Habitat utilization of the Glanville fritillary butterfly in the Tianshan Mountains, China, and its implication for conservation

In 2004 and 2005, we investigated the distribution of the Glanville fritillary butterfly (Melitaea cinxia) and its host plant in the Tianshan Mountains of China, in order to clarify its habitat utilization. Female butterflies were almost captured on dry meadows on slopes, where plenty of host plant (Veronica spicata) is distributed. Although there are host plants on meadows at higher altitude (>2,050 m), no butterfly was found there. Among the meadows with host plants, a patch of dry meadow (newly-found meadow) was considered as a natal site because of the high density and freshness of butterflies. Unlike females, male butterflies were almost captured in valleys, where there are no host plants and fewer nectar plants. It might be related to specific mating system of M. cinxia in the study site, or specific environmental factors. Thus valleys are important habitat for males. Although the status of M. cinxia is yet unknown in China, we give some suggestions for conservation based on this study. First, dry meadows with host plants are the essential component for persistence of M. cinxia, among which the specific sites with more favourable conditions, such as natal site, are most important. Second, as a site with adult resource requirement, valleys should be included as part of the habitat of the butterfly. Finally, meadows at higher altitude are not utilized at present but they are potential habitats which need to be maintained for any shifts in altitudinal range in response to global warming in the future. Hence, the trade-off of present cost and future benefit should be taken into account when formulating a conservation strategy for M. cinxia in the Tianshan Mountains of China.     

Metapopulation structure and movements in five species of checkerspot butterflies

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

Host plant use by the Heath fritillary butterfly, Melitaea athalia: plant habitat, species and chemistry

We present a study of habitat use, oviposition plant choice, and food plant suitability for the checkerspot butterfly Melitaea athalia Rottemburg (Lepidoptera: Nymphalidae) in Åland, Finland. We found that in Åland, unlike in the mainland of Finland and many parts of its range, M. athalia flies mainly in open meadows. When offered an array of plants in a large (32 × 26 m) field cage, they predominately oviposited upon Veronica chamaedrys L., V. spicata L. and Plantago lanceolata L. (Plantaginaceae), which grow in open meadows. The relative abundance of the butterfly in Åland, and its habitat and host plant use there, may reflect local adaptation to land use practices and geology that maintain clusters of small open meadows with little successional change. At the scale of a plant patch, preferred species were used as frequently in mixed species patches as in mono-specific patches, and more oviposition occurred in open than in grassy patches. All of the host plants used by M. athalia are defended by iridoid glycosides (IGs). However, oviposition choice among species and among individual plants within species was largely independent of IG concentration. This contrast with the more discerning congener, M. cinxia, supports the idea that host discrimination decreases with increasing host range. Finally, although the adult butterflies chose specific plant species for oviposition, as larvae they performed well on twelve out of thirteen species of plants, including both known hosts and related novel plants that occur in Åland, indicating a much wider range of larval food plant species than adult oviposition species.     

Habitat exploration in butterflies – an outdoor cage experiment

A large outdoor cage, measuring 7 × 30 m, was used to study the willingness of butterflies to move through unsuitable habitat in search of neighbouring patches. The area inside the cage was divided into two grassland parts by a 7 m long shady part of unsuitable habitat that the butterflies had to fly through to move between the grassland parts. In 1999 and 2000 we performed experiments on three Melitaeini species (Melitaea cinxia and Mellicta athalia were used both years and Euphydryas aurinia in 2000) and three additional species (Brenthis ino and Aphantopus hyperantus in 1999 and Clossiana euphrosyne in 2000). In both years the Melitaeini species moved at considerably lower rates through the shady part than the other species. Among the Melitaeini species Mell. athalia moved most frequently through the shady part while E. aurinia and M. cinxia moved at lower rates. The distribution of these butterflies differ from widespread to localized and the results are discussed in the context of their habitat preferences and distribution patterns.     

Effects of local and landscape factors on the abundance of an endangered multivoltine butterfly at riverbanks

Plebejus argyrognomon is one of the grassland‐dwelling butterflies undergoing rapid decline in recent decades. Grassland habitats for butterflies are generally threatened by fragmentation and invasive species, hence are among the most vulnerable ecosystems. We studied the seasonal abundance of P. argyrognomon at habitat patches along the banks of the Kinugawa River in eastern Japan, to identify environmental factors suitable for population persistence of this species, including habitat patch connectivity. Results showed that the patch's host plant cover had a positive effect on abundance in all three seasons, while the shading of the host plants by surrounding non‐host plants and nearby forested area showed negative effects. Additionally, habitat patch connectivity and nectar richness could be considered as positive factors in autumn and summer, respectively. Analysis of habitat connectivity also showed that the Kinugawa River did not appear to act as a dispersal barrier for P. argyrognomon. Our findings emphasize the importance of understanding environmental factors that may vary among seasons, and such understanding could contribute to habitat management of multivoltine butterflies in fragmented landscapes.     

Habitat use and population biology of the Danube Clouded Yellow butterfly Colias myrmidone (Lepidoptera: Pieridae) in Romania

The Danube Clouded Yellow (Colias myrmidone) is one of the most endangered butterflies in Europe. Its distribution range shrank dramatically in the last few decades due to the extinction of populations in Western and Central Europe. Ecological studies were commenced when populations were already at the verge of extinction, thus our knowledge on the population ecology and habitat use of this butterfly is very limited. Here we report the results of a study on habitat preferences, egg distribution and demography of C. myrmidone in Romania, perhaps the last remaining stronghold of the species in Central Europe. We found that Danube Clouded Yellow adults occurred mainly in mesophilous grasslands created by forest clearing and then maintained by low intensity grazing allowing bushes and forest-edge vegetation to develop and host Chamaecytisus species to grow. Butterflies highly preferred lightly grazed pastures over hay meadows and abandoned grasslands, their density was positively related to host plant density. Egg-laying females preferred habitat patches with relatively high cover of the host plant and tall vegetation. Both apparent survival rate and encounter probability were lower for females than males, and parameter estimations also had much higher errors for females. These indicate that much higher sampling effort is needed to estimate and monitor population parameters for females than for males. Our results provide guidelines for the habitat management and population monitoring of Colias myrmidone, thus may significantly contribute to its successful conservation.     

Local populations of endangered Maculinea (Phengaris) butterflies are flood resistant

Semi-natural wet meadows are threatened by drainage, the abandonment of traditional management and climate change. The large blue butterflies Maculinea teleius and M. nausithous are flagship species associated with wet meadows and are the targets of many conservation programmes. However, there is little knowledge on the impact of natural catastrophes, such as floods, on the persistence of these butterflies. In our study we tested how a flood that resulted in the temporary inundation of meadows affected populations of M. teleius and M. nausithous. Studies were conducted in two consecutive seasons of 2009 (with ‘normal’ weather) and 2010 (with extreme rainfall and a consequent flood in May) in a wet meadow complex located in the Vistula River valley in southern Poland. In both years the abundance of adults was estimated for each local habitat patch (n = 55) within sympatric metapopulations of both species. Additionally, in June 2010, i.e. directly after the flood and shortly before the flight period, a total of 754 Myrmica ant nests in 10 habitat patches (6 inundated vs. 4 not inundated) were checked for the presence of Maculinea larvae and pupae. We found no impact of inundation on year-to-year changes in adult population sizes. The probability of occurrence of Maculinea larvae and pupae in ant nests was higher in temporarily inundated meadows. Our results indicate that temporary inundation occurring after long-term downpours does not negatively affect the investigated species even during the larval period in ant nests at ground level. This provides an argument against drainage works in wet meadows with Maculinea butterflies.     

Patterns of genetic and taxonomic differentiation in three Melitaea (subg. Mellicta) species (Lepidoptera, Nymphalidae, Nymphalinae)

In conservation genetics the existence of Evolutionarily Significant Units (ESU) is a crucial question in threatened or vulnerable species. It is of particular concern to determine whether different subspecies or ecotypes of a species can be considered as separate ESUs. Some Melitaea subg. Mellicta species (e.g. Mellicta aurelia, Mellicta britomartis) are declining or scarce in Europe. Therefore, the level of genetic differentiation and pattern of genetic variation were surveyed in three Melitaea (subg. Mellicta) species. Their habitat requirements and food plants partly overlap; accordingly they often co-occur in the same habitat. M. britomartis and M. aurelia have one brood per year in Hungary, while Mellicta athalia has a monovoltine and a bivoltine ecotype. The purpose of the study was to estimate the number of genetically differentiated ESUs among these species in the Carpathian basin. Samples were taken from 5 Hungarian regions and a few samples were collected in Transylvania as well. Enzyme polymorphism was studied using polyacrylamide gel electrophoresis. The structure of genetic variation was analysed by F-statistics, AMOVA, PCA and Bayesian clustering method. UPGMA dendrogram was constructed on the basis of Cavalli-Sforza and Edwards chord distances. The three species are clearly differentiated from each other in all statistical analyses. They are evidently different Evolutionary Significant Units. The two ecotypes of M. athalia, however, do not show any genetic differentiation.     

Resource use of specialist butterflies in agricultural landscapes: conservation lessons from the butterfly Phengaris (Maculinea) nausithous

Most of the European grassland butterfly species are dependent on species rich grasslands shaped by low intensity farming. Conservation of these specialist species in agricultural landscapes relies on knowledge of their essential resources and the spatial distribution of these resources. In The Netherlands, the dusky large blue Phengaris (Maculinea) nausithous butterflies were extinct until their reintroduction in 1990. In addition, a spontaneous recolonization of road verges in an agricultural landscape occurred in 2001 in the southern part of The Netherlands. We analyzed the use of the essential resources, both host plants and host ants, of the latter population during the summers of 2003 and 2005. First we tested whether the distribution of the butterflies during several years could be explained by both the presence of host plants as well as host ants, as we expected that the resource that limits the distribution of this species can differ between locations and over time. We found that oviposition site selection was related to the most abundant resource. While in 2003, site selection was best explained by the presence of the host ant Myrmica scabrinodis, in 2005 it was more strongly related to flowerhead availability of the host plant. We secondly compared the net displacement of individuals between the road verge population and the reintroduced population in the Moerputten meadows, since we expected that movement of individuals depends on the structure of their habitat. On the road verges, butterflies moved significantly shorter distances than on meadows, which limits the butterflies in finding their essential resources. Finally we analyzed the availability of the two essential resources in the surroundings of the road verge population. Given the short net displacement distances and the adverse landscape features for long-distance dispersal, this landscape analysis suggests that the Phengaris population at the Posterholt site is trapped on the recently recolonized road verges. These results highlight the importance of assessing the availability of essential resources across different years and locations relative to the movement of the butterflies, and the necessity to careful manage these resources for the conservation of specialist species in agricultural landscapes, such as this butterfly species.     

Host associations of <Emphasis Type="Italic">Coenonympha hero</Emphasis> (Lepidoptera: Nymphalidae) in northern Europe: microhabitat rather than plant species

Understanding ecological requirements of endangered species is a primary precondition of successful conservation practice. Regrettably, we know surprisingly little about the life history of numerous threatened insects, and about their use of larval host plants in particular. The brown butterflies (Nymphalidae: Satyrinae) have traditionally been considered polyphagous on grasses and indiscriminatory in their oviposition behavior. However, detailed studies on several species have revealed local specialization in host plant use as well as the decisive role of microlimatic conditions as determinants of habitat quality. The present study addresses host plant relationships in the endangered brown butterfly Coenonympha hero (L.) at the northern limit of its European distribution. We combine laboratory-based host preference and performance tests with an analysis of microhabitat use by adult butterflies in the field. Both lines of evidence suggest that C. hero is polyphagous enough not to be associated with one particular host species. Oviposition choices of C. hero are not driven by host plant species but rather by structural characteristics of the substrate. The preferred rigid needle-like structures may serve as cues of ‘transparent’ vegetation which allows the larvae to benefit from sunlight reaching the lower strata of the tuft. Our results suggest that conservation efforts should prioritize microclimatic parameters, rather than the presence of any particular host plant species, as decisive determinants of habitat quality in C. hero.     

Species traits help explain butterfly habitat use in a tropical forest

1. When species can access all parts of the landscape, species-sorting metacommunity theory predicts that community composition depends on habitat choice and interactions with other species and the environment. These filtering processes can also depend on species' traits.
2. The authors investigated how traits mediate a species-sorting process in determining butterfly community composition in a naturally patchy landscape in the tropical Western Ghats, India. The authors asked, do traits mediate access to certain habitats and does seasonality affect these patterns? The authors surveyed 56 habitat patches in three habitat types: laterite plateau grasslands, ridge grassland, and moist-deciduous forest, in a 65-km² landscape.
3. Non-palatable butterflies showed similar occurrences across seasons and habitats, but palatable butterflies were less commonly encountered in open habitats in the dry season. Polyphagous butterflies occurred infrequently in the dry season in laterite habitats, potentially indicating emigration or diapause patterns are linked to diet breadth.
4. All species were present in all habitats, implying dispersal does not limit access to different habitat patches, consistent with the species-sorting metacommunity concept. Nevertheless, butterfly occurrence was strongly influenced by the interaction of mobility and habitat type with sedentary species occurring less often in low-resource open laterite patches than mobile species.
5. Species sorting is typically regarded as occurring directly through environmental filters, but here the authors suggest that the environmental filter acts through movement limitations. Studies integrating landscape heterogeneity and species characteristics will help us better understand metacommunities and species distributions in nature.

     

Life history of the Glanville fritillary butterfly in fragmented versus continuous landscapes

Habitat loss and fragmentation threaten the long‐term viability of innumerable species of plants and animals. At the same time, habitat fragmentation may impose strong natural selection and lead to evolution of life histories with possible consequences for demographic dynamics. The Baltic populations of the Glanville fritillary butterfly (Melitaea cinxia) inhabit regions with highly fragmented habitat (networks of small dry meadows) as well as regions with extensive continuous habitat (calcareous alvar grasslands). Here, we report the results of common garden studies on butterflies originating from two highly fragmented landscapes (FL) in Finland and Sweden and from two continuous landscapes (CL) in Sweden and Estonia, conducted in a large outdoor cage (32 by 26 m) and in the laboratory. We investigated a comprehensive set of 51 life‐history traits, including measures of larval growth and development, flight performance, and adult reproductive behavior. Seventeen of the 51 traits showed a significant difference between fragmented versus CL. Most notably, the growth rate of postdiapause larvae and several measures of flight capacity, including flight metabolic rate, were higher in butterflies from fragmented than CL. Females from CL had shorter intervals between consecutive egg clutches and somewhat higher life‐time egg production, but shorter longevity, than females from FL. These results are likely to reflect the constant opportunities for oviposition in females living in continuous habitats, while the more dispersive females from FL allocate more resources to dispersal capacity at the cost of egg maturation rate. This study supports theoretical predictions about small population sizes and high rate of population turnover in fragmented habitats selecting for increased rate of dispersal, but the results also indicate that many other life‐history traits apart from dispersal are affected by the degree of habitat fragmentation.     

Landscape context affects the relationship between local and landscape species richness of butterflies in semi‐natural habitats

Local species richness of butterflies can be expected to benefit from both local habitat properties as well as the availability of suitable habitats and source populations in the surrounding landscape. Whether local species richness is dependent on local or landscape factors can be assessed by examining the relationship between local and landscape species richness. Here we studied how local species richness is related to landscape‐level species richness in landscapes differing in agricultural intensity. The relationship was linear for field boundaries in intensively cultivated landscapes and non‐linear in less‐intensively cultivated landscapes. In landscapes containing semi‐natural grasslands (on average 4% of overall land‐use), the relationship was non‐linear for field boundaries, but linear when considering local species richness of the grasslands themselves. These results show that local factors are more important than landscape factors in determining local species richness in landscapes which contained semi‐natural grasslands. Local species richness was limited by landscape factors in intensively cultivated landscapes. This interpretation was supported by the relationship between local species richness and landscape‐scale average mobility and generalist percentage of butterfly assemblages. We conclude that the management of field boundary habitat quality for butterflies is expected to be most effective in landscapes with semi‐natural grasslands, the species composition of which in turn is dependent on the regional occurrence of grasslands. Based on our results, managing non‐crop habitats for the conservation of habitat specialists and species with poor mobility will be most efficient in regions where patches of semi‐natural grasslands occur.     

Not only the butterflies: managing ants on road verges to benefit Phengaris (Maculinea) butterflies

Obligate myrmecophilic butterfly species, such as Phengaris (Maculinea) teleius and P. nausithous, have narrow habitat requirements. Living as a caterpillar in the nests of the ant species Myrmica scabrinodis and M. rubra, respectively, they can only survive on sites with both host ants and the host plant Great Burnet Sanguisorba officinalis. After having been reintroduced into a nature reserve in the Netherlands in 1990, both butterfly species expanded their distribution to linear landscape elements such as road verges and ditch edges outside this reserve. As additional habitat of both butterfly species, vegetation management of these landscape elements became important. Our results show that a management beneficial for Phengaris butterflies should aim to increase the nest density of Myrmica species, at the same time reducing the density of nests of the competitor Lasius niger or at least keeping them at a low density. Unfavourable grassland management under which L. niger thrives, includes not mowing or flail-cutting the grass, or depositing dredgings along the side of the ditch. Management favourable for the two Myrmica species differs, demanding some flexibility if both species are to benefit. M. scabrinodis is best supported with early mowing of the road verge vegetation or late mowing in the nature reserve, both of which result in an open vegetation and warm microclimate. In contrast, the nest sites of M. rubra should be left undisturbed during the summer, and mown in late autumn. Mowing of butterfly habitat should be avoided between mid-June and mid-September as this would remove the flowerheads of the Sanguisorba plants, on which the butterflies lay their eggs.     

Population level correlation between pre-alighting and post-alighting host plant preference in the Glanville fritillary butterfly

1. In many populations of the Glanville fritillary butterfly Melitaea cinxia, ovipositing females exhibit a post‐alighting preference for one of the potential host plant species available. The work reported here aimed to establish whether females with different post‐alighting preferences can discriminate between their host plant species prior to alighting, and whether pre‐alighting and post‐alighting preferences are correlated at the population level. 2. Alighting and oviposition events were recorded for groups of females from six populations in greenhouse and field experiments. 3. Landing frequencies did not change with experience, indicating that M. cinxia females did not learn from previous encounters with host plants. 4. Females from populations exhibiting post‐alighting preference searched efficiently for their host plants in the sense that they landed mainly on the species on which they oviposited predominantly. Pre‐alighting and post‐alighting preferences were correlated at the population level. 5. The correlation between pre‐alighting and post‐alighting preferences helps to explain why in nature, where the host plants often occur in distinct patches, females are more likely to colonise habitat patches in which their preferred host plant is abundant.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

Thermal biology of flight in a butterfly: genotype,flight metabolism,and environmental conditions

Knowledge of the effects of thermal conditions on animal movement and dispersal is necessary for a mechanistic understanding of the consequences of climate change and habitat fragmentation. In particular, the flight of ectothermic insects such as small butterflies is greatly influenced by ambient temperature. Here, variation in body temperature during flight is investigated in an ecological model species, the Glanville fritillary butterfly (Melitaea cinxia). Attention is paid on the effects of flight metabolism, genotypes at candidate loci, and environmental conditions. Measurements were made under a natural range of conditions using infrared thermal imaging. Heating of flight muscles by flight metabolism has been presumed to be negligible in small butterflies. However, the results demonstrate that Glanville fritillary males with high flight metabolic rate maintain elevated body temperature better during flight than males with a low rate of flight metabolism. This effect is likely to have a significant influence on the dispersal performance and fitness of butterflies and demonstrates the possible importance of intraspecific physiological variation on dispersal in other similar ectothermic insects. The results also suggest that individuals having an advantage in low ambient temperatures can be susceptible to overheating at high temperatures. Further, tolerance of high temperatures may be important for flight performance, as indicated by an association of heat‐shock protein (Hsp70) genotype with flight metabolic rate and body temperature at takeoff. The dynamics of body temperature at flight and factors affecting it also differed significantly between female and male butterflies, indicating that thermal dynamics are governed by different mechanisms in the two sexes. This study contributes to knowledge about factors affecting intraspecific variation in dispersal‐related thermal performance in butterflies and other insects. Such information is needed for predictive models of the evolution of dispersal in the face of habitat fragmentation and climate change.     

Local and landscape effects on butterfly density in northern Idaho grasslands and forests

Understanding butterfly response to landscape context can inform conservation management and planning. We tested whether local-scale resources (host and nectar plants, canopy cover) or landscape context, measured at two scales, better explained the densities of four butterfly species. The density of Coenonympha tullia, which has host plants strongly associated with grassland habitats, was positively correlated with the amount of grassland in 0.5- and 1-km radius landscapes and only occurred in forests when they bordered grasslands. For the other species, Celastrina ladon, Cupido amyntula, and Vanessa cardui, local-scale resources better explained butterfly densities, emphasizing the importance of local habitat quality for butterflies. These three species also used host plants that were distributed more heterogeneously within and among habitat types. Our findings demonstrate the importance of host plant spatial distributions when determining the scale at which butterfly density relates to resources, and we recommend that both these distributions and landscape context be evaluated when developing butterfly monitoring programs, managing for species of concern, or modeling potential habitat.     

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.     

Larval habitat preferences of a threatened butterfly species in heavy-metal grasslands

Understanding the factors that determine habitat quality is of vital importance in ensuring appropriate habitat management. Here we used the Niobe fritillary (Argynnis niobe) as a study system to analyse the larval habitat preferences in a small network of heavy-metal grasslands in western Germany. The data were compared with the results of a previous study in coastal dune grasslands of the German North Sea. Based on this knowledge, we give management recommendations for the conservation of this threatened species. The key factors for the survival of A. niobe in heavy-metal grasslands were (i) open vegetation with a warm microclimate and (ii) sufficient host plants for the larvae. This reflects similar results from the previous study in coastal grey dune grasslands. However, in the heavy-metal grasslands, physiological stress generally slows down succession and favours the fritillary’s host plant, the metallophyte Viola calaminaria. As a result, the cover of the host plant was nearly twice as high in heavy-metal grasslands compared to the dune grasslands. Heavy-metal grasslands are of great significance for the conservation of A. niobe and overall butterfly diversity. Usually, the speed of succession in heavy-metal grasslands is slow and, hence, sites with high heavy-metal concentrations are characterised by relatively stable plant composition and vegetation structure. However, on soils with low heavy-metal content a loss of habitats of A. niobe and associated species of conservation concern may occur without management. On those sites sheep grazing seems to be an appropriate way to keep the habitats open and rich in violets.