Phylogenetic relationships of Hamadryas (Nymphalidae: Biblidinae) based on the combined analysis of morphological and molecular data

A new phylogenetic hypothesis for the Neotropical butterfly genus Hamadryas, based on the combination of a morphological matrix, one mitochondrial (COI) and four nuclear markers (CAD, RpS5, EF1a, and Wingless), is presented. Results from analyses of the molecular evidence are compared with a previously published morphological phylogeny. Molecular data and the analysis of the complete dataset support the monophyly of Hamadryas and most sister groups suggested by morphological data alone. The addition of DNA sequences to the morphological matrix helped define species groups for which no morphological synapomorphies were found. Partitioned Bremer support indicates that COI, CAD, and morphology were consistently in agreement with the combined evidence tree. In contrast, signal from the nuclear markers Rps5, EF1a, and Wingless showed indifference at most levels of the tree, and minor conflict at nodes solving the relationships between species groups. Though resolved, the combined evidence tree shows low resample values, particularly among species groups whose relationships were characterized by short internodes. A reassessment about the pattern of character change for sound production is presented and discussed.     

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.     

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.     

Biometrical evidence for adaptations of the salivary glands to pollen feeding in Heliconius butterflies (Lepidoptera: Nymphalidae)

The Neotropical genus Heliconius (Nymphalidae) is unique among butterflies for its pollen-feeding behaviour. With the application of saliva, they extract amino acids from pollen grains on the outside of the proboscis. We predicted that the salivary glands of pollen-feeding Heliconiinae would show adaptations to this derived feeding behaviour. A biometrical analysis of the salivary glands revealed that pollen-feeding butterflies of the genus Heliconius have disproportionately longer and more voluminous salivary glands than nonpollen-feeding Nymphalidae. The first two components in the principal component analysis explained approximately 95% of the total variance. The size-dependent factor score coefficients of body length and salivary gland parameters were predominately represented on axis 1. They significantly discriminated pollen-feeding from nonpollen-feeding heliconiines on that axis. Factor score coefficients for the volume of the secretory region of the salivary glands separated heliconiines from the outgroup species. The detailed biometrical analysis of salivary glands features thus provides strong evidence that the secretory regions of the salivary glands are larger in pollen-feeding butterflies. We concluded that pollen feeding is associated with a high production of salivary fluid.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 604–612.     

The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods

We have inferred the most comprehensive phylogenetic hypothesis to date of butterflies in the tribe Satyrini. In order to obtain a hypothesis of relationships, we used maximum parsimony and model‐based methods with 4435 bp of DNA sequences from mitochondrial and nuclear genes for 179 taxa (130 genera and eight out‐groups). We estimated dates of origin and diversification for major clades, and performed a biogeographic analysis using a dispersal–vicariance framework, in order to infer a scenario of the biogeographical history of the group. We found long‐branch taxa that affected the accuracy of all three methods. Moreover, different methods produced incongruent phylogenies. We found that Satyrini appeared around 42 Mya in either the Neotropical or the Eastern Palaearctic, Oriental, and/or Indo‐Australian regions, and underwent a quick radiation between 32 and 24 Mya, during which time most of its component subtribes originated. Several factors might have been important for the diversification of Satyrini: the ability to feed on grasses; early habitat shift into open, non‐forest habitats; and geographic bridges, which permitted dispersal over marine barriers, enabling the geographic expansions of ancestors to new environments that provided opportunities for geographic differentiation, and diversification.

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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.     

An integrative approach elucidates the systematics of Sea Hayward and Cybdelis Boisduval (Lepidoptera: Nymphalidae: Biblidinae)

The systematics of the enigmatic and rather uncommon species of Sea Hayward and Cybdelis Boisduval is assessed through morphological and molecular analyses. The aim of this paper is three‐fold: (i) to ascertain the phylogenetic position of Sea and Cybdelis in Biblidinae; (ii) to determine the evolutionary relationships among species of Sea and Cybdelis; and (iii) to review the taxonomy of Sea and Cybdelis. The result of the molecular analysis recovers Sea sister to Cybdelis, and this clade sister to Eubagina (sensu Lamas, 2004 ), rendering Epicaliina (sensu Lamas, 2004 ) paraphyletic. A new tribe, Cybdelini trib.n. is erected to include Sea and Cybdelis; the other genera formerly included in Epicaliina (sensu Lamas, 2004 ), Eunica Hübner and Sevenia Koçak are assigned to Eunicini trib.rest. , and the remaining genera of Biblidinae to other previously recognized tribes, similar to those implicitly proposed by Wahlberg et al. ( 2009 ). The morphological analysis recovers similar results and the following relationships among species of Cybdelis: C. phaesyla (Hübner) (C. boliviana Salvin (C. mnasylus Doubleday (C. petronita Burmeister stat.rest. + C. thrasylla C. Felder & R. Felder stat.rest. ))). Sea and Cybdelis are recognized as distinct genera; a new subspecies, S. sophronia jenkinsi Dias, Siewert & Freitas ssp.n. , distributed from Venezuela to Bolivia along the Andes eastern foothills, is described; and C. petronita and C. thrasylla are accepted as valid species. Lectotypes for C. boliviana, C. peruviana Staudinger, C. mnasylus, C. thrasylla, C. petronita and C. mnasylus var. meridionalis Hall and a neotype for Vanessa sophronia Godart are designated. Additionally, dichotomous keys and distribution maps to all taxa are provided; male genitalia and hypandria are illustrated for all species; generic characters of the head, labial palpi and legs are illustrated; immature stages of some species also are figured. The published work has been registered on ZooBank: http://zoobank.org/urn:lsid:zoobank.org:pub:D8EBB649‐5E9D‐4218‐B1F1‐7A69DA775FD0 .     

蛱蝶翅鳞片的超微结构观察

对我国东北地区典型常见蛱蝶科15属20种蝴蝶翅鳞片的超微结构进行了扫描电镜观察。结果显示：蛱蝶翅鳞片形态上可分为窄叶形、阔叶形和圆叶形3种,鳞片长65～135 μm,宽35～85 μm,间距48～112 μm。蛱蝶翅鳞片的超微结构可分为拱桥形、棋盘形和筛孔形3 种。拱桥形结构和棋盘形结构比较接近,二者与筛孔形结构差异较明显。在已观察的种类中,线蛱蝶属红线蛱蝶翅鳞片上的纵肋突起最小（200 nm×300 nm）,闪蛱蝶属柳紫闪蛱蝶翅鳞片上的纵肋突起最大（590 nm×560 nm）。鳞片具有相似的形状、结构和排列,尤其是同属蝴蝶翅鳞片超微结构的形状和尺寸差异较小,表明它们之间的亲缘关系接近。     

Phylogeny of Polygonia, Nymphalis and related butterflies (Lepidoptera: Nymphalidae): a total-evidence analysis

We investigated the phylogeny of butterflies in the tribe Nymphalini sensu Harvey 1991, comprising the genera Vanessa, Cynthia, Bassaris, Aglais, Inachis, Nymphalis, Polygonia, Kaniska, Antanartia, Hypanartia, Symbrenthia, Mynes and Araschnia . Evidence from the mitochondrial gene ndl, the nuclear gene 'wingless' and from morphology/ ecology/behaviour were used separately and combined to analyse relationships. Phylogenies based on the different types of data agreed in many aspects of basic topology. We show that an analysis of only wing pattern characters (based on Nijhout's homology system) results in a topology broadly similar to the one resulting from analysis of the complete matrix. We found support for a monophyletic Nymphalini, where Hypanartia may be the sister clade to all other genera. Mynes, Symbrenthia and Araschnia together seem to form another basal clade. Evidence presented gives only moderate support for a monophyletic Vanessa in the wide sense, including also Cynthia and Bassaris , but strong support for the monophyly of the largely holarctic clade Aglais + Inachis + Nymphalis + Polygonia + Kaniska + Roddia . Within the latter group there is strong support for a clade consisting of Aglais + Inachis and for a second clade which includes Nymphalis, Polygonia (and its sister clade, the monotypic Kaniska) as well as Roddia l-album (= Nymphalis vaualbum ). As a consequence of this topology, Aglais is recognized as a taxon separate from Nymphalis . We present a hypothesis of species relationships within the focal group of genera. We also analyse and discuss the implications of excluding or including ecological data in phylogenetic tree construction, when the tree is to be used for studies in phylogenetic ecology.     

Molecular evolution of a long wavelength-sensitive opsin in mimetic Heliconius butterflies (Lepidoptera: Nymphalidae)

This study examines the pattern of opsin nucleotide and amino acid substitution among mimetic species 'rings' of Heliconius butterflies that are characterized by divergent wing colour patterns. A long wavelength opsin gene, OPS1 , was sequenced from each of seven species of Heliconius and one species of Dryas (Lepidoptera: Nymphalidae). A parsimony analysis of OPS1 nucleotide and amino acid sequences resulted in a phylogeny that was consistent with that presented by Brower & Egan in 1997, which was based on mitochondrial cytochrome oxidase I and II as well as nuclear wingless genes. Nodes in the OPS1 phylogeny were well supported by bootstrap analysis and decay indices. An analysis of specific sites within the gene indicates that the accumulation of amino acid substitutions has occurred independently of the morphological diversification of Heliconius wing colour patterns. Amino acid substitutions were examined with respect to their location within the opsin protein and their possible interactions with the chromophore and the G-protein. Of the 15 amino acid substitutions identified among the eight species, one nonconservative replacement (A226Q) was identified in a position that may be involved in binding with the G-protein.     

The phylogenetics and biochemistry of host-plant specialization in Melitaeine butterflies (Lepidoptera: Nymphalidae)

Butterflies in the tribe Melitaeini (Lepidoptera: Nymphalidae) are known to utilize host plants belonging to 16 families, although most host-plant records are from four families. Of the 16 host-plant families, 12 produce secondary plant metabolites called iridoids. Earlier studies have shown that larvae of several melitaeine species use iridoids as feeding stimulants and sequester these compounds for larval defense. I investigate the evolutionary history of host-plant use in the tribe Melitaeini by testing a recent phylogenetic hypothesis of 65 species representing the four major species groups of the tribe. By simple character optimization of host-plant families and presence/absence of iridoids in the host plants, I find that plant chemistry is a more conservative trait than plant taxonomy. The ancestral host plant(s) of the entire tribe most likely contained iridoids and were likely to be in the plant family Plantaginaceae. A major host shift from plants containing iridoids to plants not containing iridoids has happened three times independently. The results show that the evolution of host-plant use in melitaeines has been (and still is) a dynamic process when considering plant taxonomy, but is relatively stable when considering host-plant chemistry.     

Mimicry in North American checkerspot butterflies: Euphydryas phaeton and Chlosyne harrisii (Nymphalidae)

ABSTRACT.

• 1 An examination of the phenetics, ecology and distribution of Chlosyne harrisii suggested that it was a potential mimic of the unpalatable Euphydryas phaeton.
• 2 The wing undersurfaces of the adults are very similar and the larvae are virtually identical. The geographic range of both butterflies is the northeastern United States and both are inhabitants of wet meadow habitats where they may fly together. The life histories of both species are closely synchronized.
• 3 A series of feeding experiments using blue-jays as predators showed first, that C.harrisii is palatable, and second, suggested that the birds could not distinguish between C.harrisii and E.phaeton on the basis of visual cues. Thus, Charrisii seems to be an effective Batesian mimic of E.phaeton.

     

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.     

The evolution of ecologically important characters in Heliconius butterflies (Lepidoptera: Nymphalidae): a cladistic review

The biology of Heliconius butterflies has provided a rich source of data to test theories of ecological genetics, coevolution and community ecology. Many putatively adaptive characters have been discussed with reference to a phylogenetic hypothesis based on a variety of morphological and life-history traits interpreted from an evolutionary taxonomic perspective. Here, alternate interpretations of characters on the traditional tree and a more recent mitochondrial DNA cladogram with a substantially different topology are compared and contrasted. It is shown that many characters ostensibly providing support for the traditional phylogenetic hypothesis are almost equally parsimoniously distributed and in some cases more parsimoniously distributed on the mtDNA tree than on the tree inferred from those characters. Discussion of alternate evolutionary scenarios based on the mDNA-based topology is presented for pupal mating, pollen feeding, foodplant coevolution, and other ecologically significant features.     

Phylogenetic relationships of butterflies of the tribe Acraeini (Lepidoptera, Nymphalidae, Heliconiinae) and the evolution of host plant use

The tribe Acraeini (Nymphalidae, Heliconiinae) is believed to comprise between one and seven genera, with the greatest diversity in Africa. The genera Abananote, Altinote, and Actinote (s. str.) are distributed in the Neotropics, while the genera Acraea, Bematistes, Miyana, and Pardopsis have a Palaeotropical distribution. The monotypic Pardopsis use herbaceous plants of the family Violaceae, Acraea and Bematistes feed selectively on plants with cyanoglycosides belonging to many plant families, but preferentially to Passifloraceae, and all Neotropical species with a known life cycle feed on Asteraceae only. Here, a molecular phylogeny is proposed for the butterflies of the tribe Acraeini based on sequences of COI, EF-1alpha and wgl. Both Maximum Parsimony and Bayesian analyses showed that the tribe is monophyletic, once the genus Pardopsis is excluded, since it appears to be related to Argynnini. The existing genus Acraea is a paraphyletic group with regard to the South American genera, and the species of Acraea belonging to the group of "Old World Actinote" is the sister group of the Neotropical genera. The monophyly of South American clade is strongly supported, suggesting a single colonization event of South America. The New World Actinote (s. str.) is monophyletic, and sister to Abananote+Altinote (polyphyletic). Based on the present results it was possible to propose a scenario for the evolution in host plant use within Acraeini, mainly concerning the use of Asteraceae by the South American genera.     

Dynamics of host plant use and species diversity in Polygonia butterflies (Nymphalidae)

The ability of insects to utilize different host plants has been suggested to be a dynamic and transient phase. During or after this phase, species can shift to novel host plants or respecialize on ancestral ones. Expanding the range of host plants might also be a factor leading to higher levels of net speciation rates. In this paper, we have studied the possible importance of host plant range for diversification in the genus Polygonia (Nymphalidae, Nymphalini). We have compared species richness between sistergroups in order to find out if there are any differences in number of species between clades including species that utilize only the ancestral host plants ('urticalean rosids') and their sisterclades with a broader (or in some cases potentially broader) host plant repertoire. Four comparisons could be made, and although these are not all phylogenetically or statistically independent, all showed clades including butterfly species using other or additional host plants than the urticalean rosids to be more species-rich than their sisterclade restricted to the ancestral host plants. These results are consistent with the theory that expansions in host plant range are involved in the process of diversification in butterflies and other phytophagous insects, in line with the general theory that plasticity may drive speciation.     

Adaptive radiations in butterflies: evolutionary history of the genus Erebia (Nymphalidae: Satyrinae)

We studied the speciose butterfly genus Erebia by reconstructing its phylogenetic relationships using parsimony and Bayesian approaches. We estimated times and rates of diversification for its lineages and employed a biogeographical analysis in order to reconstruct its evolutionary history. DNA sequence data from one mitochondrial gene and three nuclear genes were analyzed for a total of 74 species in Erebia. The estimated dates of origin and diversification for clades, in combination with a biogeographical analysis, suggest that the genus originated in Asian Russia and started its diversification process around 23 Myr. An important event was the dispersal of a lineage from Asia to Western Europe between 23 and 17 Myr, which allowed the radiation of most of species in the genus. The diversification pattern is consistent with a model of diversity limited by clade richness, which implies an early rapid diversification followed by deceleration due to a decrease in speciation. We argue that these characteristics of the evolutionary history of Erebia are consistent with a density‐dependent scenario, with species radiation limited by filling of niche space and reduced resources. We found that the Boeberia parmenio appears strongly supported in the genus Erebia and therefore we place Boeberia Prout, 1901 as a junior synonym of Erebia Dalman, 1816 ( syn. nov. ).     

On recognition of sound communication signals in bush crickets (Orthoptera,Tettigoniidae)

Two hypotheses have been proposed to explain the mechanisms of calling signal recognition in orthopterans: the filtration and resonance ones. To test these hypotheses, conspecific male calling songs and their models with modified temporal parameters were presented to females of bush crickets in ethological experiments. The models with a double pulse rate evoked positive phonotaxis of females while phase shift significantly complicated the recognition process. These data fit the resonance hypothesis.     

DNA barcoding of nymphalid butterflies (Nymphalidae: Lepidoptera) from Western Ghats of India

We have checked the utility of DNA barcoding for species identification of nymphalid butterflies from Western Ghats of India by using 650 bp sequence of mitochondrial gene cytochrome c oxidase subunit I. Distinct DNA barcoding gap (i.e. difference between intraspecies and interspecies nucleotide divergence), exists between species studied here. When our sequences were compared with the sequences of the conspecifics submitted from different geographic regions, nine cases of deep intraspecies nucleotide divergences were observed. In spite of this, NJ (Neighbour Joining) clustering analysis successfully discriminated all species. Observed cases of deep intraspecies nucleotide divergences certainly warrant further study.