Mapping the Habitat Quality of Patch Networks for the Marsh Fritillary Euphydryas aurinia (Rottemburg, 1775) (Lepidoptera, Nymphalidae) in Wales

The marsh fritillary is widely distributed in Wales, with over 200 populations identified in the past 15 years. However, agricultural improvement, habitat fragmentation and changes in agricultural practices continue to impact on status and a decline in range of 23.5% was recorded over a 10-year period. Solutions must be applied at the landscape scale to improve metapopulation viability and, since 2000, surveys of habitat extent and quality have been carried out on 116,373 ha of grassland surrounding 111 populations. Analysis of the results for standard ‘core landscapes’ (based on circles of 1 km radius around known populations) shows that overall cover of suitable breeding habitat is 3.44%. However, only 11.85% of the grassland that was suitable for marsh fritillaries was classified as being in Good Condition. The remainder was regarded as having sub-optimal vegetation structure and/or the foodplant was at low density; 33% of the habitat resource that was not in Good Condition was regarded as inappropriately or excessively managed and 67% was suffering from neglect. An Index of Landscape Quality is used to rank landscapes for evaluation. Thirty-five ‘core landscapes’ (incorporating 98 marsh fritillary populations) were assessed and just four exceeded a threshold value deemed to represent viable landscape configurations. The results have demonstrated that most marsh fritillary populations in Wales exist within depauperate patch networks that lack sufficient breeding habitat of the right quality for long-term persistence. Without targeted conservation action the marsh fritillary will continue to decline in Wales. This paper reports on efforts to collect information on the quality of breeding habitat for marsh fritillaries across Wales in order to identify priorities for conservation action.     

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.     

Mimicry in North American checkerspot butterflies: Euphydryas phaeton and Chlosyne harrisii (Nymphalidae)

ABSTRACT.

• 1 An examination of the phenetics, ecology and distribution of Chlosyne harrisii suggested that it was a potential mimic of the unpalatable Euphydryas phaeton.
• 2 The wing undersurfaces of the adults are very similar and the larvae are virtually identical. The geographic range of both butterflies is the northeastern United States and both are inhabitants of wet meadow habitats where they may fly together. The life histories of both species are closely synchronized.
• 3 A series of feeding experiments using blue-jays as predators showed first, that C.harrisii is palatable, and second, suggested that the birds could not distinguish between C.harrisii and E.phaeton on the basis of visual cues. Thus, Charrisii seems to be an effective Batesian mimic of E.phaeton.

     

两种共存网蛱蝶幼期的生命表研究

在河北省赤城县大海坨国家自然保护区内,从2002年到2004年应用生命表方法研究了在同一网络斑块中共存的金堇蛱蝶Euphydryas aurinia和大网蛱蝶Melitaea phoebe两个世代的幼期各阶段的死亡情况,目的是了解影响种群动态的重要因子,为它们的长期保育提供信息。结果表明,金堇蛱蝶幼期总累积死亡率都较小,两个世代分别为59%和72%; 而大网蛱蝶总累积死亡率较高,两个世代分别为89%和80%。影响大网蛱蝶死亡的最重要因子是放牧,两个世代与放牧相关的k值分别为0.559和0.167;尤其是在越冬后大网蛱蝶幼虫组聚集取食阶段,导致两个世代分别有50%和25%的幼虫组死亡。寄生蜂在大网蛱蝶小种群下也分别使两个世代4%和9%的5～6龄期幼虫以及13%和24%的蛹死亡。金堇蛱蝶死亡主要发生在越冬期,与越冬死亡相关的k值两个世代分别为0.073和0.199, 而寄主植物的质量影响越冬期幼虫组死亡; 寄生蜂则对金堇蛱蝶种群的调控作用极小,只有在2003～2004世代有4.0% 的越冬后幼虫被寄生和7%的蛹被寄生。影响两种蛱蝶种群动态的关键因子不同,采取的保护措施应有所不同。在春季减少源斑块内的放牧,对于以源-汇集合种群形式存在的大网蛱蝶种群恢复和增长十分必要; 而对以经典集合种群形式存在的金堇蛱蝶, 通过适当植被管理提高繁殖区域内寄主植物质量,可以提高越冬期幼虫组存活率,有利于其长期持续生存。     

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.     

Surprising diversity in the Pannonian populations of Marsh Fritillary (Euphydryas aurinia,Lepidoptera: Nymphalidae): Morphometric and molecular aspects

Since genetic variation is the basis of evolutionary potential of a species, its structure needs to be understood. Thus, the aim of this study was to analyze and contrast the structure of genetic and phenotypic variation in the Euphydryas aurinia populations of southeastern central Europe. Genetic variation was studied by two types of molecular genetic markers: mtDNA COI sequences and allozymes. As the great hiatus in the European distribution of E. aurinia is located in the central part of the Carpathian Basin, we expected that the populations East and West to this gap would be highly differentiated. Populations of Central Transdanubia actually represent the easternmost margin of the West European distribution of E. aurinia. In view of the peripheral position of these populations, we supposed to find some genetic sign of local adaptation, as a consequence of diversifying selection and an increased level of fluctuating asymmetry as a result of environmental stress. The analyses of the molecular genetic markers revealed a basic East–West differentiation among the populations of southeastern central Europe which was further structured in the western part of the study area. The results suggested that the genetic differentiation between the two western regions is probably the consequence of diversifying selection. The pattern of phenotypic differentiation among the western populations, however, was different. A geographic cline was revealed (decreasing wing size) toward the eastern margin of the distribution in parallel with increasing fluctuating asymmetry. The conservation inferences of the results are considered.     

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.     

Caterpillars of Euphydryas aurinia (Lepidoptera: Nymphalidae) feeding on Succisa pratensis leaves induce large foliar emissions of methanol

A major new discovery made in the last decade is that plants commonly emit large amounts and varieties of volatiles after damage inflicted by herbivores, and not merely from the site of injury. However, analytical methods for measuring herbivore-induced volatiles do not usually monitor the whole range of these compounds and are complicated by the transient nature of their formation and by their chemical instability. Here we present the results of using a fast and highly sensitive proton transfer reaction-mass spectrometry (PTR-MS) technique that allows simultaneous on-line monitoring of leaf volatiles in the pptv (pmol mol(-1)) range. The resulting on-line mass scans revealed that Euphydryas aurinia caterpillars feeding on Succisa pratensis leaves induced emissions of huge amounts of methanol--a biogeochemically active compound and a significant component of the volatile organic carbon found in the atmosphere--and other immediate, late and systemic volatile blends (including monoterpenes, sesquiterpenes and lipoxygenase-derived volatile compounds). In addition to influencing neighboring plants, as well as herbivores and their predators and parasitoids, these large emissions might affect atmospheric chemistry and physics if they are found to be generalized in other plant species.     

B chromosome variation in Euphydryas colon (Lepidoptera: Nymphalidae)

Cytogenetic analysis of an Idaho population of the checkerspot butterfly, Euphydryas colon, has revealed considerable inter- and intra-individual variation in chromosome number which turns out to be a classic case of B chromosome variation. The basic chromosome complement of the species is n (, )=31. The A chromosomes were aligned equatorially at mitotic metaphase and metaphase II, and axially at metaphase I, indicating a restriction of centric activity at the first meiotic division. No failure of pairing between homologous A chromosomes was observed and, although a marked asynchrony of chromatid separation was found to be characteristic of mitotic telophase and telophase II, the frequency of macrospermatid formation was low. The B chromosomes were at least partly heterochromatic but exhibited some variation in both pycnosity and size. Mitotically stable B-containing individuals showed a preponderance of unpaired Bs at first metaphase and these divided at either first or second anaphase. The presence of Bs was associated with a heightened production of abnormal spermatids particularly in individuals with high numbers of B chromosomes. Among the 25 individuals sampled, 21 carried from 1–6 B chromosomes, and of these 14 were mitotically stable. In all 7 unstable individuals the mean number of B chromosomes per cell exceeded the modal number. Assuming that the modal number represents the zygotic number, these results suggest that a mechanism to boost the number of B chromosomes exists in males of E. colon.     

Variable chemical defence in the checkerspot butterfly Euphydryas gillettii (Lepidoptera: NymphaIidae)

Abstract.

• 1 Like other checkerspots, Euphydryas gillettii butterflies may contain the defensive chemicals, iridoid glycosides, which are sequestered from their hostplants during larval feeding.
• 2 We analysed the iridoid glycoside content of E.gillettii adults from two different populations, Warm Lake, Idaho, and Granite Creek, Wyoming, that have different patterns of hostplant use.
• 3 Gas chromatographic analysis of thirty butterflies from the Wyoming population showed that they contained a mean of 1.27 (±0.19 SE) % dry weight iridoid glycosides. Notably, 20% of these butterflies contained no detectable iridoid glycosides.
• 4 In contrast, nineteen butterflies from the Idaho population contained a mean of 3.89 (±0.38 SE) % dry weight iridoid glycosides, and all butterflies contained iridoid glycosides.
• 5 These results illustrate how the chemical defence of herbivorous insects varies according to differential use of potential hostplants.

     

Molecular evolution of a long wavelength-sensitive opsin in mimetic Heliconius butterflies (Lepidoptera: Nymphalidae)

This study examines the pattern of opsin nucleotide and amino acid substitution among mimetic species 'rings' of Heliconius butterflies that are characterized by divergent wing colour patterns. A long wavelength opsin gene, OPS1 , was sequenced from each of seven species of Heliconius and one species of Dryas (Lepidoptera: Nymphalidae). A parsimony analysis of OPS1 nucleotide and amino acid sequences resulted in a phylogeny that was consistent with that presented by Brower & Egan in 1997, which was based on mitochondrial cytochrome oxidase I and II as well as nuclear wingless genes. Nodes in the OPS1 phylogeny were well supported by bootstrap analysis and decay indices. An analysis of specific sites within the gene indicates that the accumulation of amino acid substitutions has occurred independently of the morphological diversification of Heliconius wing colour patterns. Amino acid substitutions were examined with respect to their location within the opsin protein and their possible interactions with the chromophore and the G-protein. Of the 15 amino acid substitutions identified among the eight species, one nonconservative replacement (A226Q) was identified in a position that may be involved in binding with the G-protein.     

Skipping flights in Ypthima butterflies (Lepidoptera: Nymphalidae)

The skipping flight patterns of three species of Ypthima (Lepidoptera: Nymphalidae) were analyzed using high‐speed video recordings to clarify how wings move and how driving forces are produced. All three species showed a flight pattern that includes a pause that accounts for about 50% of a flap cycle when their wings completely close after each upstroke. The observed pause causes the “skipping” flight trajectory based on the clap–fling mechanism. Pause duration was correlated with upstroke wing motion, suggesting the contribution of the latter to a long pause duration. This is also supported by the temporal relationship between the wing and body motions. The aerodynamic power necessary for the pause flight was calculated for the three species.     

The evolution of ecologically important characters in Heliconius butterflies (Lepidoptera: Nymphalidae): a cladistic review

The biology of Heliconius butterflies has provided a rich source of data to test theories of ecological genetics, coevolution and community ecology. Many putatively adaptive characters have been discussed with reference to a phylogenetic hypothesis based on a variety of morphological and life-history traits interpreted from an evolutionary taxonomic perspective. Here, alternate interpretations of characters on the traditional tree and a more recent mitochondrial DNA cladogram with a substantially different topology are compared and contrasted. It is shown that many characters ostensibly providing support for the traditional phylogenetic hypothesis are almost equally parsimoniously distributed and in some cases more parsimoniously distributed on the mtDNA tree than on the tree inferred from those characters. Discussion of alternate evolutionary scenarios based on the mDNA-based topology is presented for pupal mating, pollen feeding, foodplant coevolution, and other ecologically significant features.     

Food plant density,patch isolation and vegetation height determine occurrence in a Swedish metapopulation of the marsh fritillary <Emphasis Type="Italic">Euphydryas aurinia</Emphasis> (Rottemburg, 1775) (Lepidoptera,Nymphalidae)

The occurrence pattern of the marsh fritillary was studied within a patch network on the Baltic island Öland, Sweden. Presence/absence was established for potentially suitable habitat patches (n = 158) on calcareous moist grassland and analyzed in a multiple logistic regression model where patch area, patch isolation and nine habitat quality variables were included as explanatory variables. Larval food plant density was positively, and patch isolation negatively, correlated to the presence of Euphydryas aurinia. Area did not contribute to the explanation of the occurrence pattern. Significant interactions between larval food plant density times patch isolation, and larval food plant density times vegetation height, show that with low food plant density the butterfly primarily occurs in patches with a vegetation height of 4–10 cm, within a distance of 250 m from nearest occupied patch. In patches with a high food plant density the butterfly occurs in patches where the vegetation height is higher, 4–16 cm, and the distance to nearest occupied patch can be longer, up to 1.4 km. This study supports earlier findings in other regions, suggesting that a network of adjacent patches with a high food plant density and a vegetation height within the preferred threshold, despite their size, is an apparent conservation goal.     

Chromosome number variation within Philaethria butterflies (Lepidoptera: Nymphalidae,Heliconiini)

The haploid chromosome number of the South American butterfly Philaethria dido varies from 12 to 88. Eight different numbers have been found in this species complex. The related Ph. pygmalion and Ph. wernickei usually show only n=29, a very frequent number in the Lepidoptera; numbers of n=15 and n=21 for these species need confirmation. The most common chromosome number for Ph. dido is also the highest, n=88, and is found in many parts of northern and central Brazil on the Amazon river and its tributaries, as well as adjacent parts of other countries. The other numbers were observed mainly in northern South America and along the east coast. Two very different numbers were found together in four localities. We did not find specimens with meiotic features suggesting hybridization between individuals with different chromosome numbers. The diverse numbers in Ph. dido may belong to good sibling species, distinguishable externally by very minor characters. Since Ph. dido is a very primitive species in the tribe Heliconiini, dating probably from the early Tertiary, it probably has had many opportunities to undergo divergent chromosome evolution in isolation. Its strong, high flight and broad ecological valence would then permit rapid spreading out and coexistence of different chromosome forms, which in some cases have been noted to show diverse behaviour in the field.Dedicated to Professor Hans Bauer on the occasion of his eightieth birthday     

蛱蝶翅鳞片的超微结构观察

对我国东北地区典型常见蛱蝶科15属20种蝴蝶翅鳞片的超微结构进行了扫描电镜观察。结果显示：蛱蝶翅鳞片形态上可分为窄叶形、阔叶形和圆叶形3种,鳞片长65～135 μm,宽35～85 μm,间距48～112 μm。蛱蝶翅鳞片的超微结构可分为拱桥形、棋盘形和筛孔形3 种。拱桥形结构和棋盘形结构比较接近,二者与筛孔形结构差异较明显。在已观察的种类中,线蛱蝶属红线蛱蝶翅鳞片上的纵肋突起最小（200 nm×300 nm）,闪蛱蝶属柳紫闪蛱蝶翅鳞片上的纵肋突起最大（590 nm×560 nm）。鳞片具有相似的形状、结构和排列,尤其是同属蝴蝶翅鳞片超微结构的形状和尺寸差异较小,表明它们之间的亲缘关系接近。     

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

Microsatellite 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.     

Phylogeny of Polygonia, Nymphalis and related butterflies (Lepidoptera: Nymphalidae): a total-evidence analysis

We investigated the phylogeny of butterflies in the tribe Nymphalini sensu Harvey 1991, comprising the genera Vanessa, Cynthia, Bassaris, Aglais, Inachis, Nymphalis, Polygonia, Kaniska, Antanartia, Hypanartia, Symbrenthia, Mynes and Araschnia . Evidence from the mitochondrial gene ndl, the nuclear gene 'wingless' and from morphology/ ecology/behaviour were used separately and combined to analyse relationships. Phylogenies based on the different types of data agreed in many aspects of basic topology. We show that an analysis of only wing pattern characters (based on Nijhout's homology system) results in a topology broadly similar to the one resulting from analysis of the complete matrix. We found support for a monophyletic Nymphalini, where Hypanartia may be the sister clade to all other genera. Mynes, Symbrenthia and Araschnia together seem to form another basal clade. Evidence presented gives only moderate support for a monophyletic Vanessa in the wide sense, including also Cynthia and Bassaris , but strong support for the monophyly of the largely holarctic clade Aglais + Inachis + Nymphalis + Polygonia + Kaniska + Roddia . Within the latter group there is strong support for a clade consisting of Aglais + Inachis and for a second clade which includes Nymphalis, Polygonia (and its sister clade, the monotypic Kaniska) as well as Roddia l-album (= Nymphalis vaualbum ). As a consequence of this topology, Aglais is recognized as a taxon separate from Nymphalis . We present a hypothesis of species relationships within the focal group of genera. We also analyse and discuss the implications of excluding or including ecological data in phylogenetic tree construction, when the tree is to be used for studies in phylogenetic ecology.