Euphydryas Aurinia Research Articles

Genetic Divergence Among Host-Specific Cryptic Species in <I>Cotesia melitaearum</I> Aggregate (Hymenoptera: Braconidae), Parasitoids of Checkerspot Butterflies

We used mitochondrial DNA sequence data and 12 microsatellite loci to examine the genetic structure of Cotesia melitaearum (Wilkinson) (Hymenoptera: Braconidae), a parasitoid wasp reared from two common butterfly species, Melitaea cinxia (L.) (Lepidoptera: Nymphalidae) and Euphydryas aurinia (Rottemburg) (Lepidoptera: Nymphalidae), across many localities in Europe and Asia, as well as from four more narrowly distributed related European species. The haplotypes of wasps reared from M. cinxia and E. aurinia show a complex geographic pattern presumably reflecting long-term history, but the microsatellite data yield two host-associated groups, each including populations across Europe and suggesting that currently there is no gene flow between the parasitoid populations attacking these two widely distributed host species. The cryptic species attacking E. aurinia also parasitizes the closely related Euphydryas desfontainii (Godart), but the three remaining host species have yet another, or possibly several, previously unrecognized parasitoid species. We support the cryptic species status of two C. melitaearum aggregate forms parasitizing M. cinxia and Melitaea athalia (Rottemburg) in the Aland Islands in Finland and provide behavioral and ecological data in addition to the molecular data.

Isolation and characterization of polymorphic microsatellites in Parnassius apollo and Euphydryas aurinia (Lepidoptera)

AbstractMicrosatellite loci were developed from Parnassius apollo and Euphydryas aurinia, two endangered Palaearctic butterfly species. Respectively, six and five polymorphic loci were characterized from an enriched partial genomic library. Genetic diversity range from three to 25 alleles for the first species and from seven to 21 for the second. Although the presence of null alleles is suspected, these polymorphic loci are likely to provide important information on the fine scale genetic structure among populations of these species.

Marsh Fritillary (Euphydryas aurinia) in the Czech Republic: monitoring, metapopulation structure, and conservation of an endangered butterfly

Thirty colonies of the Marsh Fritillary (Euphydryas aurinia) have recently been recorded in Western Bohemia, Czech Republic. The majority of colonies are small, their total area are 1.5 km2. Small size and intensive grazing/mowing were positively associated with observed declines/extinctions, while abandonment threatens the colonies in the longer term. Short distances to nearest colonies buffered against declines. High colony turnover, asynchronous local dynamics pointed and the species’ biotope requirements all point to a dynamic metapopulation structure; patterns of connectivity revealed that there are several metapopulations within the region. Because conserving the species within its extant sites seems unsustainable in the long term, restoration of its habitats is proposed.

Contrasting movement patterns in two species of chequerspot butterflies,Euphydryas auriniaandMelitaea phoebe, in the same patch network

Abstract. 1. Mark–release–recapture studies were conducted on two species of chequerspot butterfly,Euphydryas auriniaandMelitaea phoebe, in the same habitat patch network in Yanjiaping, a small basin in the Taihang Mountains, north‐west of Beijing, China, in 2000.2.Euphydryas auriniatended to stay in the habitat patches and to move to neighbouring patches, whereasM. phoebemoved widely among patches in the entire network.3. The parameters of the virtual migration model showed higher daily emigration propensity inM. phoebeand inE. auriniamales than inE. auriniafemales, and significantly greater average daily movement distance inM. phoebethan inE. aurinia.4. The results are consistent with the previous findings showing genetic structuring among local populations ofE. auriniabut not among local populations ofM. phoebe.5. Based on the genetic and ecological results, it was concluded thatE. auriniahas a classic metapopulation in the study area, whereasM. phoebeappears to have a source–sink metapopulation.6. In 2000, when there was an overall increase in the abundance of the two species, the limited mobility ofE. auriniaresulted in an increase in the average local population size, whereas the increase in the number of local populations inM. phoebewas due to its high mobility.

High similarity between flanking regions of different microsatellites detected within each of two species of Lepidoptera:Parnassius apolloandEuphydryas aurinia

Microsatellite flanking regions have been compared in two butterfly species. Several microsatellite flanking regions showed high similarity to one another among different microsatellites within a same species, but very few similarities were found between species. This can be the consequence of either duplication/multiplication events involving large regions containing microsatellites or of microsatellites imbedded in minisatellite regions. The multiplication of microsatellites might also be linked to mobile elements. Furthermore, crossing over between nonhomologous microsatellites can lead to the exchange of the flanking regions between microsatellites. The same phenomenon was observed in both studied butterfly species but not in Aphis fabae (Hemiptera), which was screened at the same time using the same protocol. These findings might explain, at least partially, why microsatellite isolation in Lepidoptera has been relatively unsuccessful so far.

Habitat of pre-hibernating larvae of the endangered butterfly Euphydryas aurinia (Lepidoptera: Nymphalidae): What can be learned from vegetation composition and architecture?

Habitats of pre-hibernating gregarious larvae of the endangered Marsh Fritillary butterfly (Euphydryas aurinia) were studied in field in Western Bohemia, Czech Republic. The species inhabits moist seminatural meadows managed by light grazing and haymaking; the only local host plant is Succisa pratensis. The redundancy analyses of the vegetation composition (around 166 nest-bearing and 381 unoccupied host plants) showed that nest presence was positively associated with short cushion-forming grasses (esp. Nardus stricta). It was negatively associated with competitively superior tall grasses (e.g., Deschampsia caespitosa) and tall herbs. Comparison of Ellenberg's indicator values of vegetation in occupied vs. unoccupied plots revealed that the nests were more often found in drier, nitrogen-poorer and more acidic conditions than unoccupied plants. Multiple regressions of nest presence against architecture of the host plants (170 occupied, 1280 unoccupied) revealed that the nests were associated with densely clumped host plants, low to medium height of sward and mechanical disturbance. The patterns agree with our knowledge of the biology of the larvae: short sward (related to low nitrogen, humidity, and low pH) facilitates larval basking; high host density reduces the likelihood of starvation. Sod disturbance facilitates host plant germination. The conditions favourable for the nests were also favourable for the host plant, but the plants grew in broader range of conditions than that occupied by the nests. The abandoning of a site results into a situation when the conditions first become intolerable for the butterfly, and ultimately for the plant. Since grazing and mowing have a different impact on vegetation composition and architecture, we propose that the studied populations have persisted in a semi-dynamic state in the two modes of management. Conservation management should mimic the dynamics of traditional land use on the smaller scales of extant colonies.

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 FST value was 0.1542 but between population groups within the suspected colonisation range of the butterfly (ca. 20 km), FST values were not significantly different from zero. Genetic diversity in terms of number of alleles and heterozygosity was reasonably high in natural populations (He=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.

Combining larval habitat quality and metapopulation structure – the key for successful management of pre-alpine Euphydryas aurinia colonies

This study aims to analyse larval habitat preferences and landscape level population structure of the threatened Marsh Fritillary butterfly, Euphydryas aurinia, and discusses implications for the conservation and management of this strongly declining species in central Europe. Whereas current management strategies are mainly based on studies of habitat requirements of adult individuals, we intend to emphasise larval habitat quality and population processes at the landscape level as additional key factors. Microhabitat preference analysis of egg-laying females showed that eggs were predominantly laid on prominent large-sized host plant individuals. Additionally, when Succisa pratensis was used as a host plant (as opposed to Gentiana asclepiadea), host individuals in open vegetation structure were preferred. Optimal oviposition conditions were present in recently abandoned calcareous fen meadows and at the edges of such meadows currently in use. A two-year patch-occupancy study in the northern pre-alpine region of south-west Germany indicated that E. aurinia lives in a metapopulation. In a logistic-regression model, patch size, isolation, and habitat quality explained 82% of the observed patch-occupancy pattern in 2001. Our data suggest that a suitable conservation strategy must incorporate both the conservation of a network of suitable habitat patches, and efforts to maximise local habitat quality by ensuring that host plants can grow to a large size and are surrounded by sparse and low vegetation cover.

Genetic differentiation within metapopulations of Euphydryas aurinia and Melitaea phoebe in China.

We analyzed genetic differentiation within metapopulations of two species of checkerspot butterfly, Euphydryas aurinia and Melitaea phoebe, in China. To generate genetic information, we used a new molecular technique, DALP - direct amplified length polymorphism. AMOVA results showed that most of the variation occurred among individuals within local populations of both E. aurinia and M. phoebe. However, while there was differentiation among local population in E. aurinia (P < 0.001), there was no subdivision in metapopulation of M. phoebe (P = 0.210). This is consistent with the behavior of M. phoebe adults being more dispersive than E. aurinia. Within the M. phoebe metapopulation, three neighboring patches were always occupied during the observation period (1998-2000). In addition, the number of individuals in these three populations accounted for the majority of M. phoebe larvae, and hence we conclude that the M. phoebe metapopulation might exist as a source-sink metapopulation. On the other hand, the E. aurinia metapopulation is an example of a classical metapopulation. Therefore, the conservation management of these two species should reflect these differences.

Dynamic populations in a dynamic landscape: the metapopulation structure of the marsh fritillary butterfly

The marsh fritillary butterflyEuphydryas auriniais an endangered species in most of northern Europe. We describe the metapopulation structure ofE. auriniain Finland, where the species has declined drastically in the past decades. We found two types of habitat patches suitable for the species: semi‐permanent meadows and transient clearcuts in the forest. Patch area was the most significant variable predicting the occurrence ofE. auriniain a habitat patch. The species tended to be found in young rather than old clearcuts, apparently because the vegetation became too high in the latter. We used the incidence function model to simulate the metapopulation dynamics ofE. auriniain its dynamic landscape and discovered that the continued presence of the semi‐permanent meadows is essential for the survival of the species in the study area in southeast Finland.

When random sampling does not work: standard design falsely indicates maladaptive host preferences in a butterfly

In experiments that investigate species’ interactions, individuals are often chosen at random to represent their populations. However, this practice can generate misleading results when individuals of different species do not interact at random. We illustrate this effect by examining oviposition preferences of Euphydryas aurinia butterflies from three populations using three different plant genera. We first offered each insect a randomly chosen member of its own host population and a foreign host (Succisa pratensis) not present in the insect’s habitat. The butterflies uniformly preferred the foreign Succisa over their own hosts. Preferences were apparently maladaptive because insects wasted time searching for a nonexistent plant. We repeated the experiment using individual hosts that had naturally received eggs in the field. The overall preference for Succisa and the appearance of maladaptation both disappeared. In the original experiments, our random choice of experimental host individuals had combined with strong within‐species discrimination by the butterflies and with overlap of acceptability between host species to obscure the true nature of host preference.

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.

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.

Phylogeography of the Marsh Fritillary Euphydryas aurinia (Lepidoptera: Nymphalidae) in the UK

We sampled extant (and extinct) populations of Euphydryas aurinia to examine the phylogeography of the species in the UK. We were interested in whether the genetic structuring of populations reflects anything other than a single recent post-glacial colonization event. Four hundred base pairs of the mitochondrial Cytochrome b gene were sequenced from individuals from populations throughout the UK, as well as populations in France and Portugal and seven polymorphic allozyme loci resolved. The mean number of allozyme alleles per locus was 4 and isolation by distance was shown not to be a factor in the geographic structuring of genetic diversity either with or without the inclusion of the French data (Mantel statistic Z=0.015 and 0.112 respectively, P>0.5). Cytochrome b nucleotide diversity (average number of nucleotide differences per site between two sequences) was low overall (0.003, n=63) but mean cytochrome b gene diversity over all populations was 0.77. The presence of the Portuguese and French haplotype in Scottish populations indicates that the Iberian peninsula was likely to be one glacial refugium for E. aurinia populations. The pattern of mitochondrial DNA found in the UK could be interpreted in one of two ways: (1) two separate colonization events or (2) a single slow colonization event. Allele frequency distributions followed a similar geographical pattern as mitochondrial DNA haplotypes. An AMOVA assigned just 2.68% of allozyme genetic variation to the grouping tested, providing more support for the single colonization event theory.

Phylogeography of the Marsh Fritillary Euphydryas aurinia (Lepidoptera: Nymphalidae) in the UK

Abstract We sampled extant (and extinct) populations of Euphydryas aurinia to examine the phylogeography of the species in the UK. We were interested in whether the genetic structuring of populations reflects anything other than a single recent post-glacial colonization event. Four hundred base pairs of the mitochondrial Cytochrome b gene were sequenced from individuals from populations throughout the UK, as well as populations in France and Portugal and seven polymorphic allozyme loci resolved. The mean number of allozyme alleles per locus was 4 and isolation by distance was shown not to be a factor in the geographic structuring of genetic diversity either with or without the inclusion of the French data (Mantel statistic Z=0.015 and 0.112 respectively, P >0.5). Cytochrome b nucleotide diversity (average number of nucleotide differences per site between two sequences) was low overall (0.003, n =63) but mean cytochrome b gene diversity over all populations was 0.77. The presence of the Portuguese and French haplotype in Scottish populations indicates that the Iberian peninsula was likely to be one glacial refugium for E. aurinia populations. The pattern of mitochondrial DNA found in the UK could be interpreted in one of two ways: (1) two separate colonization events or (2) a single slow colonization event. Allele frequency distributions followed a similar geographical pattern as mitochondrial DNA haplotypes. An AMOVA assigned just 2.68% of allozyme genetic variation to the grouping tested, providing more support for the single colonization event theory.

Use of space and resources in a Mediterranean population of the butterfly Euphydryas aurinia

Population parameters of a central Spanish population of the marsh fritillary butterfly, Euphydryas aurinia (Rott.), were studied between 1990 and 1991. Numbers were very similar in the two study years, but adult emergence was delayed 15 days in the second year as a consequence of variation in weather conditions. The nectar sources used by adults were different in the two years and were correlated with the most abundant plants available during the flight period. Adult abundance was correlated with the presence of the larval foodplant ( Lonicera periclymenum) in open woodland and was higher where this habitat was mixed with open areas, where nectar sources abound. Estimates of male population numbers were higher than those of females, suggesting that males used wider ranges than females, or a higher residence rate for males. Nevertheless mobility did not differ significantly between the sexes. The conservation of the species in central Spain depends on maintaining patches of oak woodlands, important for the presence of larval foodplants. Large numbers of the butterfly in the Natural Park study area are favoured by traditional management including charcoal production and extensive grazing by cattle and goats.

A review of the distribution, ecology and status of the marsh fritillary Euphydryas aurinia Rottemburg, 1775 (Lepidoptera: Nymphalidae) in Ireland : T. A. Lavery, Irish Naturalists' Journal, 24(5), 1993, pp 192–199

The status of the marsh fritillary in Ireland is re-assessed, based on comparison between pre and post 1980 distribution data and consideration of what is now known of the biology of this butterfly in Ireland. It is concluded that E. aurinia cannot realistically be regarded as a threatened species in Ireland but that man-induced habitat loss and habitat modification which have caused its decrease elsewhere in Europe are occurring in Ireland. It is recommended that consideration be given to protection of the few Irish sites supporting very large populations of this species, currently regarded as internationally threatened. Attention is drawn to the need for better data on present site management where large populations occur in Ireland and for more information on the controlling effects of predation and parasitization. It is suggested that re-assessment of the international status of this species would be appropriate. The marsh fritillary Euphydryas aurinia (Rott.) is an attractive butterfly associated with certain types of old pasture in Ireland. In various parts of Europe its distribution and abundance have declined noticeably during the latter part of the 20th century, resulting in its recognition as an internationally threatened species. In the present account, the biology and status of E. aurinia in Ireland are reviewed, using an extended data sheet format. The biological data are derived from direct observation by the author and others on Irish populations of this butterfly. The distribution data are

Basking Behaviour in Larvae of the Butterfly Euphydryas Aurinia

The body temperatures of basking Euphydryas aurinia Rott. (Nymphalidae) larvae were measured using thermistor probes under field conditions. These showed that basking larvae could elevate their body temperatures to well above ambient. Under conditions of low ambient temperature but high solar radiation intensity, the temperature excess (i.e. observed body temperature minus ambient) of basking larvae can attain 30'C. Increases in body temperature were positively correlated with periods of high solar radiation intensity. The contribution of larval colour and behaviour is discussed.