Experimental Density-Related Emigration in the Cranberry Fritillary Boloria aquilonaris

Studies on sedentary butterflies often show that most individuals cover only short distances and rarely leave their original patch. An important question for metapopulation dynamics and conservation is “When and why do sedentary species disperse?” To imitate high population densities I harassed individuals of the sedentary butterfly Boloria aquilonaris with a dummy and recorded the time that passed until they emigrated from their original bogpatch of 800 m². Harassed individuals emigrated significantly sooner than unharassed ones. Repeated harassment following their return to the marked patch led to an increase in the residence times of these individuals outside the patch. Individuals receiving a potentially detrimental stimulus (e.g., harassment) are more likely to emigrate sooner than other individuals. The results support the hypothesis that emigration is density dependent in this butterfly, such that gene flow and colonization may be episodic.     

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.     

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.     

Boloria phylogeny (Lepidoptera: Nymphalidae): tentatively reconstructed on the basis of male and female genitalic morphology

Abstract. The phylogeny of Boloria s.l. is reconstructed on the basis of sixty-three characters from the genital morphology of both sexes of twenty-nine taxa. Three unique and several homoplasious autapomorphies support the monophyly of Boloria. The genus is divided into two well-supported clades, one comprising the subgenera Boloria s.str. and Proclossiana and one comprising the subgenus Clossiana. Although the phylogeny of Clossiana remains largely uncertain, perhaps due to a rapid and recent speciation in some groups, it can be divided into two groups. A more subordinate group (the polaris group) appears well supported by a highly specialized phallus. Morphological evidence supports a clade comprising B.(C.) chariclea and both Palaearctic and North American B.(C.) titania populations. Possibly these taxa form a superspecies complex where B.(C.) chariclea is paraphyletic with regard to the different B.(C.) titania populations: molecular studies will be needed to test this hypothesis.     

Extensive clonality in the endangered shrub Haloragodendron lucasii (Haloragaceae) revealed by allozymes and RAPDs

The occurrence of clonality in threatened plants can have important implications for their conservation. In this study, allozymes and RAPDs were used to determine the extent of clonality in the endangered shrub Haloragodendron lucasii (Haloragaceae), which is known from only four sites within an 8 km range. Allozyme markers identified only six multilocus genotypes among the 53 ramets sampled across the four sites, although a total of 54 different genotypes were possible with the three polymorphic allozyme loci detected. The polymorphic bands detected in the RAPD analysis were capable of producing 2⁴⁶ genotypes, but again only six multilocus genotypes were delineated. The allozyme and RAPD data were congruent at three of the four sites. At the fourth site two genotypes were detected by each marker; however, once combined, three multilocus genotypes were observed. The probabilities that the observed number of replicates of each combined allozyme and RAPD genotype could be generated by sexual reproduction were less than 10^–18, leaving little doubt that clonality is the explanation for the observed patterns of genotypes. The genetic conclusions are supported by root excavations which show potential for vegetative reproduction and the observation of no sexual reproduction in the species. The recognition of extensive clonality in H. lucasii has had immediate implications for the conservation management of the species and resulted in changes to the management priorities for the species. Thus it is clear that appropriate genetic studies can play an important role in the management of threatened species.     

Population demography of alpine butterflies: Boloria pales and Boloria napaea (Lepidoptera: Nymphalidae) and their specific adaptations to high mountain environments

High mountain ecosystems are extreme habitats, and adaptation strategies to this ecosystem are still poorly understood in most groups. To unravel such strategies, we performed a MRR study in the Hohe Tauern National Park (Salzburg, Austria) with two nymphalid butterfly species, Boloria pales and B. napaea. We analysed their population structure over one flight period by studying the development of population size and wing wear. B. pales had more individuals and a higher survival probability than B. napaea; the sensitivity to extreme weather conditions or other external influences was higher in B. napaea. We only observed proterandry in B. pales. Imagines of both species survived under snow for at least some days. Additionally, we observed a kind of risk-spreading, in that individuals of both species, and especially B. pales, have regularly emerged throughout the flight period. This emergence pattern divided the population's age structure into three phases: an initial phase with decreasing wing quality (emergence > mortality), followed by an equilibrium phase with mostly constant average wing condition (emergence = mortality) and a final ageing phase with strongly deteriorating wing condition (mortality » emergence). Consequently, neither species would likely become extinct because of particularly unsuitable weather conditions during a single flight period. The observed differences between the two species suggest a better regional adaptation of B. pales, which is restricted to high mountain systems of Europe. In contrast, the arctic-alpine B. napaea might be best adapted to conditions in the Arctic and not the more southern high mountain systems. However, this needs to be examined during future research in the Arctic.     

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.     

Host plant relationships of an endangered butterfly, Lopinga achine (Lepidoptera: Nymphalidae) in northern Europe

The aim of the present study was to evaluate—in a geographic perspective—the role of host plant as a determinant of habitat quality for Lopinga achine, a satyrine butterfly endangered over much of its European range. Laboratory trials were performed to record host choices made by the ovipositing females as well as by neonate larvae. In rearing experiments, growth performance and mortality on different host plants was determined. Oviposition was found to be indiscriminate but larvae were shown to be able to choose between host plants, with the choices made broadly consistent with growth performance of the larvae on particular hosts. Nevertheless, most grasses and sedges offered were found to support larval development reasonably well. No clear superiority of the previously suggested primary host plant Carex montana could be shown. Importantly, no differences in host plant relationships were found between the populations of Sweden, western Estonia and eastern Estonia. In particular, the larvae originating from eastern Estonian populations developed on C. montana equally well even if the plant is absent from their native habitat. In the context of species conservation, one should conclude that L. achine is polyphagous enough on various grasses and sedges so that the presence of any particular host species cannot be a critical component of habitat quality. Nevertheless, some preference to broad- and soft-leaved hosts, as well as sensitivity to host wilting, may partly explain the butterfly’s preference to moist forest habitats, further emphasizing the central role of habitat management in the conservation practice of this species. In turn, the absence of ecological differences between geographic populations should enable conservationists to successful transfer their experience across national boundaries.     

Adult movements between populations in the specialist butterfly Proclossiana eunomia (Lepidoptera, Nymphalidae)

Abstract.

• 1 The structure of local populations of a monophagous butterfly, the bog fritillary Proclossiana eunomia, was studied in a complex of suitable habitat patches separated by spruce plantations or fertilized pasture.
• 2 An unexpected high level of adult movements between habitat patches was detected by a mark—release—recapture technique. Local populations were connected by adult movements across unsuitable habitats, leading to a meta-population structure.
• 3 This evidence of the metapopulation structure of a specialist butterfly challenges the supposed relationship between habitat specialization and closed, isolated populations.
• 4 Males and females of P.eunomia exhibited different spatial behaviours; females were more likely to emigrate and dispersed further than males. These differences in spatial behaviour are related to the mating system.

     

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.     

Patterns of genetic and morphometric differentiation in Melitaea (Mellicta) athalia (Lepidoptera: Nymphalidae)

Patterns of genetic and morphometric differentiation were surveyed in Melitaea (Mellicta) athalia populations of the Carpathian Basin. This species has a wide distribution and exists under a wide variety of ecological conditions. It has two ecotypes in Hungary: with either one or two broods per year. It is of particular interest to reveal the main factors driving differentiation patterns in this species. Samples in our study were obtained from five Hungarian and one Transylvanian (Romanian) regions. Enzyme polymorphism, wing characters and male external genitalia were analysed using traditional morphometric methods. Statistical methods were optimized to compare morphological and genetic data. The results of genetic surveys revealed a clear regional pattern of differentiation in M. athalia. Moreover, the results of principal component analysis, Bayesian clustering and the dendrogram all suggested that the regions can be classified into two groups corresponding to the East or West zones of the Carpathian Basin. In contrast, differentiation between the two ecotypes was less expressed in the genetic variation of M. athalia. Results of the analyses conducted on phenotypic variation also suggested a regional pattern for both sets of morphometric characters (wings and external genitalia). At the same time, neither East–West regional division nor ecotype differentiation was detected in the morphometric studies. In sum, our analyses confirmed that both genetic and phenotypic variations of M. athalia exhibit a regional pattern rather than the differentiation between the two ecotypes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

The species of the genus Zethera Felder (Lepidoptera: Nymphalidae,Satyrinae)

The butterfly genus Zethera Felder, endemic to the Philippines and northern Sulawesi, is shown to include six species, four of which comprise the Z. pimplea (Erichson) superspecies. Type data is given for all nominal taxa, and a key to the species recognised is presented, together with data on their variation and distribution. Attention is drawn to the zoogeographic subregions of the Philippine islands, as originally described by Semper, and illustrated by the Zethera species. Their mimicry is also noted, with particular reference to its sex-limitation in the Z. pimplea superspecies, and non-limitation in Z. incerta (Hewitson) and Z. hestioides C. & R. Felder. The need to demonstrate whether or not Zethera represents a true monophyletic unit is briefly discussed.     

Ithomiini Butterflies (Lepidoptera: Nymphalidae) of Antioquia, Colombia

Colombia is one of the most biodiverse countries on the planet. However, economic and scientific investment in completing inventories of its biodiversity has been relatively poor in comparison with other Neotropical countries. Butterflies are the best studied group of invertebrates, with the highest proportion of known to expected species. More than 3,200 species of butterflies have been recorded in Colombia, although the study of the still many unexplored areas will presumably increase this number. This work provides a list of Ithomiini butterflies collected in the department of Antioquia and estimates the total number of species present, based on revision of entomological collections, records in the literature and field work performed between 2003 and 2011. The list includes 99 species and 32 genera, representing 27% of all Ithomiini species. We report 50 species of Ithomiini not formerly listed from Antioquia, and found the highest diversity of ithomiine species to be at middle elevations (900–1,800 m). The mean value of the Chao2 estimator for number of species in Antioquia is 115 species, which is close to a predicted total of 109 based on known distributions of other Ithomiini not yet recorded from the department. Nine species are potentially of particular conservation importance because of their restricted distributions, and we present range maps for each species. We also highlight areas in Antioquia with a lack of biodiversity knowledge to be targeted in future studies. This paper contributes to mapping the distribution of the Lepidoptera of Antioquia department in particular and of Colombia in general.     

Influence of spatial structure on genetic isolation in Plebejus argus populations (Lepidoptera: Lycaenidae)

Populations of Plebejus argus were sampled in southwest Finland, both on the mainland and on islands, and in and around the Do?ana National Park in southwest Spain. A total of 453 individuals coming from 14 locations were investigated using allozyme electrophoresis on a total of 10 polymorphic allozyme loci. Contrary to earlier studies, all conducted in Britain, our samples showed little differentiation between sampled locations. In Spain, the populations of the Dona?a area showed no differentiation despite being up to 36 km apart; only the population to the south of the Guadalquivir river showed a significant difference to the others. In Finland the population on one island showed marked genetic differentiation from all the others, which showed little or no difference from each other. No isolation-by-distance effect could be detected in either system. We hypothesise that emigration-immigration events are more frequent in the Spanish and Finnish populations than in the British ones. We did, however, find two isolated populations, one in Spain and one in Finland; both were small and geographically isolated and shared evident drift.     

Glands, muscles and genitalia. Morphological and phylogenetic implications of histological characters in the male genitalia of Fritillary butterflies (Lepidoptera: Nymphalidae: Argynnini)

The histology and internal morphology of the male valve in a broad sample of Fritillary butterflies are examined. A large internal ridge to which the extensor of the valve muscle fastens is probably an autapomorphy for the tribe Argynnini. A large glandular pocket is found in the valves of all examined Heliconiinae, but is missing in other examined Nymphalidae. The pocket shows considerable variation between the higher taxa. In the higher Fritillaries (subtribe Argynnina) the pocket probably has an effective closing mechanism allowing the male to store a considerable amount of glandular secretion. It has earlier been suggested that the secreta from the pocket glands are transferred to the female during copulation, where they probably have an antiaphrodisiac effect, preventing other males from mating a once mated female. All examined male Fritillaries also have glands in the anellus, the terminal membrane between the valves. In Boloria these glands are extremely well developed and abundant, a condition that is probably an autapomorphy for that genus. The obtained characters are tested against a known phylogeny of the tribe Argynnini.     

Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini)

Species-level phylogeny of the Butterfly tribe Argynnini is established based on 141 characters derived from wing and genitalia morphology of both sexes. The Argynnini can be divided into three subtribes; Yrameina comprising Yramea and Boloria s . l ., Argynnina comprising Prokuekenthaliella , Issoria , Brenthis , and all the 'large fritillary' species joined in the genus Argynnis s . l . and a new subtribe Euptoietina comprising only the genus Euptoieta . The classical genus Issoria s . l . is polyphyletic regarding Yramea and possibly paraphyletic regarding the two Afrotropic species baumanni and hanningtoni ; these two species are tentatively transferred to the old genus/subgenus Prokuekenthaliella . Surprisingly, one Afrotropic species, Issoria smaragdifera is closely related to the East Palaearctic Issoria species. A revised classification of Argynnini is proposed based on the obtained phylogeny. Studies of larval host plants based on the obtained phylogeny suggest that the ancestral Argynnini used Passiflora and Violaceae, but already the ancestor of Yrameina + Argynnina was probably specialized on Violaceae. Whereas the Boloria species have turned to other food plants such as Dryas , Vaccinium and Salix on several occasions, only Brenthis among the Argynnina use other host plants than Viola (mainly Rosaceae). The habit of laying eggs away from the food plant has probably evolved twice within Argynnina.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 627–673.     

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.     

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.     

Variation in thermally induced melanism in monarch butterflies (Lepidoptera: Nymphalidae) from three North American populations

• 1.As ectotherms, insects often experience varying temperatures throughout their life cycle, and some respond by becoming more or less melanistic (dark coloring) during development to increase or decrease thermal energy absorption as larvae or adults.
• 2.Monarch butterflies (Danaus plexippus) breed in temperate and tropical environments worldwide and are exposed to different average and extreme temperatures in different parts of their geographic range. In this study, we compared variation in thermally induced melanism among monarch butterflies from eastern and western North America and from South Florida.
• 3.We raised the progeny of wild-captured adult butterflies from these populations in a common garden experiment, rearing individuals in cold (19 °C), moderate (26 °C), and hot (32 °C) temperatures to examine population variation in larval and adult pigmentation.
• 4.Across all populations, monarch larvae developed the darkest coloration in the cold treatment and were lightest when reared in hot temperatures. Similar results were observed for measures of adult wing melanism, with the exception of adult females, which developed darker colored wings in warmer temperatures.
• 5.Significant population-level differences in average measures of melanism among larvae and adult butterflies were observed. Larvae from the eastern population became substantially darker in colder temperatures than S. Florida or western larvae. Western larvae were lightest overall, which might be adaptive to high temperatures experienced throughout portions of their summer breeding range. S. Florida larvae showed a lower response to cold temperatures relative to monarchs from either migratory population.
• 6.Population level differences were also observed for thermal responses in wing melanism, particularly among adult females. Moreover, we found significant family level effects for each measure of larval and adult melanism, pointing to a genetic basis or strong maternal effects influencing these traits in monarch butterflies.