Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses

Adults of the Euthalia phemius complex, which is composed of three South‐East Asian nymphalid species, Euthalia phemius, Euthalia ipona, and Euthalia euphemia, were genetically analysed by examining mitochondrial and nuclear genes. The E. phemius complex was also examined morphologically, with particular emphasis on wing markings and male genitalia. No significant differences amongst the three species in the complex were detected with respect to either genetic distance or genital morphology. We therefore conclude that the three currently recognized Euthalia species belong to a single species. Accordingly, E. ipona is synonymized with E. phemius. Euthalia euphemia is treated as a subspecies of E. phemius. Type specimens of all taxa and a synonymic list for the E. phemius complex are also given. In addition, we briefly discuss the evolution and biogeography of the species complex. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 304–327.     

Phylogenetic relationships among Heliconiinae genera based on morphology (Lepidoptera: Nymphalidae)

Abstract.  Phylogenetic relationships among Heliconiinae genera are proposed based on early-stage and adult morphology. Parsimony analyses of forty-nine species in twenty-nine genera indicate that Heliconiinae can be divided into four main groups: (1) Pardopsis , Acraea and Actinote ; (2) Cethosia plus Neotropical genera; (3) Oriental genera and (4) fritillaries. Analyses of adult characters suggested that Oriental genera form a monophyletic group, whereas those of adult plus early-stage characters artificially split this group into three separate lineages. Our character set does not contain enough phylogenetic information to resolve relationships among fritillary genera with confidence, and further studies of this group are needed. The classification of the Heliconiinae is revised based on our results.     

A classification of Danaus butterflies (Lepidoptera: Nymphalidae) based upon data from morphology and DNA

Classification of the cosmopolitan butterfly genus Danaus (Nymphalidae: Danainae) is revised at subgeneric, specific and subspecific levels, combining for the first time mitochondrial and nuclear DNA sequence information with morphological data. Tree topologies based on the nuclear genome (allozymes, pheromone components, the morphology of all life history stages and nuclear DNA sequences), on the one hand, and mitochondrial DNA, on the other, are incongruent and challenge the current taxonomy of the genus. Although earlier classifications, based on adult morphology alone, are, in general, well supported by an analysis of total evidence, the mitochondrial phylogeny shows that the species D. chrysippus and its subgenus Anosia are deeply paraphyletic. Subspecies dorippus of D. chrysippus is the basal clade of the genus and is reinstated as the species D. dorippus. The former species D. plexaure is demoted to a subspecies of D. eresimus. The specific status of D. erippus, as distinct from D. plexippus, is tentatively supported. On the strength of the new data, division of the monophyletic genus Danaus s.l. into three subgenera Danaus s.s., Salatura and Anosia is unsustainable and is abandoned. Of the 15 terminal clades (taxa) of Danaus s.l. included in the study, 11 are species that broadly conform to the biological species concept. (The West Indian species D. cleophile, missing from our analysis, is the twelfth species). The remaining terminal clades are subspecies of D. chrysippus comb. nov. and D. dorippus stat. rev. Two sympatric Neotropical species, D. eresimus and D. gilippus, are morphologically distinct and sexually isolated but have nearly identical mitochondrial genomes. In contrast, two partially sympatric Palaeotropical species, D. chrysippus and D. dorippus, are cryptic species that share structural morphology and hybridize but have highly differentiated mitochondrial genomes. D. dorippus is polymorphic for two anciently diverged haplotypes and its history has possibly involved recombinational speciation and/or hybridism. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144 , 191?212.     

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.     

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.     

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.     

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.     

External morphology of adult Libythea celtis (Laicharting [1782]) (Lepidoptera: Nymphalidae)

Libytheinae is a morphologically unusual group of butterflies whose external anatomy has never been described in detail. The subfamily is considered basal within the diverse family Nymphalidae, and libytheines retain many plesiomorphic features, such as a functional female foreleg and smooth larva. In this paper, we describe and illustrate the adult morphology of Libythea celtis Laicharting, make comparisons to other nymphalid subfamilies, and homologize terms whenever possible. We describe major sclerites, sulci, and the presence of cranial and lateral pits on the tibia, the latter which appear to be previously unreported from the lepidopteran literature. When present, sexual dimorphism and geographic variation are noted. It is hoped that the present paper will help stabilize inconsistencies in morphological terms in nymphalid butterflies.     

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.     

Proboscis sensilla in Vanessa cardui (Nymphalidae, Lepidoptera): functional morphology and significance in flower-probing

 Morphology and distribution of the proboscis sensilla in Vanessa cardui have been investigated in order to contribute to the understanding of flower-probing behaviour in butterflies. The proboscis has a bend region approximately one-third of the length from the base. A short tip region is characterized by rows of intake slits leading into the food canal. Along the dorsal, lateral and ventral sides of the proboscis, sensilla trichodea, sensilla basiconica and sensilla styloconica are distributed in varying patterns depending on their distance from the b ase. The medial food canal bears one longitudinal row of sensilla basiconica only. The bristle-shaped sensilla trichodea are longer in the proximal region of the proboscis and become gradually shorter towards the tip. They are most frequent in number near to the bend region and near the beginning of the tip region. Sensilla basiconica arranged in longitudinal rows increase in number the more distal they are on the proboscis. The tip region is characterized by rows of sensilla styloconica on the dorsal side whereas the sensilla trichodea are mostly restricted to the ventral side. The ultrastructure suggests that the aporous sensilla trichodea function as mechanosensilla while the uniporous sensilla basiconica act as contact chemosensilla. The sensilla styloconica are regarded as bimodal contact chemo/mechanosensilla since their sensory cones are equipped with a single terminal pore and a tubular body at the base. The mouthpart sensilla appear to provide tactile cues on the positioning of the proboscis and on the degree of its insertion into a floral tube. Furthermore, they receive chemical stimuli on the availability of nectar and on the immersion status of the food canal. Accepted: 12 September 1997     

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.     

Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data

We have inferred the first empirically supported hypothesis of relationships for the cosmopolitan butterfly subfamily Satyrinae. We used 3090 base pairs of DNA from the mitochondrial gene COI and the nuclear genes EF-1alpha and wingless for 165 Satyrinae taxa representing 4 tribes and 15 subtribes, and 26 outgroups, in order to test the monophyly of the subfamily and elucidate phylogenetic relationships of its major lineages. In a combined analysis, the three gene regions supported an almost fully resolved topology, which recovered Satyrinae as polyphyletic, and revealed that the current classification of suprageneric taxa within the subfamily is comprised almost completely of unnatural assemblages. The most noteworthy findings are that Manataria is closely related to Melanitini; Palaeonympha belongs to Euptychiina; Oressinoma, Orsotriaena and Coenonympha group with the Hypocystina; Miller's (1968). Parargina is polyphyletic and its components group with multiple distantly related lineages; and the subtribes Elymniina and Zetherina fall outside the Satyrinae. The three gene regions used in a combined analysis prove to be very effective in resolving relationships of Satyrinae at the subtribal and tribal levels. Further sampling of the taxa closely related to Satyrinae, as well as more extensive sampling of genera within the tribes and subtribes for this group will be critical to test the monophyly of the subfamily and establish a stronger basis for future biogeographical and evolutionary studies.     

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.     

Revision of the Bicyclus sciathis species group (Lepidoptera: Nymphalidae) with descriptions of four new species and corrected distributional records

In this paper we present a thorough revision of the sciathis species group of the butterfly genus Bicyclus (Kirby). Type materials are discussed and in several cases lectotypes are assigned to specimens from original type series. Four new, and morphologically distinct, species are described (B. elishiae Brattström sp.n. , B. heathi Brattström sp.n. , B. sigiussidorum Brattström sp.n. and B. subtilisurae Brattström sp.n. ), along with a comprehensive molecular phylogeny that includes exemplar taxa of all currently recognized species. We also investigate the types of all previously synonymized taxa and in the process invalidate the name B. ewondo Libert. This was done after finding the previously missing holotype of B. makomensis (Strand), which clearly belongs to the same species and thereby gives the older name priority. The phylogeny showed that some distinctly different species were surprisingly closely related, suggesting a high rate of morphological evolution in parts of the sciathis group. The distributional records for the group are updated after investigating over 1700 specimens kept in a range of museum collections. Many species previously thought to be broadly sympatric were found to have much more restricted ranges, with the previous overestimations probably based on misidentified specimens. The higher level of allopatry now established will make identification of many morphologically similar species easier. The fact that species often have smaller ranges than previously known, meaning that the level of endemism for African butterflies is likely to be higher than current estimates, has important implications for conservation management. An identification key for males of all 13 currently recognized species in the species group is included. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:837A9D4C‐779A‐4497‐8176‐7151D409DFA5 .     

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.     

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。     

Simulation of movement and egglaying inDanaus plexippus (Lepidoptera: Nymphalidae)

A model simulating the movement and oviposition of monarch butterflies over a female's life time is presented. The model's predictions compare favourably with observed data and suggest that females who lay eggs in an optimal fashion should have low directionalities in areas with high host plant density (patches and single plants) and high directionalities in areas with low host plant density. The model also provides one means of combining individual animal processes and spatial heterogeneity into population dynamics.     

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.     

Relationships within the Melitaea phoebe species group (Lepidoptera: Nymphalidae): new insights from molecular and morphometric information

The genus Melitaea consists of about 80 species, divided into ten species groups, which are all restricted to the Palaearctic region. The Melitaea phoebe group was defined by Higgins based on morphological characters such as wing pattern and genital structures. According to his interpretation, the M. phoebe group included seven species: M. phoebe, M. sibina, M. scotosia, M. aetherie, M. collina, M. consulis and M. turkmanica. The taxonomy of the phoebe species group has been poorly resolved and recent results on the species composition within the group suggest the need for a re‐evaluation. In this study molecular sequences (5985 bp) including one mitochondrial (COI) and up to six nuclear (CAD, EF‐1α, GAPDH, MDH, RpS5 and wingless) gene regions from 38 specimens of the Melitaea phoebe species group sensu Higgins and some closely related taxa from the Palaearctic region were analysed. The possible evolution of the processus posterior of the male genitalia was also reconstructed based on a shape mapping technique. The analysis of the combined data shows a very clear pattern and almost all relationships are highly supported. Based on the combined Bayesian tree and the shape of the processus posterior of the male genitalia, four main groups are recognised: (i) collina group, (ii) arduinna group, (iii) aetherie group and (iv) phoebe group. The status of M. ornata, M. zagrosi and M. scotosia as species is confirmed, and the results also indicate that M. telona (s.s.) from Israel is a separate species.