographic processes. Since the formation and deterioration of suitable tree are partly stochastic processes, there is a considerable extinction risk for many O. eremita populations, because they mainly rely on only one or a few trees with large amounts of wood mould.     

Rapid development of a restored oyster reef facilitates habitat provision for estuarine fauna

Oyster reef restoration has become a principal strategy for ameliorating the loss of natural Crassostrea virginica populations and increasing habitat provision. In 2014, a large‐scale, high‐relief, 23‐ha subtidal C. virginica reef was restored at the historically productive Half Moon Reef in Matagorda Bay, TX, using concrete and limestone substrates. Encrusting and motile fauna were sampled seasonally until 17 months postrestoration at the restored reef and at adjacent unrestored sites. Restored oysters developed rapidly and were most abundant 3 months postrestoration, with subsequent declines possibly due to interacting effects of larval settlement success on new substrate versus post‐settlement mortality due to competitors and predators. Oyster densities were 2× higher than in a restored oyster population in Chesapeake Bay that was reported to be the largest reestablished metapopulation of native oysters in the world. Resident fauna on the restored reef were 62% more diverse, had 433% greater biomass, and comprised a distinct faunal community compared to unrestored sites. The presence of three‐dimensional habitat was the most important factor determining resident faunal community composition, indicating that substrate limitation is a major hindrance for oyster reef community success in Texas and other parts of the Gulf of Mexico. There were only minor differences in density, biomass, and diversity of associated fauna located adjacent (13 m) versus distant (150 m) to the restored reef. The two substrate types compared had little influence on oyster recruitment or faunal habitat provision. Results support the use of reef restoration as a productive means to rebuild habitat and facilitate faunal enhancement.     

Long‐term metapopulation study of the Glanville fritillary butterfly (Melitaea cinxia): survey methods,data management,and long‐term population trends

Long‐term observational studies conducted at large (regional) spatial scales contribute to better understanding of landscape effects on population and evolutionary dynamics, including the conditions that affect long‐term viability of species, but large‐scale studies are expensive and logistically challenging to keep running for a long time. Here, we describe the long‐term metapopulation study of the Glanville fritillary butterfly (Melitaea cinxia) that has been conducted since 1991 in a large network of 4000 habitat patches (dry meadows) within a study area of 50 by 70 km in the Åland Islands in Finland. We explain how the landscape structure has been described, including definition, delimitation, and mapping of the habitat patches; methods of field survey, including the logistics, cost, and reliability of the survey; and data management using the EarthCape biodiversity platform. We describe the long‐term metapopulation dynamics of the Glanville fritillary based on the survey. There has been no long‐term change in the overall size of the metapopulation, but the level of spatial synchrony and hence the amplitude of fluctuations in year‐to‐year metapopulation dynamics have increased over the years, possibly due to increasing frequency of exceptional weather conditions. We discuss the added value of large‐scale and long‐term population studies, but also emphasize the need to integrate more targeted experimental studies in the context of long‐term observational studies. For instance, in the case of the Glanville fritillary project, the long‐term study has produced an opportunity to sample individuals for experiments from local populations with a known demographic history. These studies have demonstrated striking differences in dispersal rate and other life‐history traits of individuals from newly established local populations (the offspring of colonizers) versus individuals from old, established local populations. The long‐term observational study has stimulated the development of metapopulation models and provided an opportunity to test model predictions. This combination of empirical studies and modeling has facilitated the study of key phenomena in spatial dynamics, such as extinction threshold and extinction debt.     

Genetic spatial structure in a butterfly metapopulation correlates better with past than present demographic structure

The Glanville fritillary butterfly ( Melitaea cinxia ) has been studied in the Åland Islands in Finland since 1991, where it occurs as a classic metapopulation in a large network of 4000 dry meadows. Much ecological work has been conducted on this species, but population genetic studies have been hampered by paucity of suitable genetic markers. Here, using single nucleotide polymorphisms and microsatellites developed for the Glanville fritillary, we examine the correspondence between the demographic and genetic spatial structures. Given the dynamic nature of the metapopulation, the current genetic spatial structure may bear a signal of past changes in population sizes and past patterns of gene flow rather than reflect the current demographic structure or landscape structure. We analyse this question with demographic data for 10 years, using the Rand index to assess the similarity between the genetic, demographic, and landscape spatial structures. Our results show that the current genetic spatial structure is better explained by the past rather than by the current demographic spatial structure or by the spatial configuration of the habitat in the landscape. Furthermore, current genetic diversity is significantly explained by past metapopulation sizes. The time lag between major demographic events and change in the genetic spatial structure and diversity has implications for the study of spatial dynamics.