Euphydryas Aurinia Research Articles

Cluster biodiversity as a multidimensional structure evolution strategy: checkerspot butterflies of the group Euphydryas aurinia (Rottemburg, 1775) (Lepidoptera: Nymphalidae)

AbstractIn the present paper, we explore the evolution of cluster structure in closely related species in the Euphydryas aurinia complex based on morphological (wing pattern, genital armatures) and molecular (cytochrome c oxidase subunit I) characters. Male genitalia differ in the length and shape of the uncus, harpe and juxta branches, by the shape of some parts of the phallus, and by the amount of spikes on the ventral section of the valva. The main trends in the vertical distribution of the E. aurinia group are dwarfism with increasing altitude, coupled with enlargement of paler and darker‐coloured elements of the wing pattern, increasing the overall contrast. Unlike the Euphydryas maturna, the E. aurinia complex forms many local populations specialized under different ecological conditions, probably affected by different evolutionary scenarios. The phylogenetic analysis of the group reveals two ecologically distinct subgroups: one associated with the boreal forest‐mesophyllic meadow biome and one associated with the xeromesophyllic steppe biome. Within each group, two major ecological strategies have evolved in parallel: montane and lowland. Based on the results of the analyses, we revise the nomenclature as follows: E. aurinia pyrenesdebilis (Verity, 1928), stat. rev. (= debilis Oberthür, 1909, syn.n., nomen nudum), E. aurinia bulgarica (Fruhstorfer, 1916), stat. rev., E. aurinia provincialis (Boisduval, 1828), stat. rev. and E. beckeri (Lederer, 1853), stat. rev. The following name‐bearing types are designated: neotype of Papilio aurinia Rottemburg, 1775, neotype of Papilio merope de Prunner, 1798, lectotype of Melitaea beckeri Lederer, 1853, and lectotype of Melitaea aurinia banghaasi Seitz, 1908. All name‐bearing types are figured. A new subspecies, Euphydryas laeta ostracon Korb, Bolshakov, Fric, ssp.n., is described (type locality by holotype data: Kazakhstan, Vostochno‐Kazakhstanskaya Oblast, Shemonaikha).

Surrounding vegetation mediates frequency of plant–herbivore interactions in leaf-feeders but not in other herbivore groups

The overall impact of herbivores on plant population dynamics is determined by a combination of herbivore impact on individuals and frequency of herbivore occurrence within the plant population. While the first component is well-explored, the second is much less studied and may depend both on plant individual traits and on local environmental conditions. In the present study, we focus on determinants of occurrence of damage by five herbivore functional groups to a wet meadow perennial, Succisa pratensis. We assess the role of plant individual traits (size and conspicuousness) and the surrounding vegetation (regarded as a proxy of local biotic and abiotic conditions) in a field study spread over several spatial (and partly temporal) scales.Overall occurrence of damage both by invertebrate and vertebrate folivores and by the specialist folivore Euphydryas aurinia depended on the surrounding vegetation. The preferences to plant individual traits of the two former groups changed with the surrounding vegetation type as well. On the other hand, occurrence of stalk grazing did not vary with any of the tested factors and predispersal seed predators responded only to local inflorescence density. Herbivore damage occurrence varied most at the population scale in all five herbivore groups. The relative importance of the subpopulation scale was lower for vertebrates than for invertebrates. Temporal variation was higher in vertebrate herbivore groups.Our results demonstrate that occurrence of damage by three important herbivore groups changed substantially on a relatively finely delimited gradient of local conditions even at the spatial scale of metres. Such spatial heterogeneity could give rise to a mosaic of different subpopulations within the plant population each with different dynamics. This would in turn substantially change the plant species’ long-term persistence prospects in landscape.

Three in One--Multiple Faunal Elements within an Endangered European Butterfly Species.

Ice ages within Europe forced many species to retreat to refugia, of which three major biogeographic basic types can be distinguished: "Mediterranean", "Continental" and "Alpine / Arctic" species. However, this classification often fails to explain the complex phylogeography of European species with a wide range of latitudinal and altitudinal distribution. Hence, we tested for the possibility that all three mentioned faunal elements are represented within one species. Our data was obtained by scoring 1,307 Euphydryas aurinia individuals (46 European locations) for 17 allozyme loci, and sequencing a subset of 492 individuals (21 sites) for a 626 base pairs COI fragment. Genetic diversity indices, F statistics, hierarchical analyses of molecular variance, individual-based clustering, and networks were used to explore the phylogeographic patterns. The COI fragment represented 18 haplotypes showing a strong geographic structure. All but one allozyme loci analysed were polymorphic with a mean F ST of 0.20, supporting a pronounced among population structure. Interpretation of both genetic marker systems, using several analytical tools, calls for the recognition of twelve genetic groups. These analyses consistently distinguished different groups in Iberia (2), Italy, Provence, Alps (3), Slovenia, Carpathian Basin, the lowlands of West and Central Europe as well as Estonia, often with considerable additional substructures. The genetic data strongly support the hypothesis that E. aurinia survived the last glaciation in Mediterranean, extra-Mediterranean and perialpine refugia. It is thus a rare example of a model organism that combines attributes of faunal elements from all three of these sources. The observed differences between allozymes and mtDNA most likely result from recent introgression of mtDNA into nuclear allozyme groups. Our results indicate discrepancies with the morphologically-based subspecies models, underlining the need to revise the current taxonomy.

THE PROTECTED SPECIES OF LEPIDOPTERA IN THE LANDSCAPE RESERVE ‘ZVANETS’ (BELARUS)

The article contains the data on distribution, population, habitats and phenology of 16 species of lepidopteran insects (Insecta: Lepidoptera), inhabiting the landscape reserve “Zvanets” (Belarus, Brest region) and listed in the Red Book of the Republic of Belarus: Rhyparioides metelkana, Pericallia matronula, Callimorpha dominula, Arytrura musculus, Diachrysia zosimi, Chariaspilates formosaria, Scopula caricaria, Gagitodes sagittata, Lycaena dispar, Euphydryas aurinia, Eu. maturna, Melitaea phoebe, M. diamina, M. aurelia, M. britomartis, Coenonympha oedippus. The data conserning the habitat of Parnassius mnemosyne in the reserve are proposed to be considered unreliable. The study also provides the information on Polyommatus daphnis which had previously been a protected species.It was found that the preferred habitat for C. oedippus, Ch. formosaria, Rh. metelkana, A. musculus, D. zosimi is fen mire. It is possible that for Rh. metelkana and A. musculus the reserve “Zvanets” is the key area concerning their preservation in Europe. On the background of the ongoing extinction of these two species in Europe and in connection with burning of vegetation practised in “Zvanets”, the further study of Rh. metelkana and A. musculus, it is particularly relevant for the development and implementation of action plans for their saving.The widespread burning of vegetation on swamps affecting mineral islands (especially small ones) has negative impact on the Papilionoidea species. In larger forest areas which burn only fragmentary the Lepidoptera population is quickly restored in burnt places due to the resettlement from the areas not affected by fire. It requires the development of special protection measures for the Papilionoidea in the reserve area.With the legalization of target burnings in the reserve “Zvanets” the best way out of this situation is the differentiation of ways to fight the bush encroachment and overgrowth of reeds. It is expedient to split the reserve into a number of areas which will be subject to burning alternately in different years. To localize the fire they must be separated by pre-cutting of the low dividing strips. To prevent any further damage by fire to the mineral islands and margins of swamps and dams the unwanted vegetation along the margins of swamps, near the channels and on the borders of swamps with mineral islands should be removed by mowing and cutting out where possible.Key words: Lepidoptera, protection, Red Data Book, wildlife area, burning of vegetation, Belarus.

Testing for local monophagy in the regionally oligophagous Euphydryas aurinia (Lepidoptera: Nymphalidae)

Regionally oligophagous insects are often host plant specialists at smaller geographical scales, so conservation planning should preferably rely on locally derived information. Host use of the endangered butterfly Euphydryas aurinia was investigated close to the northern limit of its European distribution. We experimentally studied host preference of ovipositing females and growth performance of pre-diapause larvae on different hosts, as well as made field observations on host use by surveying larval webs. In the field, Succisa pratensis has remained the only confirmed host of E. aurinia in Estonia. Also, in the laboratory, the females consistently preferred S. pratensis as oviposition substrate. Larval performance was the highest on S. pratensis though the insects were also able to develop on Knautia arvensis. By contrast, species of Plantago, or Valeriana officinalis, were not suitable for larval development. E. aurinia appears thus to be a host specialist in Estonia, even if the regional list includes a higher number of plant species. Our results confirm a pattern repeatedly recorded in Melitaeine butterflies, i.e. geographic differences in host use and local monophagy. As a methodological contribution, we showed that different lab-based methods yield consistent information on host use, though tend to somewhat overestimate the actual host range of the species in the field. Using larval growth rate as an index of host suitability appears problematic for species not time-stressed during their larval development, and survival is a more appropriate parameter to record.

Oviposition site selection of an endangered butterfly at local spatial scales

As pre-hibernating larvae of the marsh fritillary (Euphydryas aurinia) have limited mobility essential resources need to be available at a very local scale. We surveyed larval webs (2011–2013), the host plant devil’s bit scabious (Succisa pratensis) (2012), and derived variables from digital orthophotos and digital elevation models (Normalized Differenced Vegetation Index, accumulated sun hours, slope, aspect) to explain the presence–absence and abundance of larval webs at three different spatial grain sizes (5 × 2.5 m, 10 × 10 m, 25 × 25 m) across seven study sites in northern Jutland, Denmark. Two-component hurdle models indicated that host plant abundance was the only important predictor of presence–absence and abundance of larval webs across the seven sites. The strength of the host plant effect on larval web prevalence increased when enlarging spatial grain size. For presence–absence (and less for abundance), the effect of host plants on larval webs varied across study sites. Using mixed effects models, we additionally analysed presence–absence of larval webs (in 1 × 1 m plots) in relation to detailed host plant measurements (abundance and size), vegetation height, and environmental variables (soil temperature, air temperature and soil moisture) across four of the sites. This showed that larval webs were located in the densest parts of the host plant patches. Given the low mobility of pre-hibernating larvae (<0.5 m), our results suggest that females select dense parts within large patches of host plants as oviposition sites. Future management should concentrate on establishing large patches of the larval host plant.

Dispersal and connectivity effects at different altitudes in the Euphydryas aurinia complex

Across its European range, the Euphydryas aurinia complex (Annex II of the Habitats Directive) includes a series of distinct populations. At least 3 taxa occur in Italy, each showing slight morphological differences and distinct eco-ethological features. For the first time, we compared metapopulation dynamics of E. (a.) glaciegenita inhabiting a site in the NW Alps (2,100–2,300 m) with E. (a.) provincialis occurring in the Mediterranean biogeographical region in hilly dry grasslands (700 m). To describe patterns of dispersal, we applied the virtual migration model (VMM) to data collected using Mark-Release-Recapture (MRR). We used parameters of survival and migration to explore metapopulation characteristics. In particular we investigated the relative role of connectivity and patch quality in affecting migration rates. We observed differences between the two metapopulation systems, with the “Alpine” population occurring at higher altitude and in more open habitats, showing lower dispersal propensity. In contrast, even though the “Mediterranean” population is more prone to disperse, migration appears to have higher costs. Dispersal abilities affect metapopulation dynamics, which are at the basis of long-term perspectives of survival for butterfly populations. We discuss our results in the framework of conservation and management options for habitats occupied by these Italian taxa of the E. aurinia complex.

Spatial ecology of host–parasitoid interactions: a threatened butterfly and its specialised parasitoid

Habitat conservation for threatened temperate insect species is often guided by one of two paradigms: a metapopulation approach focusing on patch area, isolation and number; or a habitat approach focusing on maintaining high quality habitat for the focal species. Recent research has identified the additive and interacting importance of both approaches for maintaining populations of threatened butterflies. For specialised host-parasitoid interactions, understanding the consequences of habitat characteristics for the interacting species is important, because (1) specialised parasitoids are particularly vulnerable to the consequences of fragmentation, and (2) altered interaction frequencies resulting from changes to habitat management or the spatial configuration of habitat are likely to have consequences for host dynamics. The spatial ecology of Cotesia bignellii, a specialist parasitoid of the threatened butterfly Euphydryas aurinia, was investigated at two spatial scales: within habitat patches (at the scale of individual aggregations of larvae, or ‘webs’) and among habitat patches (the scale of local populations). Parasitism rates were investigated in relation to larval web size, vegetation sward height and host density. Within patches, the probability of a larval webs being parasitized increased significantly with increasing number of larvae in the web, and parasitism rates increased significantly with increasing web isolation. The proportion of webs parasitized was significantly and negatively correlated with cluster density. Among habitat patches the proportion of parasitized webs decreased as cluster density increased. Clusters with a high proportion of larval webs parasitized tended to have lower parasitism rates per larval web. These results support the call for relatively large and continuous habitat patches to maintain stable parasitoid and host populations. Conservation efforts directed towards maintenance of high host plant density could allow E. aurinia to reduce parasitism risk, while providing C. bignellii with sufficient larval webs to allow population persistence.

Host‐associated genomic differentiation in congeneric butterflies: now you see it, now you do not

Ecotypic variation among populations may become associated with widespread genomic differentiation, but theory predicts that this should happen only under particular conditions of gene flow, selection and population size. In closely related species, we might expect the strength of host-associated genomic differentiation (HAD) to be correlated with the degree of phenotypic differentiation in host-adaptive traits. Using microsatellite and Amplified Fragment Length Polymorphism (AFLP) markers, and controlling for isolation by distance between populations, we sought HAD in two congeneric species of butterflies with different degrees of host plant specialization. Prior work on Euphydryas editha had shown strong interpopulation differentiation in host-adapted traits, resulting in incipient reproductive isolation among host-associated ecotypes. We show here that Euphydryas aurinia had much weaker host-associated phenotypic differentiation. Contrary to our expectations, we detected HAD in Euphydryas aurinia, but not in E. editha. Even within an E. aurinia population that fed on both hosts, we found weak but significant sympatric HAD that persisted in samples taken 9 years apart. The finding of significantly stronger HAD in the system with less phenotypic differentiation may seem paradoxical. Our findings can be explained by multiple factors, ranging from differences in dispersal or effective population size, to spatial variation in genomic or phenotypic traits and to structure induced by past histories of host-adapted populations. Other infrequently measured factors, such as differences in recombination rates, may also play a role. Our result adds to recent work as a further caution against assumptions of simple relationships between genomic and adaptive phenotypic differentiation.

European checkerspots (Melitaeini: Lepidoptera, Nymphalidae) are not aposematic – the point of view of great tits (Parus major)

Abstract Checkerspots (Melitaeini, Nymphalidae, Lepidoptera) are usually considered a textbook example of aposematic butterflies and several studies on Nearctic species confirm this. The responses of the avian predator, the great tit (Parus major L., Paridae, Passeriformes), to five species of Palaearctic checkerspots and one control palatable ringlet (Aphantopus hyperanthus L., Satyrinae, Nymphalidae, Lepidoptera) were observed. None of the tested checkerspots was attacked more often than the control palatable ringlet, which suggests that birds originating from the wild had no previous bad experiences with them. Nonetheless, certain tested butterfly species (Melitea didyma, Esper; Melitea aurelia, Nickerl, and probably Melitea athalia, Rottemburg; Melitaeini, Nymphalidae, Lepidoptera) elicited some aversion in great tits (a long time handling them, a small portion of the body eaten, and some discomfort after ingestion). Larvae of commonly eaten species (Melitea diamina, Lang; Euphydryas aurinia, Rottemburg; Melitaeini, Nymphalidae, Lepidoptera) feed on plants containing secoiridoids (in contrast to the earlier mentioned protected species, which feed mostly on plants containing iridoid glycosides); therefore the efficiency of secoiridoids in the chemical protection of butterflies is discussed.

Microsatellites for the marsh fritillary butterfly: de novo transcriptome sequencing, and a comparison with amplified fragment length polymorphism (AFLP) markers.

BackgroundUntil recently the isolation of microsatellite markers from Lepidoptera has proved troublesome, expensive and time-consuming. Following on from a previous study of Edith's checkerspot butterfly, Euphydryas editha, we developed novel microsatellite markers for the vulnerable marsh fritillary butterfly, E. aurinia. Our goal was to optimize the process in order to reduce both time and cost relative to prevailing techniques. This was accomplished by using a combination of previously developed techniques: in silico mining of a de novo assembled transcriptome sequence, and genotyping the microsatellites found there using an economic method of fluorescently labelling primers.Principal FindingsIn total, we screened nine polymorphic microsatellite markers, two of which were previously published, and seven that were isolated de novo. These markers were able to amplify across geographically isolated populations throughout Continental Europe and the UK. Significant deviations from Hardy-Weinberg equilibrium were evident in some populations, most likely due to the presence of null alleles. However, we used an Fst outlier approach to show that these markers are likely selectively neutral. Furthermore, using a set of 128 individuals from 11 populations, we demonstrate consistency in population differentiation estimates with previously developed amplified fragment length polymorphism (AFLP) markers (r = 0.68, p<0.001).SignificanceRapid development of microsatellite markers for difficult taxa such as Lepidoptera, and concordant results with other putatively neutral molecular markers, demonstrate the potential of de novo transcriptional sequencing for future studies of population structure and gene flow that are desperately needed for declining species across fragmented landscapes.

Mark-recapture on large spatial scale reveals long distance dispersal in the Marsh Fritillary, Euphydryas aurinia

1. Long distance dispersal (LDD), or movements far beyond the occupied habitat borders, maintains the integrity of metapopulations in fragmented landscapes. Recent studies on butterflies increasingly reveal that LDD exists even in species that were long regarded as sedentary. Mark–recapture (MR) studies covering larger study areas typically reveal movements among distant colonies. 2. We studied dispersal of the EU-protected, regionally endangered Euphydryas aurinia Rottemburg butterfly in the Czech Republic, using two complementary MR approaches. The single system study was carried out for eight seasons within 30 habitat patches covering 28 ha. The multiple populations study was carried out for a single season, but covering almost all Czech colonies of the species (82 colonies, 110 distinct patches, total area 324 ha within ca 1500 km2). 3. Single system mean lifetime movements were consistently higher for males, but slopes of dispersal kernel power functions were shallower for females, implying that higher proportions of females crossed distances of several kilometres. 4. The multiple populations study allowed detection of 51 lifetime movements exceeding 5 km (41 males, 10 females) and 14 movements exceeding 10 km (13 males, 1 female). Both mean lifetime movements and slopes of the dispersal kernels varied among systems, with no consistent pattern between sexes. All Czech Republic populations are within 0.1% movement probability of both sexes, whereas 1% movement probability delimits three separate management units. 5. Dispersal predictions from local data underestimate total mobility, warning against the use of local MR data for extrapolating long-distance movements. Local dispersal data, however, remain useful for analysing finer details of insect mobility.

Demography of adults of the Marsh fritillary butterfly, Euphydryas aurinia (Lepidoptera: Nymphalidae) in the Czech Republic: Patterns across sites and seasons

The Marsh fritillary (Euphydryas aurinia) (Lepidoptera: Nymphalidae) has declined across Europe, including the Czech Republic. Current conservation strategies rely on prevention of habitat loss and degradation, and increase in habitat quality and connectivity via promoting traditional grassland management. The population structure and adult demography parameters of a single population was investigated for eight years (single system), and of all the known Czech populations (multiple populations) for a single year, using mark-recapture. There was substantial variation in the patterns of adult demography, both among years in the single system and among the multiple populations in a single year. In the single system, the date of the first flight of an adult varied by 18 days over the 8 years and total annual numbers varied with a coefficient of variation of 0.40 (females fluctuating more than males). The average density was ca 80 adults/ha. The population size displayed density-dependence, i.e. decreased following years with high adult numbers, with an equilibrium density of 90 individuals/ha. The average density of the multiple populations was ca 120 individuals/ha. The estimated total population for the Czech Republic was 25,000 individuals (17,000 males / 8,000 females) in 2007, which does not indicate an imminent threat of extinction. The regional persistence of E. aurinia is likely to depend on re-colonisation of temporarily vacant sites by dispersing individuals, facilitated by local shifts in adult flight phenology to that better adapted to local conditions.

Challenges of microsatellite development in Lepidoptera: Euphydryas aurinia (Nymphalidae) as a case study

Currently it remains difficult to obtain robust microsatellite markers for Lepidoptera. In an attempt to overcome the problems associated with developing microsatellite markers for this insect order we combined (i) biotin-enrichment protocol, (ii) next generation pyrosequencing (through 454 GS-FLX Titanium technology) and (iii) the use of individuals collected from eight geographically distant European populations representing three subspecies of Euphydryas aurinia. Out of 96 stringently designed primer pairs, 12 polymorphic microsatellite loci amplified without obvious evidence of null alleles in eight individuals from different subspecies. Between five and seven of these loci showed full within population applicability and three revealed to be robust and transferable between populations and sub-species, providing a first step towards the development of a valuable and robust tool for studying conservation issues and evolution in E. aurinia populations. Nevertheless, as in most studies dealing with Lepidoptera microsatellites, null alleles were detected in most of the developed markers. Our results emphasize the need for further research in order to better understand the complex evolution and organization of Lepidopteran genomes.

The effects of habitat fragmentation on niche requirements of the marsh fritillary, Euphydryas aurinia, (Rottemburg, 1775) on calcareous grasslands in southern UK

The marsh fritillary, Euphydryas aurinia, has declined greatly in distribution across its range within Europe, resulting in its designation as a protected species under Annex II of the 1979 Bern Convention and the EC Habitats and Species Directive. The decline has been linked to a marked reduction in the extent of suitable calcareous and wet grassland habitats, habitats which have been lost through conversion of land to agriculture or urban areas, or reduced in quality due to inappropriate management. The UK is now one of the major strongholds for this butterfly in Europe, although much of the remaining habitat is small, isolated and highly fragmented. E. aurinia populations fluctuate greatly due to the combined effects of biotic (e.g. parasitoids) and abiotic (e.g. climate change) factors. We quantified the habitat associations of larval webs of E. aurinia on fragmented versus extensive (unfragmented) calcareous grassland habitat in southern England to test the hypothesis that habitat requirements of E. aurinia are more constrained within fragmented landscapes. Within both fragmented and unfragmented landscapes the quality and quantity of its main host plant in the UK, Succisa pratensis, was positively related to numbers of E. aurinia larval webs found. The sward height was also important at predicting the distribution of larval webs in both landscapes, although the heights were greater within sites in the unfragmented (≈20 cm) compared to fragmented (≈15 cm) landscape. We also found significant effects of elevation and the cover of bare ground on numbers of larval webs. Elevation was strongly correlated with the availability of host plant, whilst bare ground was only significant on sites within the fragmented landscape, showing a negative relationship with number of larval webs. Our results further emphasise the importance of not only maintaining the habitat quality of extant calcareous grassland sites for E. aurinia in the UK, but also increasing the size and connectivity of these sites to increase the chances and rate of (re)colonisation of unoccupied but suitable habitat. In addition, we show that the habitat requirements of E. aurinia on sites in a large unfragmented landscape may be less specific and thus require less extensive management than that required to create optimal conditions necessary at smaller, more isolated sites in fragmented landscapes.

Butterflies on the brink: habitat requirements for declining populations of the marsh fritillary (Euphydryas aurinia) in SW England

1. The marsh fritillary Euphydryas aurinia is one of our most endangered butterflies, and the only to be protected under European legislation as well as British. It persists in fragile subpopulations threatened by habitat fragmentation and degradation. 2. A combination of swaling and cattle grazing are accepted to be best practice for managing wet, unimproved grasslands—the favoured habitat for E. aurinia in Cornwall. These two well-endorsed methods of management were used to increase and improve the quality of habitat for E. aurinia over a 5 years period, 2004–2008, at a stronghold network of habitat patches in mid Cornwall, south-west England. 3. Analyses of adult and larval densities over 5 years in fifty-four transects across nine sites found E. aurinia to favour habitat patches with higher densities of the larval food plant (Devil’s-bit scabious Succisa pratensis), higher sward height in autumn, and intermediate optimum levels of stock grazing. 4. Main findings indicated most sites experienced significant declines in numbers. Unfavourable weather in the last 2 years of monitoring was likely to have had a significant impact on the response of individual subpopulations to habitat management though poor recovery rates may also reflect a time-lag in colonisation events after habitat improvement has occurred. 5. Habitat management produced an improvement, albeit an inconsistent improvement in habitat variables across patches—S. pratensis shows a clear recovery at some sites. Autumn sward height increased significantly at one site, and a quadratic relationship between stock grazing and important habitat variables has been found which will aid further improvement over all sites for the long term persistence of E. aurinia.

Securing viable metapopulations of the Marsh Fritillary butterfly, Euphydryas aurinia, (Lepidoptera: Nymphalidae) in Northern England

The Marsh Fritillary butterfly, Euphydryas aurinia (Rott.), is regarded as the classic species that requires landscape-scale management in order to maintain viable populations. This paper describes the background to an extinction of this species in Northern England, United Kingdom, and the action taken to reintroduce and monitor the progress of sub-populations between 2007 and 2009. A captive breeding stock was secured using the last 155 known wild caterpillars in 2004 and supplemented by 95 caterpillars collected under licence from the nearest extant populations in western Scotland. Through captive breeding the source population for subsequent reintroduction was increased dramatically and 42,400 caterpillars were used in 2007 to reintroduce the butterfly to four locations in North and West Cumbria. Each of the release sites is within known historical network areas where the butterfly had thrived in past decades. The intention was to restore multiple habitat patches across each network area and eventually establish viable metapopulations across these landscapes. Adults and larval webs have been monitored since 2007 and show that introduction has been successful at three out of the four locations. The failure at the fourth site appears to be unrelated to habitat condition and the causes are currently being investigated. Work is continuing to secure appropriate management of suitable habitat within each landscape area and the number of sub-populations in one area will be increased in the 2010 season.

Changing demography and dispersal behaviour: ecological adaptations in an alpine butterfly

High mountain ecosystems are extreme habitats for all organisms and therefore demand specific adaptations. In this context, we studied the ecology of the butterfly Euphydryas aurinia debilis in the High Tauern (Austria) and compared the obtained data against the ecology of the species in lower elevation habitats. We performed mark-release-recapture studies over the entire flight periods (end of June to end of July) in 2007 and 2008 to analyse the fundamental ecological parameters of a population. The demography of males and females was similar in both years, and no indication of typical protandry was detected. We observed a generally low dispersal of the individuals in both years, but males dispersed significantly more than females in 2008; this finding of low vagility was supported by allozyme analyses. Furthermore, butterflies survived periods of several days of continuously closed snow cover without any indication of increased mortality rates. In these three traits, this alpine population of E. aurinia apparently has ecological and physiological adaptations to the extreme requirements of high-altitude habitats and strongly deviates from the lower elevation populations.

Dispersal of four fritillary butterflies within identical landscape

AbstractBoth species‐specific traits and landscape configuration, such as area and connectivity of habitat patches plus the character of uninhabitable matrix, affect animal movements in fragmented landscapes. Difficulties with disentangling species‐specific and landscape effects have obscured comparisons among species, hindering the understanding of dispersal in metapopulations. To circumvent this complication, we performed a mark–recapture study of four related nymphalid butterflies within identical landscape and in single season. The studied species were three Melitaeinae checkerspots (Euphydryas aurinia,Melitaea athalia,Melitaea diamina) and one Argynnini fritillary (Brenthis ino). Applying the Virtual Migration model revealed that (1) except for mortality within habitat, model parameters differed from those found for the studied species elsewhere; (2) the three Melitaeinae species were more akin in movement parameters than the Argynnini representative (i.e.,B. ino); (3) within Melitaeinae, differences between sexes were more prominent than differences among species; (4) Melitaeinae males left natal patches more readily than females, while the opposite applied toB. ino; (5) males ofM. diaminaand both sexes ofB. inoexhibited highest values of dispersal mortality; (6) except for females ofM. diaminaand both sexes ofB. ino, immigration and emigration scaled with area in females but not in males. Finding (1) demonstrates that geometry of habitat network affects mobility considerably and that transferring dispersal parameters across systems is unwarranted. Still, (2–6) demonstrate that within identical networks, related species follow similar dispersal patterns, suggesting that conservation scenarios suitable for a well‐studied model species would suite related species as well.

Influence of experimental warming and shading on host–parasitoid synchrony

AbstractAs the climate warms, many species are showing altered phenology patterns, potentially disrupting synchrony between interacting species. Recent studies have documented disrupted synchrony in plant–herbivore and predator–prey interactions. However, studies investigating climate‐related asynchrony in host–parasitoid interactions and exploring the relative responses of interacting hosts and parasitoids to climate change are lacking. This is an important gap in knowledge given the ubiquity of insect parasitoids and their importance in influencing the abundance and dynamics of their hosts. In the threatened marsh fritillary butterflyEuphydryas aurinia(Lepidoptera: Nymphalidae) and its specialized parasitoid,Cotesia bignellii(Hymenoptera: Braconidae) phenological synchrony (and consequently population fluctuations) are thought to be weather‐dependent. To assess the likely influence of climate and microenvironment change on synchrony betweenE. auriniaandC. bignellii, we experimentally manipulated the exposure of sensitive‐stage host larvae and parasitoid pupae to temperature (ambient or elevated) and shading (shaded or unshaded) regimes. We also analysed a 20‐year population dynamic dataset from the United Kingdom forE. auriniato investigate whether population variations could be explained by interannual variations in the thermal and sunshine environment. Development times were affected significantly by the experimental temperature and shading treatments forE. auriniabut not forC. bignellii. However, the contrasting responses were insufficient to significantly affect host availability for parasitoids. In the field, thermal and sunshine conditions did not influence population fluctuations, and population variations across a large (UK‐wide) scale were uncorrelated. Changes to the thermal and sunshine environment of the magnitude investigated in our experiment and within the range experienced by wildE. auriniapopulations over the last 20‐years thus seem unlikely to cause breakdown in host–parasitoid synchrony. We suggest that experiments investigating the mechanistic responses of interacting species to environmental change are needed to support the analysis and interpretation of observational data on species' phenology.