Metapopulation dynamics: brief history and conceptual domain

We review the early development of metapopulation ideas, which culminated in the well-known model by Levins in 1969. We present a survey of metapopulation terminology and outline the kinds of studies that have been conducted on single-species and multispecies metapopulations. Metapopulation studies have important conceptual links with the equilibrium theory of island biogeography and with studies on the dynamics of species living in patchy environments. Metapopulation ideas play an increasingly important role in landscape ecology and conservation biology.     

Finite metapopulation models with density-dependent migration and stochastic local dynamics

The effects of small density-dependent migration on the dynamics of a metapopulation are studied in a model with stochastic local dynamics. We use a diffusion approximation to study how changes in the migration rate and habitat occupancy affect the rates of local colonization and extinction. If the emigration rate increases or if the immigration rate decreases with local population size, a positive expected rate of change in habitat occupancy is found for a greater range of habitat occupancies than when the migration is density-independent. In contrast, the reverse patterns of density dependence in respective emigration and immigration reduce the range of habitat occupancies where the metapopulation will be viable. This occurs because density-dependent migration strongly influences both the establishment and rescue effects in the local dynamics of metapopulations.     

The role of setts in badger (Meles meles) group size, breeding success and status of TB (Mycobacterium bovis)

This paper examines the relationship between the number of occupied setts in a badger social group territory and badger group size, breeding success, and status of infection with Mycobacterium bovis (TB). The data used were from a long-term epidemiological and ecological study of a high-density population of badgers Meles meles in south-west England. The number of occupied setts in a social group was significantly and positively related to the number of badgers caught in the social group, so that as a social group increases in size, badgers occupy more of the available setts. This relationship remained significant when numbers of adults, adult males and adult females were examined. The number of breeding females, number of cubs and sex ratio was not related to the number of occupied setts in a social group. It is possible that the advantages to breeding females of a larger number of setts available to breed in might be outweighed by the increased aggression found in larger groups. The TB score for prevalence and for incidence of social groups was significantly and positively related to the number of occupied setts in a social group, such that the more occupied setts there were in a territory, the higher the TB index of the group. Possibly the setts themselves contribute to the persistence of TB within social groups, or badgers infected with TB might show a difference in behaviour from uninfected badgers resulting in their increased use of outlying setts.     

Interplay between local dynamics and dispersal in discrete-time metapopulation models

The effects of synchronous dispersal on discrete-time metapopulation dynamics with local (patch) dynamics of the same (compensatory or overcompensatory) or mixed (compensatory and overcompensatory) types are explored. Single-species metapopulation models behave as single-species single-patch models, whenever all local patches are governed by compensatory dynamics. Dispersal gives rise to multiple attractors with complex basin structures, whenever some local patches are under overcompensatory dynamics. In mixed systems, dispersal is capable of altering the local dynamics from compensatory to overcompensatory dynamics and vice versa. Examples are provided of metapopulation models supporting multiple attractors with intermingled basins of attraction.     

Testing metapopulation dynamics using genetic, demographic and ecological data

The metapopulation concept is a cornerstone in the recent history of ecology and evolution. However, determining whether a natural system fits a metapopulation model is a complex issue. Extinction-colonization dynamics are indeed often difficult to quantify because species detectability is not always 100%, resulting in an imperfect record of extinctions. Here, we explore whether combining population genetics with demographic and ecological surveys can yield more realistic estimates of metapopulation dynamics. We apply this approach to the freshwater snail Drepanotrema depressissimum in a fragmented landscape of tropical ponds. In addition to studying correlations between genetic diversity and demographical or ecological characteristics, we undertake, for the first time, a detailed search for genetic signatures of extinction-recolonization events using temporal changes in allele frequencies within sites. Surprisingly, genetic data indicate that extinction is much rarer than suggested by demographic surveys. Consequently, this system is better described as a set of populations with different sizes and immigration rates than as a true metapopulation. We identify several cases of apparent extinction owing to nondetection of low-density populations, and of aestivating individuals in desiccated ponds. More generally, we observed a frequent mismatch between genetic and demographical/ecological information at small spatial and temporal scales. We discuss the causes of these discrepancies and show how these two types of data provide complementary information on population dynamics and history, especially when temporal genetic samples are available.     

Colonization in metapopulations: a review of theory and observations

In metapopulation dynamics turnover of populations in isolated patches may be frequent. Regional survival of a species in such a system with frequent extinctions hinges on its colonization ability. Colonization is more than just dispersal; when a propagule reaches a new patch it faces higher extinction probabilities than does an established population. Extinction models as well as empirical data suggest that a large propagule with a potential for rapid increase in a varying environment, or with a low mortality rate in an environment perceived as constant, has a higher probability of successful colonization. Large variation in population size when it is still small increases the risk of failure. Factors introducing such variation are demographic stochasticity and environmental variation. It is hard to single out demographic traits that ensure good colonizing ability, since colonization can be achieved in many different ways, but generalists and species with self-fertilization seem to be superior.     

Reproductive skew and relatedness in social groups of European badgers, Meles meles

Reproductive skew is a measure of the proportion of individuals of each sex that breed in a group and is a valuable measure for understanding the evolution and maintenance of sociality. Here, we provide the first quantification of reproductive skew within social groups of European badgers Meles meles , throughout an 18-year study in a high-density population. We used 22 microsatellite loci to analyse within-group relatedness and demonstrated that badger groups contained relatives. The average within-group relatedness was high ( R =  0.20) and approximately one-third of within-group dyads were more likely to represent first-order kin than unrelated pairs. Adult females within groups had higher pairwise relatedness than adult males, due to the high frequency of extra-group paternities, rather than permanent physical dispersal. Spatial clustering of relatives occurred among neighbouring groups, which we suggest was due to the majority of extra-group paternities being attributable to neighbouring males. Reproductive skew was found among within-group candidate fathers ( B  = 0.26) and candidate mothers ( B  = 0.07), but not among breeding individuals; our power to detect skew in the latter was low. We use these results to evaluate reproductive skew models. Although badger society best fits the assumptions of the incomplete-control models, our results were not consistent with their predictions. We suggest that this may be due to female control of paternity, female–female reproductive suppression occurring only in years with high food availability resulting in competition over access to breeding sites, extra-group paternity masking the benefits of natal philopatry, and/or the inconsistent occurrence of hierarchies that are linear when established.     

Dispersal, distribution, patch network and metapopulation dynamics of the dingy skipper butterfly (Erynnis tages)

Two general approaches have usually been taken towards understanding the distributions and dynamics of localised species in heterogeneous landscapes, namely habitat characterisation and metapopulation dynamics. We show how habitat and metapopulation dynamics interact to generate a highly localised distribution of a butterfly, despite the extremely widespread nature of the butterfly’s host plant. Egg placement, macro-habitat requirements and dispersal were studied for the butterfly Erynnis tages, in North Wales, where it shows a restricted distribution relative to that of its host plant, Lotus corniculatus. Females laid eggs disproportionately on large plants growing in hollows, with intermediate cover of bare ground and high cover of L. corniculatus. Ideal macro-habitat, studied at 100-m grid resolution, consisted of areas with high host plant densities, sheltered from wind, with light or no grazing or cutting. These specialised conditions are represented as localised patches in the landscape, and define the potential habitat network, within which metapopulation dynamics take place. Although there was a moderate (22%) level of exchange of individual E. tages among local populations, the total number of potential colonists in the whole system was low because source population sizes were small (≤200 individuals at peak in any site in 1997 and 1998). Four unoccupied but apparently suitable 500-m grid squares were colonised between 1997 and 1998, and isolated habitat was less likely to be occupied. Overall, our study suggests that long-term regional persistence of E. tages is very likely to depend on metapopulation processes within the restricted patch network, rather than on the long-term survival of local populations. Received: 25 May 1999 / Accepted: 9 August 1999     

Avian life-history determinants of local extinction risk in a hyper-fragmented neotropical forest landscape

The fact that species vary in their vulnerability to extinction is well documented, but the reasons for these differences remain poorly understood. Why should some species/families/guilds decline rapidly with increasing anthropogenic disturbance, while others either tolerate or proliferate in disturbed habitats? We investigated the bird species composition in 31 primary forest patches of varying size in a region of the Amazonian 'Arc of Deforestation' and assessed which species life-history traits predisposed individual species to extinction. Medium-sized non-flocking canopy frugivores/ominvores of low primary forest dependence were least likely to go extinct in small patches, while small-bodied flock-following primary-forest-dependent terrestrial insectivores were most fragmentation sensitive. We found highly idiosyncratic relationships between the minimum size of forest patches occupied by different species and their territory size requirements estimated based on other Amazonian studies. This suggests that avian assemblages in forest fragments primarily comprise species that either have good dispersal abilities or are highly tolerant to the non-forest matrix, rather than those whose minimum spatial requirements can be met by the size of available forest fragments.     

Consequences of forest fragmentation for the dynamics of bird populations: conceptual issues and the evidence

This paper reviews the consequences of forest fragmentation for the dynamics of bird populations. Owing to high mobility and large home ranges, birds usually perceive fragmented forests in a finegrained manner, i.e. embrace several forest fragments in functional home ranges. On a regional scale, however, coarse-grained clusters of fine-grained fragments (hierarchical fragmentation may sub-divide bird populations into isolated demes, which enter a domain of metapopulation dynamics. Distinctions are made between pure distance-area or population-level effects and more indirect community-level effects due to changes in landscape composition. Distance-area effects, such as insularization and decreasing fragment size, directly prevent dispersal and reduce population size. Landscape effects, such as reduced fragment-matrix and interior-edge ratios, increase the pressure from surrounding predators, competitors, parasites and disease. In short, forest fragmentation can be viewed as a two-step process. Initially, fine-grained fragmentation triggers distance-area and landscape effects on a local scale, which in turn, results in a range retraction of a population to non-fragmented or less fragmented parts of a region. At a certain point, non-fragmented areas become so widely spaced out that regional distance-area effects come into operation, giving rise to metapopulation dynamics. Although few bird metapopulations have yet been documented, metapopulation dynamics probably is a common characteristic of bird populations confined to 'hierarchical' fragmented forests.     

The strain-specific dynamics of Escherichia coli O157:H7 faecal shedding in cattle post inoculation

This study reports analysis of faecal shedding dynamics in cattle for three Escherichia coli O157:H7 (ECO157) strains (S1, S2 and S3) of different genotype and ecological history, using experimental inoculation data. The three strains were compared for their shedding frequency and level of ECO157 in faeces. A multistate Markov chain model was used to compare shedding patterns of S1 and S2. Strains S1 and S2 were detected seven to eight times more often and at 10⁴ larger levels than strain S3. Strains S1 and S2 had similar frequencies and levels of shedding. However, the total time spent in the shedding state during colonization was on average four times longer for S1 (15 days) compared to S2 (4 days). These results indicate that an ECO157 strain effect on the frequency, level, pattern and the duration of faecal shedding may need to be considered in control of ECO157 in the cattle reservoir.     

Relative contributions of local and regional factors to species richness and total density of butterflies and moths in semi-natural grasslands

Metapopulation theory predicts that species richness and total population density of habitat specialists increase with increasing area and regional connectivity of the habitat. To test these predictions, we examined the relative contributions of habitat patch area, connectivity of the regional habitat network and local habitat quality to species richness and total density of butterflies and day-active moths inhabiting semi-natural grasslands. We studied butterflies and moths in 48 replicate landscapes situated in southwest Finland, including a focal patch and the surrounding network of other semi-natural grasslands within a radius of 1.5 km from the focal patch. By applying the method of hierarchical partitioning, which can distinguish between independent and joint contributions of individual explanatory variables, we observed that variables of the local habitat quality (e.g. mean vegetation height and nectar plant abundance) generally showed the highest independent effect on species richness and total density of butterflies and moths. Habitat area did not show a significant independent contribution to species richness and total density of butterflies and moths. The effect of habitat connectivity was observed only for total density of the declining butterflies and moths. These observations indicate that the local habitat quality is of foremost importance in explaining variation in species richness and total density of butterflies and moths. In addition, declining butterflies and moths have larger populations in well-connected networks of semi-natural grasslands. Our results suggest that, while it is crucial to maintain high-quality habitats by management, with limited resources it would be appropriate to concentrate grassland management and restoration to areas with well-connected grassland networks in which the declining species currently have their strongest populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.