Homologies in the colour patterns of the genus Heliconius (Lepidoptera: Nymphalidae)

The colour patterns of Heliconius butterflies are composed from a relatively simple set of pattern elements whose homologues are recognizable throughout the genus. Although Heliconius colour patterns look quite different from those of most nymphalids, these pattern elements are seen to derive from the generalized nymphalid groundplan. The differences arise primarily from the loss or positional shift of certain pattern elements, a high degree of fusion between individual pattern elements, and, in the forewing, asymmetries of the pattern elements relative to the wing-cell midline. The scheme of homologies we present is consistent with what is currently known about the comparative morphology and developmental physiology of colour pattern formation in Lepidoptera, and provides a framework for the interpretation of developmental, evolutionary and genetic studies in Heliconius.     

An analysis of the phenotypic effects of certain colour pattern genes in Heliconius (Lepidoptera: Nymphalidae)

The colour patterns of Heliconius butterflies are built up from an array of serially homologous pattern elements known as the nymphalid groundplan. An analysis of the phenotypic effects of ten genetic loci from H. melpomene and H. cydno reveals that each alters the expression either of a single element of the groundplan or of an entire row of serially homologous elements. Five of the ten loci affect the size (or presence/absence) of specific pattern elements, two affect the colour in which a pattern element is expressed, two affect pattern-inducing activity of the wing veins, and one appears to affect an overall threshold for pattern determination. Three of the ten loci have identical effects on homologues of the fore- and hindwing. We show that most of the apparently large and qualitative phenotypic effects of these genes can be readily explained by relatively small and quantitative changes in the dimensions or positions of specific pattern elements.     

The biogeography of the western Mediterranean: elucidating contradictory distribution patterns of differentiation in Maniola jurtina (Lepidoptera: Nymphalidae)

In the western Palaearctic, the Mediterranean zone is an important region where taxa and genes of thermophilous organisms are preserved during glacial stages and new clades are generated. This is achieved through the existence of refugia over Mediterranean Europe and North Africa, where organisms persisted and continued to evolve during the cold phases. However, it is not clear in detail how these refugia function for the maintenance of ancestral taxa, the evolution of new taxa, and as launching pads during postglacial colonizations of northern Europe. One outstanding issue is the incongruence of findings from different marker systems. For the butterfly Maniola jurtina, morphometry and allozyme data analyzed for populations scattered over Europe and North Africa show congruent patterns for Sicily and the Maghreb but produce discrepant results for the Italian mainland. This discrepancy between allozyme and morphological data can be explained by recent gene flow in the wake of postglacial range expansions and shifts. It is evident that colonization histories are far more complex than originally considered. We highlight different aspects of colonization and evolutionary history emerging from the joint use of different marker systems and advocate multiple uses of different markers in paleobiogeographic reconstructions to explore evolutionary events and colonization pathways. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 571–577.     

Biology and morphology of the immature stages of Hermeuptychia atalanta (Lepidoptera: Nymphalidae: Satyrinae)

This paper describes the morphology of the immature stages (egg, larva and pupa) and larval behavior of the butterfly Hermeuptychia atalanta, a common and widespread species of satyrinae in South America. Eggs are white and spherical, laid singly on leaves of grasses. The first instar has a light-green body and dark-brown head; all remaining instars present body and head capsule light green. Larvae have no spines, and general morphology is similar to other species of the subtribe Euptychiina.     

Colour pattern homology and evolution in Vanessa butterflies (Nymphalidae: Nymphalini): eyespot characters

Ocelli are serially repeated colour patterns on the wings of many butterflies. Eyespots are elaborate ocelli that function in predator avoidance and deterrence as well as in mate choice. A phylogenetic approach was used to study ocelli and eyespot evolution in Vanessa butterflies, a genus exhibiting diverse phenotypes among these serial homologs. Forty‐four morphological characters based on eyespot number, arrangement, shape and the number of elements in each eyespot were defined and scored. Ocelli from eight wing cells on the dorsal and ventral surfaces of the forewing and hindwing were evaluated. The evolution of these characters was traced over a phylogeny of Vanessa based on 7750 DNA base pairs from 10 genes. Our reconstruction predicts that the ancestral Vanessa had 5 serially arranged ocelli on all four wing surfaces. The ancestral state on the dorsal forewing and ventral hindwing was ocelli arranged in two heterogeneous groups. On the dorsal hindwing, the ancestral state was either homogenous or ocelli arranged in two heterogeneous groups. On the ventral forewing, we determined that the ancestral state was organized into three heterogeneous groups. In Vanessa, almost all ocelli are individuated and capable of independent evolution relative to other colour patterns except for the ocelli in cells ?1 and 0 on the dorsal and ventral forewings, which appear to be constrained to evolve in parallel. The genus Vanessa is a good model system for the study of serial homology and the interaction of selective forces with developmental architecture to produce diversity in butterfly colour patterns.     

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.     

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.     

A molecular phylogeny of the neotropical butterfly genus Anartia (Lepidoptera: Nymphalidae)

While Anartia butterflies have served as model organisms for research on the genetics of speciation, no phylogeny has been published to describe interspecific relationships. Here, we present a molecular phylogenetic analysis of Anartia species relationships, using both mitochondrial and nuclear genes. Analyses of both data sets confirm earlier predictions of sister species pairings based primarily on genital morphology. Yet both the mitochondrial and nuclear gene phylogenies demonstrate that Anartia jatrophae is not sister to all other Anartia species, but rather that it is sister to the Anartia fatima-Anartia amathea lineage. Traditional biogeographic explanations for speciation across the genus relied on A. jatrophae being sister to its congeners. These explanations invoked allopatric divergence of sister species pairs and multiple sympatric speciation events to explain why A. jatrophae flies alongside all its congeners. The molecular phylogenies are more consistent with lineage divergence due to vicariance, and range expansion of A. jatrophae to explain its sympatry with congeners. Further interpretations of the tree topologies also suggest how morphological evolution and eco-geographic adaptation may have set species range boundaries.     

‘After Africa’: the evolutionary history and systematics of the genus Charaxes Ochsenheimer (Lepidoptera: Nymphalidae) in the Indo‐Pacific region

The predominantly Afrotropical genus Charaxes is represented by 31 known species outside of Africa (excluding subgenus Polyura Billberg). We explored the biogeographic history of the genus using every known non‐African species, with several African species as outgroup taxa. A phylogenetic hypothesis is proposed, based on molecular characters of the mitochondrial genes cytochrome oxidase subunit I (COI) and NADH dehydrogenase 5 (ND5), and the nuclear wingless gene. Phylogenetic analyses based on maximum parsimony and Bayesian inference of the combined dataset implies that the Indo‐Pacific Charaxes form a monophyletic assemblage, with the exception of Charaxes solon Fabricius. Eight major lineages are recognized in the Indo‐Pacific, here designated the solon (+African), elwesi, harmodius, amycus, mars, eurialus, latona, nitebis, and bernardus clades. Species group relationships are concordant with morphology and, based on the phylogeny, we present the first systematic appraisal and classification of all non‐African species. A biogeographical analysis reveals that, after the genus originated in Africa, the evolutionary history of Charaxes in the Indo‐Pacific, in particular Wallacea, may be correlated with the inferred geological and climatic history of the region. We propose that Wallacea was the area of origin of all Charaxes (excluding C. solon) occurring to the east of Wallace's [1863] Line. The earliest Indo‐Pacific lineages appear to have diverged subsequent to the initial fragmentation of a palaeo‐continent approximately 13 million years ago. Further diversification in Indo‐Pacific Charaxes appears primarily related to climatic changes during the Pliocene and possibly as recently as the Pleistocene. Although both dispersal and vicariance have played important roles in the evolution of the genus within the region, the latter has been particularly responsible for diversification of Charaxes in Wallacea. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 457–481.     

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.

     

Species diversity and seasonal abundance in Charaxes butterflies (Nymphalidae)

Species diversity in Charaxes butterflies trapped in gardens at Freetown, Sierra Leone, is calculated by two different methods. There is little evidence of seasonal changes in diversity and in abundance.     

Pupal Melanization in Heliconius erato phyllis (Lepidoptera; Nymphalidae): Genetic and Environmental Effects

This paper deals with estimates of heritability and the effect of two different kinds of environment on the melanization of pupae in the butterfly Heliconius erato phyllis. The results for heritability showed greater differences when the offspring were regressed on male or female values for pupal colour. Pupal colour followed a system of discrete scores, from 2 for the light pupae to 5 for the darkest one. Estimate of the heritability (h²) was 0.44 when the average score of the siblings were regressed on the mid-parental value. When regressed on the male parent, h² was larger (0.54), being very low when regressed on the female parent (0.09). Estimate by the analysis of variance was also 0.44 when both male and female sibs were included in the analysis (for male sibs, h²=0.43 and for female sibs, 0.47). To test for the effect of the environment, each brood was divided in three, one being the control, the second subjected to a black environment when reaching the fifth instar and the third to a white environment. Those in the black environment originated dark pupae that scored 5 or 4; for those in the white environment, there was no difference with the controls. Caterpillars when entering the prepupal stage were also subjected to the black treatment (early and late prepupal stage, respectively, for treatments called black A and black B). A significant effect was observed only when early prepupal stage was subjected to the black environment.     

Phylogenetic relationships among genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA sequences

Cladistic relationships among genera and subtribes of Danaini (the milkweed butterflies) were inferred by analysis of data combined from five sources: morphology of adults and immature stages, and DNA sequences from three gene regions. The results corroborate and greatly increase support for prior hypotheses based on morphology alone. A new index summarizing incongruence among data partitions, the Partition Congruence Proportion (PCP), is introduced. The significance of the inferred pattern of phylogenetic relationships for comparative chemical ecology of milkweed butterflies is briefly discussed.     

Adaptive radiation and convergence in subdivisions of the butterfly genus Heliconius (Lepidoptera: Nymphalidae)

The process of adaptive radiation and convergence, usually regarded as a feature of macro-evolution, can be seen in the mimetic colour patterns of the butterflies within the confines of the South American genus Heliconius. This can be shown by dividing the genus into subgroups on the basis of adult, pupal and larval morphology: the theory that the mimicry between species results solely from close systematic relationships is thereby refuted, as members of the same morphological group can display widely divergent mimetic patterns, and conversely mutual mimics may belong to several different morphological groups. Various forms of parallel and convergent evolution are thought to account for the present pattern of mimicry, the process is known to start even before full speciation has taken place. A new subgenus (Neruda) is created to contain three atypical members of the genus.     

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, ?? , ??–??.     

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.     

A new species of Parapanteles Ashmead, 1900 (Hymenoptera: Braconidae: Microgastrinae) parasitic on Charaxes athamas (Drury) (Lepidoptera: Nymphalidae) in India

A new species of gregarious endoparasitoid, Parapanteles athamasae n. sp. (Hymenoptera: Braconidae), parasitising caterpillars of Charaxes athamas (Drury) (Lepidoptera: Nymphalidae) on the host plant Senegalia catechu (=Acacia catechu) (L.f.) Hurter & Mabb., is described from Maharashtra, India. Diagnostic characters of the new species include: propodeum with areola 0.93× longer than wide, legs yellow, hind tibia 4.30× as long as ovipositor, ovipositor sheaths exerted, first metasomal tergal plate 1.24× longer than wide, with coarse sculpture merging with longitudinal striations at 3/4 of the apical region. This is the first time a species of the family Nymphalidae Rafinesque is recorded in association with Parapanteles Ashmead, 1900. A key to the Indian species of Parapanteles based on females is also provided.     

Seasonal variation in larval host plants and consequences for Heliconius erato (Lepidoptera: Nymphalidae) adult body size

Abstract Adult body size, a key life history component, varies strongly within and between Heliconius erato phyllis (Lepidoptera: Nymphalidae) populations. In the present study, we determined whether seasonal variation in adult body size is temperature related and/or determined by seasonal changes of host plants (Passifloraceae) used by the larval stage. A population of H. erato phyllis located in a Eucalyptus plantation (Barba Negra Forest, Barra do Ribeiro County, Rio Grande do Sul State, Brazil) was sampled every 45 days from March 1997 to October 1998 to quantify seasonal variation in adult body size and use of larval host plants. In the laboratory, the effects of the following factors on adult body size were quantified: (i) host plant species (Passiflora misera or Passiflora suberosa); (ii) food quantity consumed by larvae (experimentally manipulated for each passion vine species); (iii) winter and summer temperatures (15 and 25°C, respectively); and (iv) the interaction between host plant species and temperature. Adults emerging during summer were larger than those emerging in other seasons. Female butterflies oviposited selectively on P. misera even when the dominant passion vine was P. suberosa. They only switched from using P. misera to P. suberosa during later autumn and winter, when P. misera vines were completely defoliated. The laboratory feeding trials with both passion vines showed a strong positive association between food quantity consumed by larvae and adult size. They also confirmed that adults are larger when their larvae are reared on P. misera than on P. suberosa. Temperature during larval development had no effect on H. erato phyllis adult size. Thus, seasonal variation of H. erato phyllis adult size in a given place is primarily determined by the availability and quality of host plant species used by the larval stage.     

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.     

Intraspecific variation in wing colour is related to larval energy reserves in monarch butterflies (Danaus plexippus)

The physiological basis for pigment synthesis in lepidopteran wing scales is well‐studied, although less is known about the reasons why individuals of the same species vary in pigmentation. Monarch butterflies (Danaus plexippus L.) show subtle variations in the shade of orange on their wings and this is known to predict flight ability and mating success. The present study tests the possibility that the shade of orange is associated with the amount of residual energy reserves carried over from the larval stage. Using monarchs reared in captivity under identical conditions (n = 207), the residuals of a regression of wing size and mass at eclosion, which indicate larval energy reserves, are obtained. This measure is positively related to adult longevity without feeding, indicating the importance of this reserve to the monarchs, as well as the value of the measure for this investigation. The shade of orange (i.e. hue) is determined on scanned wings using image analysis. Importantly, orange hue is predicted significantly by residual mass at eclosion (individuals with more mass are redder). The linkage between these traits may explain previous findings whereby redder monarchs fly further and mate more because both behaviours would be enhanced with greater energy stores. The findings of the present study add to a growing body of work showing how intraspecific variation in pigmentation has biological significance to monarchs, and possibly other butterflies. Although much remains to be investigated regarding the physiological underpinnings of this variation, the results of the present study indicate that future efforts should be rewarding.