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.     

Grazing promotes dung beetle diversity in the xeric landscape of a Mexican Biosphere Reserve

We analyze the impact of grazing on dung beetle diversity at the Barranca de Metztitlán Biosphere Reserve, a xeric ecosystem in central Mexico with a long history of use by humans. We compared the community structure, as well as the alpha and beta diversity between two cover conditions (open and closed vegetation) that represent the impact of grazing within a habitat, and between habitat types (submountainous and crassicaule scrublands). From 576 samples we collected 75,605 dung beetles belonging to 20 taxa. While mean species richness and diversity were different between habitat types, cumulative species richness was not. The effects of grazing on vegetation structure influenced the cumulative species richness and diversity of dung beetles in the submountainous scrubland, where grazing has created land mosaics of a grassland matrix with scrubland patches. This was not the case in the crassicaule scrubland where the impact of grazing is not as evident. Beta diversity significantly responds to the effects of grazing on habitat conditions. We discuss the ecological factors that may promote these responses by landscape diversity components. We also identify the species that could act as useful indicators to monitor the effect of land management on biodiversity. Our results indicate cattle farming maintains a diversified land mosaic, and these areas support more diverse dung beetle ensembles than homogeneous areas of closed, shrubby vegetation cover. Thus, controlled grazing activity could certainly favour the conservation of dung beetle biodiversity and improve ecosystem functioning by maintaining dung decomposition rates.     

The relative importance of patch habitat quality and landscape attributes on a declining steppe-bird metapopulation

Metapopulation theory is one of the most popular approaches to identify the factors affecting the spatial and temporal dynamics of populations in fragmented habitat networks. Habitat quality, patch area and isolation are mainly focused on when analyzing distribution patterns in fragmented landscapes. The effects of landscape heterogeneity in the non-occupied matrix, however, have been largely neglected. Here, we determined the relative importance of patch quality and landscape attributes on the occurrence, density and extinction of the Dupont’s lark (Chersophilus duponti), an endangered steppe passerine whose habitat has been extremely reduced to highly isolated and fragmented patches embedded in a mainly unsuitable landscape matrix. Habitat patch quality, measured in terms of vegetation structure, grazing pressure, arthropod availability, predator abundance, and inter-specific competition, did not affect occurrence, density or extinction. At the landscape scale, however, the species’ occurrence was principally determined by the interactions among patch size, geographic isolation and landscape matrix. Isolation had the main independent contribution to explaining the probability of occurrence, followed by landscape matrix composition and patch size. The species’ density was negatively correlated to patch size, suggesting crowding effects in small fragments, while extinction events were exclusively related to isolation. Our findings suggest that landscape rather than local population characteristics are crucial in determining the patterns of distribution and abundance of non-equilibrium populations in highly fragmented habitat networks. Consequently, conservation measures for these species should simultaneously involve patch size, isolation and landscape matrix and apply to the entire metapopulation rather than to particular patches.     

Diversity patterns in butterfly communities of the Greek nature reserve Dadia

Traditionally, nature reserves have been centered mainly around areas that are important for vertebrate diversity. This practice has not gone unchallenged and may be a suboptimal choice for overall conservation planning. To investigate this problem, we sampled butterfly species richness in a nature reserve in north eastern Greece that was originally established for the protection of birds of prey. Patterns of butterfly species richness and abundance were investigated by means of transect walks across a range of the seven predominant habitat types (wet and dry meadow, pine, oak and mixed forest, grazed, and agricultural land). Data analysis, including ANOVA and DCA (detrended correspondence analysis), revealed that the main gradients in butterfly species richness (low to high) were from sites dominated by the pine forest matrix of the core areas of the reserve, to peripheral sites in landscapes of mixed or oak forest, and from sites with little human impact to more disturbed areas with high grazing pressure. Species of conservation interest were concentrated at sites of low human impact. Ten of them are endemic to Europe and/or threatened in Europe. In this respect, the most important species are Lycaena ottomanus, Thymelicus acteon, and Pseudophilotes vicrama which are declining all over Europe. Our results suggest that (1) traditional agricultural practices in areas surrounded by forest can be considered as important management tools in butterfly conservation, (2) highest butterfly species richness is found in the periphery of the reserve rather than in the core areas, and (3) for butterfly conservation the zones surrounding the strictly protected areas are equally important as the core areas.     

How does grazing intensity affect different vegetation types in arid Succulent Karoo, South Africa? Implications for conservation management

The Knersvlakte in the Succulent Karoo Biome (South Africa) is known for its high plant diversity and endemism. In the course of establishing a conservation area there, we assessed baseline data for future management. We investigated the effects of grazing on the vegetation in terms of species diversity and composition as well as reproduction of selected species. Data were sampled on four adjacent farms, which were ungrazed, moderately or intensively grazed by sheep and goats. The data were collected in 27 quartz and 24 non-quartz plots, representing two major habitat types of the region. Within each of the 1000-m² plots, 100 subplots of 400 cm² size were sampled. ANOVAs revealed that species richness and abundance of endemic species on quartz fields decreased with grazing. Abundance of annuals did not increase significantly due to grazing. Fidelity analyses indicated that species composition differed between grazing intensities and that the ungrazed and moderately grazed plots both contained unique locally endemic habitat specialists. Reproduction of two endemic dwarf shrubs Drosanthemumschoenlandianum and Argyrodermafissum (both Aizoaceae) increased under moderate grazing, which in the case of D.schoenlandianum was interpreted as an effect of grazing. We attribute the low number of seedlings and annuals on the moderately grazed farm to lower seasonal rainfall on these plots. From a conservation perspective, no or moderate grazing appear to be necessary to preserve plant diversity and vegetation patterns, and their underlying processes.     

When functional habitat does not match vegetation types: A resource-based approach to map butterfly habitat

The unambiguous recognition of a species’ habitat is a matter of debate. For terrestrial species, habitat is often defined as physical patches of a certain vegetation type in a matrix of non-habitat. Ecological resources that make up the habitat of a species may, however, only cover subsets of vegetation types or can be spatially dispersed in a complex way over different vegetation types. Here we present and test a procedure to recognize and delineate habitat according to a resource-based approach instead of a vegetation-based approach. We used the green hairstreak butterfly (Callophrys rubi) in a heathland landscape as a study case. Our resource-based habitat approach selects those zones that comprise essential resources and conditions within an appropriate spatial window. Variables that were retained in a logistic regression model were used to calculate larval, adult and combined habitat indices in a GIS, taking into account thermal constraints on resource-use, as this is a key habitat aspect for this heliothermous insect. To group different (and sometimes scattered) ecological resources into functional habitat zones, we derived a measure of space-use from mark-release-recapture data. By least-cost modelling this spatial window was adapted to the nature of the vegetation between sets of resources. The habitat zones that were delineated using this approach matched the observed distribution of butterflies significantly better than did a classic approach based on vegetation types with host plants only. Our approach provides concrete output for conservation purposes, like recognizing zones with the highest potential for habitat restoration.     

Habitat use and conservation of bumblebees (Bombus spp.) under different grassland management regimes

Declines in the natural populations of several bumblebee species across Britain and Europe are an increasing cause for concern. In this study the habitat use of bumblebees was investigated on Salisbury Plain Training Area, the largest remaining area of unimproved chalk grassland in north-west Europe. Habitat characteristics influencing the overall abundance, species richness and foraging activity of bumblebees included the diversity and abundance of flowering plant species (particularly of favoured forage plants such as Trifolium pratense), vegetation structure and height. It is suggested that different Bombus species respond to these habitat characteristics depending on their specific foraging and nesting requirements, the case of Bombus humilis being especially relevant. The effects of several grassland management practices were considered in terms of their suitability for the conservation of bumblebee habitats. Cattle grazing was shown to be preferable to both sheep grazing and the absence of any management, although the timing and intensity of such grazing was important. Small-scale disturbances caused by vehicle activity were also of value in producing locally abundant forage resources in less intensively managed grasslands.     

When is a habitat not a habitat? Dramatic resource use changes under differing weather conditions for the butterfly Plebejus argus

A key to conserving organisms is identification of the habitat bounds and essential resources within them. In population studies (metapopulations) of phytophagous arthropods it is tacitly assumed that habitat bounds portray short-term stability and that habitat is largely synonymous with hostplant areas or with a single vegetation unit comprising hostplants; structural components are usually ignored. We test these assumptions by monitoring the behaviour and relative abundance of Plebejus argus (L.) (Lepidoptera: Lycaenidae) in relation to changing weather conditions in two patches of a North Wales metapopulation for the butterfly. Our findings confirm the importance of structural habitat components. P. argus density is higher in the vicinity of shrubs which are used for roosting, resting, basking, mate location and shelter. A dominant proportion of the population adopts shrub areas in cooler, cloudy and windy weather. In warmer, sunnier and calmer conditions, the butterfly spends longer in flight and moves out onto calcareous heath dominated by hostplants. In doing so, an increasing, even dominant, proportion of the population occupies exposed slopes adjacent to and above shrub covered areas associated with the hostplant. In effect, the habitat bounds appear to change with conditions on scales of days and hours. What part of a landscape may be defined as a habitat, and what part of it may appear to be most important for an organism, depends on just when and where surveys are carried out. We argue that for correct delineation of habitats attention needs to be given to resource use in different conditions. In the face of enhanced global warming, a broad view should be taken of arthropod habitats that considers the resources required for varying conditions.     

Habitat requirements of passerines and reedbed management in southern France

Reedbeds have high conservation value in Europe. In southern France, they are the major breeding habitat of five passerine species. Yet, habitat management is done primarily by water control to serve socio-economic rather than conservation interests, because we lack information on the species' ecological requirements. Determinants of passerine abundance were assessed through a comparative analysis of water regime, plant structure, and arthropod (food) distribution at 12 sites consisting of at least 10 ha of marsh densely covered with common reed (Phragmites australis). Overall bird abundance estimated through standardised mist netting was positively correlated with food availability (sweep-netted arthropods weighted by their occurrence in birds' diet), which was in turn negatively correlated with duration of ground dryness between June and December. Abundance of four of the five bird species was associated with specific vegetation parameters (reed diameter, dry reed density, growing reed height, etc.), which could be associated with particular management practices, especially with regard to water levels and salinity. Potential impact of socio-economic activities through their water management is addressed, as well as possible ways to minimise these impacts.     

Functional conservation units for the endangered Alcon Blue butterfly Maculinea alcon in Belgium (Lepidoptera: Lycaenidae)

To organize and prioritise species-specific conservation efforts, we delineate `functional conservation units' for the threatened Alcon Blue butterfly Maculinea alcon in Belgium. We used detailed distribution data on the butterfly, its host plant and its habitat, present-day population sizes and its mobility and colonization capacity to determine functional conservation units (FCUs) on different spatial scales: FCU-1, i.e., the 12 presently occupied habitat patches plus the area within a range of 500 m surrounding them (the maximum local movement distance, based on mark-release-recapture data), FCU-2, i.e., the areas within a range of 2 km around the occupied habitat patches (the maximum observed colonization capacity) and FCU-3, i.e., potential re-introduction sites (sites where M. alcon went extinct recently). We suggest different management and planning measures for each type of functional conservation unit and discuss translocation and re-introduction as `intensive care' conservation measures for this threatened and sedentary species.     

Conservation implications of the distribution of genetic diversity at different scales: a case study using the marsh fritillary butterfly (Euphydryas aurinia)

In the UK, Euphydryas aurinia exists in fragmented habitat patches, and undergoes population fluctuations as a result of a larval parasitoid. Its range is declining in the UK and conservation is thought to require a landscape approach since populations spread over large areas in some years and contract to core breeding patches in others. We examined populations at a range of geographic scales using allozyme electrophoresis to look for evidence of gene flow and differences in genetic diversity among populations. Nationally, our F_ST value was 0.1542 but between population groups within the suspected colonisation range of the butterfly (ca. 20 km), F_ST values were not significantly different from zero. Genetic diversity in terms of number of alleles and heterozygosity was reasonably high in natural populations (H_e=0.267) but low in an introduced, isolated population. We infer that migration between closely spaced subpopulations (in a metapopulation) maintains a high genetic effective population size (large number of individuals in a population that contribute genes to the next generation) which offsets any local reductions in population numbers due to stochastic extinctions or parasitoid effects. We therefore conclude that effective conservation of the species must seek to provide networks of suitable habitat for groups of subpopulations, rather than maintaining habitat for isolated populations.     

Spatial population structure of the Cabrera vole in Mediterranean farmland: The relative role of patch and matrix effects

There is mounting evidence that both patch networks and the intervening matrix influence species persistence in fragmented landscapes, though the relative importance of each of these factors in determining spatial population structure remains poorly understood. This study examined this issue using a three-year data set on the distribution of Cabrera voles (Microtus cabrerae) in Mediterranean farmland. The spatial pattern appeared consistent with a metapopulation structure, as voles occupied discrete tall herb patches scattered across the agricultural landscape, where local extinctions and colonizations induced temporal changes in occupancy patterns. Patch dynamics determined deviations from classical metapopulation assumptions, with over half the extinctions resulting from agricultural disturbance or vegetation succession, and recolonizations often occurring after the recovery of suitable habitat conditions sometime after disturbance. Occupancy in undisturbed patches was more stable, with vole occurrence in one year strongly reflecting that in the previous year. Overall, occupancy increased with both patch size and connectivity, but the unique contribution of patch variables to explain variation in vole occurrence was far smaller than that of matrix attributes. Voles occurred more often in patches surrounded by natural pastures, while prevalence declined with increasing cover by shrubland, pine plantations, improved pastures and grazed cropland. It is hypothesised that unfavourable land uses may increase the effective isolation of habitat patches through increased predation risk of dispersing voles. Conservation of the Cabrera vole in Mediterranean farmland should thus strive to maintain lightly grazed fields surrounding well-connected networks of suitable habitat patches.     

Landscape occupancy and local population size depends on host plant distribution in the butterfly Cupido minimus

We tested whether the landscape occupancy and local population size of the monophagous butterfly Cupido minimus can be predicted by patch size and isolation of its host plant or by other habitat characteristics. C. minimus and its larval food plant Anthyllis vulneraria are classified as rare and endangered in northern Germany. Adults of C. minimus are ranked as the most sedentary butterfly species in northern Europe.Around the city of Göttingen (Germany), we checked all known locations of A. vulneraria (n=70) in June 2002 for butterfly eggs (in blooming flowerheads) and adult butterflies (within 20-min transects).We found eggs of C. minimus or a high number of adults (>7) in all habitats with A. vulneraria (which are calcareous grasslands) even when isolated up to 2-4 km. In multiple regression analyses, local population size of adult butterflies was positively related to the cover of its larval food plant A. vulneraria explaining 65% of variance. Cover of A. vulneraria increased with increasing habitat area and increasing cover of plant species in flower and decreased with increasing cover of shrub layer. Habitat isolation and further factors describing habitat quality were not related to C. minimus population size or cover of its larval food plant.The results suggest that dispersal ability of C. minimus is greater than expected and that management should focus to increase A. vulneraria patches. For conservation, low impact grazing once a year and removing of excessive shrubs in winter seems to be the most appropriate strategies.     

Habitat selection by the pale-headed brush-finch (Atlapetes pallidiceps) in southern Ecuador: implications for conservation

The pale-headed brush-finch (Atlapetes pallidiceps) is threatened with extinction due to habitat loss, but very little is known about its ecological requirements. We used multiple logistic regression to study habitat selection of this species at landscape, territory, and nest site scales in order to make recommendations about effective management. Habitat selection by the sympatric stripe-headed brush-finch (Buarremon torquatus) was examined with the same methods in order to analyse interspecific resource partitioning and potential competition. The pale-headed brush-finch selected semi-open habitat types with intermediate scrub heights, and avoided forests. Nest sites depended on the presence of vines or bamboo. By contrast, the stripe-headed brush-finch chose dense habitat with low ground cover under tall vegetation and avoided semi-open habitat. The two species had overlapping territories but differed significantly in microhabitat use and the use of vegetation strata. We found no convincing evidence that the stripe-headed brush-finch displaces the pale-headed brush-finch from optimal habitat. The preservation of semi-open scrubland maintained by low-intensity grazing is suggested for future conservation of the pale-headed brush-finch.     

Assessing habitat quality for butterflies on intensively managed arable farmland

The conservation and management of biodiversity requires accurate, repeatable and cost-effective monitoring techniques. In this study, a simple and rapid methodology was employed to measure the quality of different habitats for butterfly species on 10 arable farms in lowland Britain. This habitat monitoring was seven times more rapid than the traditional species-based butterfly monitoring. Data on butterfly abundance were combined with the information gathered on habitat quality at five of the farms using stepwise regression. These models had a consistently high degree of explanatory power for the summary variables of total butterfly abundance and species richness, and the functional groupings of mobile and immobile species. There was good agreement between observed and predicted estimates of species richness and abundance when the models were validated on a further five independent sites. Models to predict the abundance of 11 common butterfly species were also produced. These varied considerably in their predictive power and validity between species. The results clearly demonstrated the beneficial effects of targeted habitat creation for butterflies which is promoted under the Agri-environment Schemes, especially the recently introduced pollen and nectar seed mixtures (WM2) available under the Countryside Stewardship Scheme. The models confirmed the importance of shelter, floristically diverse field margins and the availability of nectar resources from legume species in explaining the abundance of immobile butterfly species in arable landscapes. These sources of nectar and the presence of larval host plants in the crucifer family were important factors in explaining the abundance of mobile species. The results are discussed in terms of the potential of this approach for: (i) assessing habitat quality for butterflies in intensively managed landscapes; (ii) enhancing the value of butterfly monitoring schemes in explaining changes in butterfly abundance at the site and national scale; and (iii) informing habitat management and restoration guidelines for butterfly conservation on arable farmland.     

Higher breeding densities of the threatened little bustard Tetrax tetrax occur in larger grassland fields: Implications for conservation

Previous studies have found that densities of little bustard Tetrax tetrax breeding males tend to be higher in areas with smaller agricultural fields, presumably due to increased habitat diversity. However, exceptionally high densities have been found in large grassland fields in Portugal, which suggests that the influence of field size varies geographically, and that the role of this factor is not yet fully understood, despite its importance as a key management issue.We studied how field size, together with vegetation structure, influences the presence and density of breeding little bustards in a region of southern Portugal. Fifty-four grassland fields were sampled in 2007 and another 29 in 2008, with sizes ranging from 23 to 172 ha. A total of 183 breeding males were found in 47 of these fields, reaching densities of up to 37 males/100 ha. A higher probability of occurrence of breeding males was found in larger fields with a vegetation height below 40 cm and field size alone explained 46% of the variability in male density. These results suggest that larger continuous areas of suitable habitat attract many males, most likely as a consequence of their lek mating system. We conclude that conservation efforts, in a landscape context of large farm sizes, should: (1) be channelled to farms with large fields; (2) ensure adequate livestock grazing to create suitable habitat and (3) promote management at a landscape level to ensure the most continuous grassland habitat patches possible.     

Habitat modelling for the conservation of the rare ground beetle species Carabus variolosus (Coleoptera, Carabidae) in the riparian zones of headwaters

Carabus variolosus is a highly endangered insect which is listed in the EC Habitats and Species Directive. Detailed knowledge of the habitat requirements of this semi-aquatic woodland carabid beetle is needed if effective conservation and management strategies are to be developed. Previous habitat models have proved to be a successful analytical and predictive tool for the conservation of species. We conducted an intensive study over a two year period on two extant C. variolosus populations in Westphalia (Germany) using live capture pitfall traps. We analysed the distribution of the beetles over their main activity period in relation to a large number of edaphic and vegetation variables in order to estimate habitat suitability models and describe optimum ranges. C. variolosus is restricted to the fringes of water bodies and to areas of high soil moisture that display patches of bare soil, shows a slight preference for sparse tree vegetation, and avoids acidic soil. Temporal and spatial transferability of the statistically significant habitat models indicates their robustness and validity. Based on the results of our study, we suggest management measures for the conservation of C. variolosus, which promote the rehabilitation of natural flood plains of headwater areas.     

The influence of livestock grazing and weed invasion on habitat use by birds in grassy woodland remnants

Remnants of native vegetation in regions dominated by agriculture are subject to degradation, especially by livestock grazing and weed invasion. Ground-foraging birds are amongst the most threatened bird groups in Australia, and these agents of degradation might be contributing to their decline by causing a reduction in food availability. We studied the foraging behaviour and microhabitat use of seven species of ground-foraging insectivores in south-eastern Australian buloke woodland remnants with native, grazed and weedy ground-layers. If birds must resort to using more energetically expensive prey-attack manoeuvres, or selectively use substrates and microhabitats that are less available in degraded habitats, then such degradation is likely to be negatively impacting on these species. We found evidence of a negative impact of one or both of these types of degradation on five of the seven bird species. Three species that employ a range of foraging manoeuvres to attack prey used potentially more energetically expensive aerial manoeuvres significantly more frequently in weedy remnants than in remnants with a native or grazed ground layer. Red-capped robins Petroica goodenovii and brown treecreepers Climacteris picumnus both selectively foraged near trees in grazed sites, and hooded robins Melanodryas cucullata, red-capped robins and willie wagtails Rhipidura leucophrys avoided foraging in microhabitats with a high percentage cover of exotic grasses in weedy sites. Brown treecreepers were also less likely to be present in weedy sites that had been protected from grazing than in either grazed or native sites. These results suggest that although grazing appears to have a detrimental impact on foraging habitat of ground-foraging birds, the exclusion of livestock grazing from previously disturbed buloke remnants alone is not adequate to restore habitat values for ground-foraging birds. A conservation strategy for this habitat type should consider the exclusion of heavy grazing from sites with an intact cryptogamic crust and the management of weeds in disturbed remnants, potentially through the use of carefully controlled light grazing.     

Effects of oak barrens habitat management for Karner blue butterfly (Lycaeides samuelis) on the avian community

The federally endangered Karner blue butterfly (Lycaeides samuelis) is the focal species for a conservation plan designed to create and maintain barrens habitats. We investigated whether habitat management for Karner blue butterflies influences avian community structure at Fort McCoy Military Installation in Wisconsin, USA. From 2007 through 2009 breeding bird point count and habitat characteristic data were collected at 186 sample points in five habitat types including two remnant barrens types, barrens habitat restored from woodland and managed specifically for the Karner blue butterfly, and two woodland habitat types. Although the bird community of managed barrens was not identical to the communities of remnant barrens, the Field Sparrow (Spizella pusilla), a species of conservation concern, and sparse canopy associated bird species, such as the Baltimore Oriole (Icterus galbula) and Eastern Bluebird (Sialia sialis) were predicted to occupy managed barrens and remnant barrens in similar proportions. Adjacent habitat was the most influential factor in determining the community of bird species using the managed barrens. In Wisconsin, and likely throughout the range of the Karner blue butterfly, management for the butterfly creates habitat that attracts a bird community similar to that of remnant barrens, and benefits several avian species of conservation concern. Additionally, the landscape context surrounding the managed habitat influences avian community composition. Managed barrens that are adjacent to remnant barrens, rather than adjacent to woodland habitats, have the highest potential for conserving barrens breeding birds.