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.     

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.     

Host plant and habitat preference of the endangered Euphydryas maturna (Lepidoptera: Nymphalidae): evidence from northern Europe

1. The aim of the present study was to evaluate host plant and habitat preferences in the Estonian populations of Euphydryas maturna, a regionally polyphagous but often locally specialised butterfly endangered in most parts of its European range. 2. Laboratory trials suggested that Fraxinus excelsior, Viburnum opulus and Melampyrum pratense are plants recognised by ovipositing females as potential hosts. These plants also supported development of the larvae, with the poorest growth performance on M. pratense. 3. Both a transect count‐based habitat occupancy analysis and a country‐wide landscape occupancy analysis revealed the abundance of F. excelsior as the primary determinant of the occurrence of E. maturna. In contrast, the occurrence of E. maturna was not associated with habitats colonised by M. pratense. 4. The results suggest that European ash, F. excelsior, is the main, if not the only, host plant of E. maturna in Estonia. The use of V. opulus cannot be excluded, although, due the relative scarcity of this plant, an important role for it as a determinant of the distribution of E. maturna is unlikely. Melampyrum pratense is not likely an alternative host of E. maturna in Estonia, which contrasts with the situation in neighbouring Finland. 5. This study adds to the evidence of geographical differences in host specialisation in melitaeine butterflies. The results imply that conservation actions should focus on securing the favourable status of the locally used host plant; the populations of F. excelsior are currently threatened by a fungal disease, ash dieback.     

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     

Within population variation in the demography of Speyeria mormonia (Lepidoptera: Nymphalidae)

The adult demography of a population of Speyeria mormonia (Lepidoptera: Nymphalidae) near Gothic, Colorado USA was studied for four years. Values of demographic parameters, including survival, dispersal and sex ratio, varied yearly, without major changes in density. Shift in sex ratio with constant population numbers entails variation in effective population size among years. Flight period, catchability, relationship between body size and eclosion date, and significance of correlation between weather and daily survival rates differed between the sexes. Sexual dimorphism in correlation of wheather with survival rate may partially explain the lack of yearly variation in total population size. There was no significant sexual dimorphism in large scale habitat use, or in dispersal, except in 1982.
The demography of this insect is compared to that of related species of Argynnis and heliconiines, from the old world temperate zone and new world tropics, respectively. While some differences are observed which can be ascribed to differences in length of the growing season, many patterns are similar. This implies conservation of demographic traits, even across broad habitat and geographic areas.     

A chromosome study ofCharaxes jasius L. (Lepidoptera,Nymphalidae)

The haploid chromosome complement ofCharaxes jasius is n=25 in both males and females. The chromosomes in the mitotic metaphase are rod⁻ and dot-shaped; the primary constriction, even after treatment with hypotonic solution, is lacking in them. Meiosis in females is achiasmatic. This research was supported by a grant from M.P.I. 40%-1984.     

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.     

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.     

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。     

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.     

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

In 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).     

Phylogeny of the Nymphalidae (Lepidoptera)

A generic-level phylogeny for the butterfly family Nymphalidae was produced by cladistic analysis of 234 characters from all life stages. The 95 species in the matrix (selected from the 213 studied) represent all important recognized lineages within this family. The analysis showed the taxa grouping into six main lineages. The basal branch is the Libytheinae, with the Danainae and Ithomiinae on the next branch. The remaining lineages are grouped into two main branches: the Heliconiinae-Nymphalinae, primarily flower-visitors (but including the fruit-attracted Coeini); and the Limenitidinae (sensu strictu), Biblidinae, and the satyroid lineage (Apaturinae, Charaxinae, Biinae, Calinaginae, Morphinae, Brassolinae, and Satyrinae), primarily fruit-attracted. Data partitions showed that the two data sets (immatures and adults) are very different, and a partitioned Bremer support analysis showed that the adult characters are the main source of conflict in the nodes of the combined analysis tree. This phylogeny includes the widest taxon coverage of any morphological study on Nymphalid butterflies to date, and supports the monophyly and relationships of most presently recognized subgroups, providing strong evidence for the presently accepted phylogenetic scheme.     

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.

     

Spatial and temporal variation in flight morphology in the butterfly Melitaea cinxia (Lepidoptera: Nymphalidae)

Some studies on insects have found a relationship between habitat structure and investment in dispersal-related traits. In this study we compared the morphology of the butterfly Melitaea cinxia from five sites on the large Baltic island Öland that differed markedly in degree of fragmentation and size. Both wild-caught adults and individuals reared in a split-plot design were compared. We found significant site differences in size-adjusted thorax mass and total body mass. Male thorax mass was on average larger among the sites with the highest degree of habitat fragmentation. However, due to significant sex–site interactions, males and females may have adapted differently to the habitat fragmentation. Using museum specimens, we also analysed changes in morphology, finding an increase in size (measured as head and thorax width) over time. Thorax width appears to have increased among females and decreased among males. Possible explanations include increasing fragmentation of the landscape and changes in population density.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 445–453.     

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.     

A fake Heliconius (Lepidoptera, Nymphalidae)

Four ‘doubtful species’ are listed in catalogues of the genus Heliconius. All are non–existent: (1) H. Carolina (Herbst) is a forgery made from trimmed fragments of at least three butterflies, the two identifiable ones both beingH. erato; (2) H.cinereo–fuscus (Goeze) is a faded specimen of H.ricini; (3) H.arcuatus (Goeze) is so inadequately described that its identity cannot be determined, although it is apparently some species of heliconiine or ithomiine;(4)H.euclea (Godart) is the valid senior name of an ithomiid in the genus Hypothyris.     

Seasonal variation in mate-locating activity in the desert hackberry butterfly (Asterocampa leilia; Lepidoptera: Nymphalidae)

Males of the desert hackberry butterfly occupy and defend encounter sites used in mate location. This activity is restricted to about 3 h each day, usually between 0800 and 1100. We monitored daily and seasonal changes in the number of males perched at encounter sites along a census route on 32 days during 1991 and 1992. The results indicate that males are active earlier in the mornings during summer than in spring or fall. However, they were not active at all times when air temperature and solar altitude would appear to permit perch site occupation. Daily and seasonal shifts in air temperature and solar radiation are not a sufficient proximate explanation for patterns of male behavior. Over a wide range of temperatures, females most often eclose around 0800, which suggests that the timing of female availability determines when males are active within the broad limits imposed by thermoregulatory requirements.     

Phylogenetic relationships of Phyciodes butterfly species (Lepidoptera: Nymphalidae): complex mtDNA variation and species delimitations

Abstract. Mitochondrial DNA variation was studied in the butterfly genus Phyciodes (Lepidoptera: Nymphalidae) by sequencing 1450 bp of the COI gene from 140 individuals of all eleven currently recognized species. The study focused on four species in particular that have been taxonomically difficult for the past century, P. tharos , P. cocyta , P. batesii and P. pulchella . A cladistic analysis of ninety-eight unique haplotypes showed that Phyciodes forms a monophyletic group with P. graphica as the most basal species. Of the three informal species groups described for Phyciodes , only one (the mylitta -group) is unambiguously monophyletic. Within the tharos -group, seven well supported clades were found that correspond to three taxa, P. tharos , P. pulchella and a grade consisting of P. cocyta and P. batesii haplotypes interdigitated with each other. None of the clades is formed exclusively by one species. The patterns of haplotype variation are the result of both retained ancient polymorphism and introgression. Introgression appears to be most common between P. cocyta and P. batesii ; however, these two species occur sympatrically and are morphologically and ecologically distinct, suggesting that the level of current introgression does not seem to be enough to threaten their genetic integrity. The results indicate that mitochondrial DNA sequences must be used with great caution in delimiting species, especially when infraspecific samples are few, or introgression seems to be rampant.     

芒蛱蝶西藏一个新纪录及川藏地区黄缘蛱蝶的历史记录

在2017年6-7月对藏东南地区蝶类资源进行调查时,于墨脱县采集到了2枚芒蛱蝶Euripusnyctelius(鳞翅目:蛱蝶科)标本,经鉴定为西藏新纪种,同时也是首次在西藏发现芒蛱蝶属;于察隅县采集到1枚黄缘蛱蝶Nymphalisantiopa(鳞翅目:蛱蝶科),兹此列举诸多川藏地区黄缘蛱蝶的历史采集记录;并结合西藏处于古北区和东洋区生物交界带的地理位置,对在西藏发现两种蝴蝶的特殊意义进行了简要讨论。