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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

中国粉眼蝶属分类研究（鳞翅目：蛱蝶科：眼蝶亚科）

系统整理分布在中国的粉眼蝶属Callarge Leech,[1892]种类,已知1种4亚种,其中含1中国新纪录亚种Callarge occidentalis fansipana Monastyrskii,2005;通过外部形态和雄性生殖器特征研究,提出将其作为Callargesagitta fansipana Monastyrskii,stat.nov.阐述各亚种的主要特征及地理分布,并附有各亚种雄性外生殖器特征图和成虫照片.     

Wing pattern evolution and the origins of mimicry among North American admiral butterflies (Nymphalidae: Limenitis)

The evolution of wing pattern diversity in butterflies has emerged as a model system for understanding the origins and maintenance of adaptive phenotypic novelty. Admiral butterflies (genus Limenitis) are an attractive system for studying wing pattern diversity because mimicry is common among the North American species and hybrid zones occur wherever mimetic and non-mimetic wing pattern races meet. However, the utility of this system has been limited because the evolutionary relationships among these butterflies remain unclear. Here I present a robust species-level phylogeny of Limenitis based on 1911 bp of two mitochondrial genes (COI and COII) and 904 bp of EF1-alpha for all five of the Nearctic species/wing pattern races, the majority of the Palearctic species, and three outgroup genera; Athyma, Moduza (Limenitidini), and Neptis (Limenitidinae: Neptini). Maximum-likelihood and Bayesian analyses indicate that the North American species are a well-supported, monophyletic lineage that is most closely related to the widespread, Palearctic, Poplar admiral (L. populi). Within North America, the Viceroy (L. archippus) is the basal lineage while the relationships among the remaining species are not well resolved. A combined maximum-likelihood analysis, however, indicates that the two western North America species (L. lorquini and L. weidemeyerii) are sister taxa and closely related to the wing pattern subspecies of the polytypic Limenitis arthemis species complex. These results are consistent with (1) an ancestral host-shift to Salicaceae by the common ancestor of the Poplar admiral and the Nearctic admiral lineage, (2) a single colonization of the Nearctic, and (3) a subsequent radiation of the North American forms leading to at least three independent origins of mimicry.     

A new genus and species of Euptychiina (Lepidoptera: Nymphalidae: Satyrinae) from southern Brazil

This paper describes a new genus and a new species of Euptychiina from open grassland habitats (campos de cima da serra) in southern Brazil. The systematic position of this new taxon is discussed based on morphological and molecular data, and it is considered sister to Taydebis Freitas. Since the campos vegetation is considered endangered due to anthropogenic activities, this butterfly species deserves attention and should be included in future conservation plans for this biome.     

Carotenoids of butterfly models and their mimics (Lep: Papilionidae and Nymphalidae)

Carotenoid concentration in model swallowtail butterflies is exceptionally high; it is suggested that they may exert a photo-protective function in species storing nitrophenanthrenes. Carotenoid concentration in the viceroy is also high. Again, it is suggested that this represents a protective mechanism for Lepidoptera feeding on Salix which contains salicin. Such a mechanism would not be required by the monarch storing cardenolides.     

The danaid genus Tellervo (Lepidoptera, Nymphalidae)—a cladistic approach

Cladistic analysis suggests that the genus Tellervo comprises six species of which four are assignable to the zoilus complex, frequently sympatric with a fifth species, T. nedusia. The sixth species, T. jurriaansei , is very distinctive and highly localized. The presence of many well-differentiated races (17 are described as new) is discussed in relation to within-species structure, subspecies concepts and biogeography, with accompanying reviews of mimicry, early-stage biology, adult behaviour and taxonomy- seven synonyms are newly established and two neotypes designated. A key is given to separate the six species involved.     

Foodplant effects on colour morphs of Eumorpha fasciata caterpillars (Lepidoptera: Sphingidae)

Caterpillars of the hawkmoth Eumorpha fasciata are highly polymorphic for colour, with green, pink, and pink-and-yellow forms in the second through fourth instars, and green and multicoloured forms in the fifth instar. Four years of field censuses on four foodplant species determined that all morphs were found on all plant species; morph frequencies were homogeneous on each plant species over time; and morph frequencies differed consistently among plant species. When larvae were reared from eggs on three of the hostplant species in the laboratory, differences in morph frequencies paralleled the census results. Thus foodplant quality is one factor affecting larval colour in E. fasciata. A literature survey reveals that foodplant effects on larval colouration may be widespread in the family Sphingidae, but most reports are anecdotal rather than experimental. The implications of this mechanism of colour determination are discussed.     

Light absorption patterns of intact Rosa flowers in relation to the flower colour

Absorption curves of fresh, intact petals from 18 rose cultivars and 2 species were measured and compared with visual evaluations of their colours and there was a reasonable correlation. The in vivo maxima of anthocyanin absorption were in the range of 520–560 nm. Five patterns of absorption spectrum in the visible region were recognized: (a) maximum range ca. 520–535 rim (red roses); (b) as (a) but low absorbance (pink roses); (c) absorption pattern varying with age of flowers; (d) absorption at long wavelengths in blue roses due to co-pigmentation of cyanin, flavonols; (e) absorption of carotenoids and anthocyanins together in yellow, orange or orange red flowers.     

Shell colour polymorphism associated with substrate colour in the intertidal snail Littorina saxatilis Olivi (Prosobranchia: Littorinidae)

Littorina saxatilis Olivi (1792), the rough winkle, is highly polymorphic in shell colour. Shell colour frequencies were studied at six locations in south-western Wales, U.K., each at a geological contact between red sandstone and grey limestone or volcanic rock. At each site shell colour frequencies were determined in samples from the contact zone and on red or grey rock on either side. Highly significant associations were found between shell colour frequencies and substrate colour. Grey shells were always more common on grey rock than on red rock, and brown shells were usually more common on red than on grey rock, suggesting selection for cryptic colouration. Shell colour frequency differences were also found between replicate samples taken only 5 m apart from the same kind of rock, and between samples from the same kind of rock at the six study sites. These latter differences suggest that selection for camouflage is not the only factor involved in maintaining shell colour polymorphism in this species.