A metapopulation perspective for salmon and other anadromous fish

Salmonids are an important component of biodiversity, culture and economy in several regions, particularly the North Pacific Rim. Given this importance, they have been intensively studied for about a century, and the pioneering scientists recognized the critical link between population structure and conservation. Spatial structure is indeed of prime importance for salmon conservation and management. At first glance, the essence of the metapopulation concept, i.e. a population of populations, widely used on other organisms like butterflies, seems to be particularly relevant to salmon, and more generally to anadromous fish. Nevertheless, the concept is rarely used, and barely tested. Here, we present a metapopulation perspective for anadromous fish, assessing in terms of processes rather than of patterns the set of necessary conditions for metapopulation dynamics to exist. Salmon, and particularly sockeye salmon in Alaska, are used as an illustrative case study. A review of life history traits indicates that the three basic conditions are likely to be fulfilled by anadromous salmon: (i) the spawning habitat is discrete and populations are spatially separated by unsuitable habitat; (ii) some asynchrony is present in the dynamics of more or less distant populations and (iii) dispersal links populations because some salmon stray from their natal population. The implications of some peculiarities of salmon life history traits, unusual in classical metapopulations, are also discussed. Deeper understanding of the population structure of anadromous fish will be advanced by future studies on specific topics: (i) criteria must be defined for the delineation of suitable habitats that are based on features of the biotope and not on the presence of fish; (ii) the collection of long‐term data and the development of improved methods to determine age structure are essential for correctly estimating levels of asynchrony between populations and (iii) several key aspects of dispersal are still poorly understood and need to be examined in detail: the spatial and temporal scales of dispersal movements, the origin and destination populations instead of simple straying rates, and the relative reproductive success of immigrants and residents.     

Resource grain scales mobility and adult morphology in butterflies

Relations between species mobility and life history traits and/or landscape and habitat features are of broad interest to ecologists and conservation biologists. Here we investigated the reliability of the relations between mobility and (1) resource grain and (2) morphological traits in butterflies. Results were used to assess the biological realism of morphological traits associated with flight as mobility proxies. We then investigated how biological, environmental and landscape variables affected these mobility proxies. We used a multi-species approach on two different sites. Morphological traits were measured on ca. 20 individuals per site, species and sex. Resource distribution was carefully monitored by investigating the spatial distribution and overlap of larval and adult feeding resources, together representing the resource grain. The spatial extent of individual station keeping movements was estimated from distances recorded between successive recaptures of individuals from mark-release-recapture experiments. Morphological traits seemed reliable proxies of mobility, as both variables were strongly correlated. Morphological variations were related to flight type and spatial dimension of nectar resource. The most striking point was the clear relation between the index of relative investment in mobility versus fecundity in females with the spatial dimension of adult feeding resource. Given the generally accepted relation between abdomen volume and female fecundity, this suggests that females might invest more in fecundity when nectar resources are widespread. Finally, we did not detected effects of landscape structure on mobility, which indicates that functional grain of resources is more likely to influence mobility and evolution of morphology in butterflies than landscape connectivity.     

Using animal movement behavior to categorize land cover and predict consequences for connectivity and patch residence times

Context

Landscape-scale population dynamics are driven in part by movement within and dispersal among habitat patches. Predicting these processes requires information about how movement behavior varies among land cover types.

Objectives

We investigated how butterfly movement in a heterogeneous landscape varies within and between habitat and matrix land cover types, and the implications of these differences for within-patch residence times and among-patch connectivity.

Methods

We empirically measured movement behavior in the Baltimore checkerspot butterfly (Euphydryas phaeton) in three land cover classes that broadly constitute habitat and two classes that constitute matrix. We also measured habitat preference at boundaries. We predicted patch residence times and interpatch dispersal using movement parameters estimated separately for each habitat and matrix land cover subclass (5 categories), or for combined habitat and combined matrix land cover classes (2 categories). We evaluated the effects of including edge behavior on all metrics.

Results

Overall, movement was slower within habitat land cover types, and faster in matrix cover types. Butterflies at forest edges were biased to remain in open areas, and connectivity and patch residence times were most affected by behavior at structural edges. Differences in movement between matrix subclasses had a greater effect on predictions about connectivity than differences between habitat subclasses. Differences in movement among habitat subclasses had a greater effect on residence times.

Conclusions

Our findings highlight the importance of careful classification of movement and land cover in heterogeneous landscapes, and reveal how subtle differences in behavioral responses to land cover can affect landscape-scale outcomes.

     

Metapopulation dynamics of the bog fritillary butterfly: comparison of demographic parameters and dispersal between a continuous and a highly fragmented landscape

We investigated the effects of habitat loss and fragmentation on population functioning. We compared demography (daily and total population sizes) and dispersal (dispersal rate and dispersal kernels) of the bog fritillary butterfly in two 6-km² landscapes differing in their degree of fragmentation. In 2000, we conducted a Capture-Mark-Recapture experiment in a highly fragmented system in the marginal part of the species distribution (Belgium) and in a more continuous system in the central part of its distribution (Finland). A total of 293 and 947 butterflies were marked with 286 and 190 recapture events recorded in the fragmented and the continuous system respectively. Our results suggest that habitat loss and fragmentation affect dispersal more than demography. Although density was lower in the continuous system, it remains in the yearly range of variation observed on 10 generations in the fragmented system. However, in the fragmented system, the dispersal rate dropped drastically (39 vs. 64%) and females moved longer distances. Patch area had a significant effect on migration in the fragmented system only. From our results, we propose the definition of a new parameter, the minimal patch area (MPA) needed to establish a local population in highly fragmented landscapes.This revised version was published online in May 2005 with corrections to the Cover Date.     

Quantifying functional connectivity: Experimental evidence for patch-specific resistance in the Natterjack toad (Bufo calamita)

Despite the importance assigned to inter-patch movements in fragmented systems, the structure of landscape between suitable habitat patches, the matrix, is often considered as to be of minor interest, or totally ignored. Consequently, models predicting metapopulation dynamics typically assume that dispersal and movement abilities are independent of the composition of the matrix. The predictions of such models should be invalided if that crucial assumption is unverified. In order to test the hypothesis of a patch-specific resistance, we led an experimental study to assess the matrix effects on the movement ability of juvenile Natterjack toads (Bufo calamita). The movement behaviour of first year toadlets, the dispersal stage in this species, was investigated in an arena experiment. Toadlet mobility was assessed in five landscape components that were mimicked in the lab: sandy soil, road, forest, agricultural field, and pasture. We analysed several movement components including move length, speed, efficiency and turning angle distribution. Our results showed that movement ability was strongly affected by the land cover, even if body size modulated the behavioural responses of toadlets. Performances were the best in the arenas mimicking sand and roads, and the worst in the forest arena, toadlet moves being three to five times less effective in the latter. The mobility was intermediate in the two other arenas. We propose here a new method to quantify functional connectivity, based on quantitative estimates of relative values for resistance of landscape components. This method offers a reliable alternative for resistance value estimates to subjective expert advice or inference from genetic population structure.     

Quantifying functional connectivity: Experimental evidence for patch-specific resistance in the Natterjack toad (Bufo calamita)

Despite the importance assigned to inter-patch movements in fragmented systems, the structure of landscape between suitable habitat patches, the matrix, is often considered as to be of minor interest, or totally ignored. Consequently, models predicting metapopulation dynamics typically assume that dispersal and movement abilities are independent of the composition of the matrix. The predictions of such models should be invalided if that crucial assumption is unverified. In order to test the hypothesis of a patch-specific resistance, we led an experimental study to assess the matrix effects on the movement ability of juvenile Natterjack toads (Bufo calamita). The movement behaviour of first year toadlets, the dispersal stage in this species, was investigated in an arena experiment. Toadlet mobility was assessed in five landscape components that were mimicked in the lab: sandy soil, road, forest, agricultural field, and pasture. We analysed several movement components including move length, speed, efficiency and turning angle distribution. Our results showed that movement ability was strongly affected by the land cover, even if body size modulated the behavioural responses of toadlets. Performances were the best in the arenas mimicking sand and roads, and the worst in the forest arena, toadlet moves being three to five times less effective in the latter. The mobility was intermediate in the two other arenas. We propose here a new method to quantify functional connectivity, based on quantitative estimates of relative values for resistance of landscape components. This method offers a reliable alternative for resistance value estimates to subjective expert advice or inference from genetic population structure.     

Metapopulation dynamics and conservation of the marsh fritillary butterfly: Population viability analysis and management options for a critically endangered species in Western Europe

This study investigates the dynamics and viability of a marsh fritillary butterfly Euphydryas aurinia metapopulation in a Belgian successional landscape. Based on capture-mark-recapture and winter nest census data, we first estimated demography (survival and recruitment rates, population size, density dependence) and dispersal parameters (emigration rate, effect of patch connectivity on dispersal, mortality during dispersal). Then using RAMAS/GIS platform, we parameterised a population viability analysis (PVA) model with these parameters to simulate the future of this metapopulation under different scenarios.The metapopulation does not seem viable even if natural reforestation is controlled by adequate management. In its present state, the patch system is not able to sustain enough individuals: due to the large temporal fluctuations in demographic parameters, a carrying capacity far higher than currently would be necessary to limit extinction risk to 1%, suggesting the existence of an extinction debt for the species in Belgium. The situation of E. aurinia appears much worse compared to two other fritillary species threatened in Belgium, for which similar PVA are available. It is therefore urgent to increase the carrying capacity of the patch system. How and where it is achieved are of secondary importance for the gain in viability: improvement of habitat quality through restoration, or increase of habitat quantity via enlargement of existing patches and/or creation of new habitat in the matrix. A regime of management based on regular re-opening and maintenance of habitat patches may be the only guarantee of long-term persistence for this critically endangered species in Belgium.