Towards a theory of the evolution of butterfly colour patterns under directional and disruptive selection

Two general models for the transspecific evolution of butterfly colour patterns are advanced: directional selection acting equally on both sexes, and disruptive selection involving periods of polymorphism. To consider possible outcomes of me latter process, a morphism notation based on an integrated classification for polymorphism and sexual dimorphism is developed. This notation is used to examine the properties of all morphism transformations possible from the minimal expressions of the nine morphism categories, as reached through defined minimum step changes. The significance of such pathway models is analysed in terms of general properties of butterfly polymorphism. The potential use of pathway models in evolutionary studies is briefly discussed, mainly with respect to phylogenetics, and ideas on the evolution of genetic dominance.     

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.     

Biogeography of milkweed butterflies (Nymphalidae: Danainae) and mimetic patterns on tropical Paciflc archipelagos

Distributions of danaine butterfly species and associated mimetic patterns were compared among fifteen archipelagos of the tropical Pacific Ocean, and within five major archipelagos (the Bismarcks, Fiji, East and West Solomon Islands, and Vanuatu). Using both simple and stepwise linear regression analysis, variation in the total number of danaine species and number of mimetic patterns was assessed with respect to island size, isolation and elevation. Relative to interarchipelago distributions, the distribution of danaine species and number of mimetic patterns on islands within archipelagos exhibited less dependence upon interisland distance and island area. Geographical features influencing the number of mimetic patterns were similar to those of danaines as a whole. Analysis of residuals from stepwise linear regression suggested that factors influencing danaine distributions were different from those for non-danaine butterflies. This result is consistent with the hypothesis of enhancement of danaine species establishment through Müllerian mimicry, although other factors such as host plant availability and similar habitat use may also be important.     

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.

     

First chromosome scale genomes of ithomiine butterflies (Nymphalidae: Ithomiini): Comparative models for mimicry genetic studies

The ithomiine butterflies (Nymphalidae: Danainae) represent the largest known radiation of Müllerian mimetic butterflies. They dominate by number the mimetic butterfly communities, which include species such as the iconic neotropical Heliconius genus. Recent studies on the ecology and genetics of speciation in Ithomiini have suggested that sexual pheromones, colour pattern and perhaps hostplant could drive reproductive isolation. However, no reference genome was available for Ithomiini, which has hindered further exploration on the genetic architecture of these candidate traits, and more generally on the genomic patterns of divergence. Here, we generated high-quality, chromosome-scale genome assemblies for two Melinaea species, M. marsaeus and M. menophilus, and a draft genome of the species Ithomia salapia. We obtained genomes with a size ranging from 396 to 503 Mb across the three species and scaffold N50 of 40.5 and 23.2 Mb for the two chromosome-scale assemblies. Using collinearity analyses we identified massive rearrangements between the two closely related Melinaea species. An annotation of transposable elements and gene content was performed, as well as a specialist annotation to target chemosensory genes, which is crucial for host plant detection and mate recognition in mimetic species. A comparative genomic approach revealed independent gene expansions in ithomiines and particularly in gustatory receptor genes. These first three genomes of ithomiine mimetic butterflies constitute a valuable addition and a welcome comparison to existing biological models such as Heliconius, and will enable further understanding of the mechanisms of adaptation in butterflies.     

Non‐target habitat exploitation by Trichogramma brassicae (Hym. Trichogrammatidae): what are the risks for endemic butterflies?

Abstract 1 Trichogramma brassicae Bezdenko is inundatively released against the European corn borer in Switzerland. Because parasitoids dispersing from the release fields might pose a threat to native butterflies, the searching efficiency of T. brassicae was investigated in nontarget habitats. 2 In field studies, T. brassicae was released at rates of 120 000 females/ha. Parasitism of sentinel Ephestia kuehniella egg clusters was 1.6–3.6% in meadows and 2.0–4.0% in flower strips. The respective figures were 57.6–66.7% and 19.2–46.9% in maize, significantly higher than the parasitism rates in the nontarget habitats. Experiments carried out in small field cages confirmed these results: Again, significantly higher parasitism rates were found in maize compared to meadows and flower strips, and also compared to hedgerows (in sleeve cages). 3 To elucidate potential factors underlying the low searching efficiency in nontarget habitats, the behaviour of individual T. brassicae females was investigated on four meadow plants comparatively to maize and a filter paper control. Mean (±SE) walking speed on maize was 2.2 ± 0.2 mm/s, similar to three of the plants tested and filter paper but significantly higher than on Trifolium pratense (0.85 mm/s). A higher turning rate was found on T. pratense, Viola wittrockiana and Plantago lanceolata, in contrast to the longer leaved maize and Alopecurus pratensis. The number of wasps leaving the plant within the observation period differed significantly between plant species, and was twice as high for T. pratense (and the filter paper control) compared to the other plant species. 4 In a choice experiment carried out in a climate cabinet with all five host plant species in cages, we obtained the highest parasitism rates on maize and the lowest parasitism on T. pratense, thus confirming the behavioural observations. 5 In conclusion, there is evidence for a decreased searching efficiency on plants in nontarget habitats compared to maize. However, the data explain only part of the differences found between parasitism in maize compared to nontarget habitats. Other factors, such as the structural complexity of a habitat, may also play a role. We conclude that the risk for butterfly populations in the tested nontarget habitat due to mass released T. brassicae is low.     

Marginality: scale determined processes and the conservation of the British butterfly fauna

Marginality describes the impact that environmental and landscape factors have in decreasing the probability of population survival and persistence. It may be imposed by extreme conditions or resource scarcity. Typically, it affects populations at the range edge but can also affect populations within the core of ranges, and produces a number of symptoms: characteristically demographic, but also morphological, physiological, biochemical and genetic. In this paper, the causes and effects of marginality on British butterflies are compared in edge and centre of range populations. Issues of temporal and spatial scales are examined, as is the relevance of marginality to the conservation of single and multiple species populations. The recognition of marginality questions the appropriateness of many so-called spatially realistic models of populations and highlights areas of research which have hitherto been ignored. Projected changes in land use and climate have implications for marginality in core and peripheral populations; in view of this, current scales of mapping are found to be unsuitable for monitoring fragmentation and the increasing marginalization of butterfly species in the British landscape.     

Variation in cuticular lipid profiles of black butterflies of the genus Papilio in Japan

Insect (epi)cuticular lipids characterize sex and species and often play an important role in mating behavior. We previously revealed that two black-colored swallowtail butterflies, Papilio polytes and Papilio protenor, show sexual dimorphism and species specificity in cuticular lipid composition and that P. polytes males use specific monoene components for mate discrimination. These findings suggest that their closely related species may have different profiles of cuticular lipids. We examined the cuticular lipid compositions of five Papilio species (P. bianor, P. maackii, P. helenus, P. macilentus, and P. memnon), closely related and sympatric to P. polytes and P. protenor, and discussed whether it is possible to discriminate between sexes, and between species based on their chemical profiles. The cuticular lipids consist mainly of C23–C31 aliphatic hydrocarbons, in which n-tricosane, n-heptacosane, and n-nonacosane are predominant. Several aliphatic ketones, aliphatic acids, and oxygenated terpenoids were also identified as major components shared by several species. There were no components exclusive to a particular species. Conspecific males and females shared most of the components but were mostly distinguishable based on their composition. Moreover, P. helenus males, P. polytes females, and P. protenor females had two different phenotypes of lipid composition. Although unrelated to the genetic lineage, the seven species were classified into four clusters based on their lipid profiles. The first cluster was composed of only P. memnon. The other six species were broadly classified into three clusters consisting of subclusters for each species: 1) P. polytes, P. helenus, P. macilentus, and several P. protenor females; 2) P. bianor and P. protenor; and 3) P. maackii and several P. helenus males. These results indicate that cuticular lipid profiles characterize the species and sex of the Papilio species and may be responsible for mate discrimination among them.