Erebia serotina Descimon & de Lesse 1953 (Lepidoptera: Nymphalidae), a recurrent hybrid between two distantly related species

Erebia serotina was described in 1953 as a scarce, low-elevation endemic Pyrenean species flying late in the season. At least 34 individuals are known from various locations. However, the absence of females suggests a hybrid origin, and E. epiphron and either E. pronoe or E. manto have been proposed as possible parents. Electrophoretic analysis of five allozyme loci and sequencing of three mitochondrial DNA segments and one nuclear gene now demonstrate that E. serotina results from the cross between E. epiphron females and E. pronoe males. We have used our and previously published sequence data to generate a molecular phylogenetic tree of the genus Erebia which shows that these two species are only distantly related. The question of why they happen to hybridize on a seemingly routine basis is thus raised.     

中国勇红眼蝶一新亚种记述(鳞翅目:蛱蝶科:眼蝶亚科)

记述中国勇红眼蝶Erebia alcmena Grum—Grshimailo,1891的1新亚种——产自河南西部的勇红眼蝶李氏亚种Erebia alcmena leechuanlungi,ssp．nov．。模式标本保存于河南省科学院。     

Phylogenetic relationships among four members of Stizostedion (Percidae) determined by mitochondrial DNA and allozyme analyses

Phylogenetic relationships among four Stizostedion species were examined using mitochondrial DNA (mtDNA) and allozyme analyses. Twenty-six allozyme loci were scored, and mtDNA variation was examined using 24 restriction endonucleases, yielding 48–57 restriction sites among the species. Genetic distance analyses show that the two North American species ( S. canadense and S. vitreum ) cluster in one group, while the two European species ( S. hciopercu and S. vogense ) form a second group. Nei's genetic distance between these two groups was 0.7 ± 0.2 for allozymes, while the corresponding mtDNA sequence divergence was 14.8 ± 2.0%, suggesting that these two groups diverged approximately 10 million years ago. Thus, these data are consistent with the hypothesis that Stizostedion colonized North America during the Pliocene.     

Complete mitochondrial genomes of three skippers in the tribe Aeromachini (Lepidoptera: Hesperiidae: Hesperiinae) and their phylogenetic implications

The mitochondrial genome is now widely used in the study of phylogenetics and molecular evolution due to its maternal inheritance, fast evolutionary rate, and highly conserved gene content. To explore the phylogenetic relationships of the tribe Aeromachini within the subfamily Hesperiinae at the mitochondrial genomic level, we sequenced and annotated the complete mitogenomes of 3 skippers: Ampittia virgata, Halpe nephele, and Onryza maga (new mitogenomes for 2 genera) with a total length of 15,333 bp, 15,291 bp, and 15,381 bp, respectively. The mitogenomes all contain 13 protein‐coding genes (PCGs), 22 transfer RNAs (tRNAs), 2 ribosomal RNAs (rRNAs), and a noncoding A + T‐rich region and are consistent with other lepidopterans in gene order and type. In addition, we reconstructed the phylogenetic trees of Hesperiinae using maximum likelihood (ML) and Bayesian inference (BI) methods based on mitogenomic data. Results show that the tribe Aeromachini in this study robustly constitute a monophyletic group in the subfamily Hesperiinae, with the relationships Coeliadinae + (Euschemoninae + (Pyrginae + ((Eudaminae + Tagiadinae) + (Heteropterinae + ((Trapezitinae + Barcinae) + Hesperiinae))))). Moreover, our study supports the view that Apostictopterus fuliginosus and Barca bicolor should be placed out of the subfamily Hesperiinae.     

Intraspecific allozymatic differentiation reveals the glacial refugia and the postglacial expansions of European Erebia medusa (Lepidoptera: Nymphalidae)

Allozyme analysis of Erebia medusa over large regions of Europe revealed a significant population differentiation (F_ST: 0.149 ± 0.016). A UPGMA-analysis showed a division into four major lineages with mean inter-group genetic distances ranging from 0.051 (±0.010) to 0.117 (±0.024). An AMOVA revealed that rather more than two-thirds of the variance between samples was being between these lineages and less than one-third within lineages. An eastern group included the samples from the Czech Republic, Slovakia and north-eastern Hungary. This genetic lineage expressed significantly higher genetic diversity than the other three. A second lineage was formed by the samples from France and Germany. The two samples from western Hungary represent a third delimited lineage and the sample from northern Italy a fourth. We suppose that this genetic differentiation took place during the last ice-age in four disjunct refugia. The genetically more diverse eastern genetic lineage might have evolved in a relatively large refugium in south-eastern Europe. We assume that the other three lineages developed in relatively small relict areas around the Alps. It is likely for the western lineage that its ice-age distribution showed at least one disjunction in late Würm with the consequence of further genetic differentiation. Most probably, the eastern lineage colonized postglacial Central Europe using two alternative routes: one north and one south of the Carpathians. Up to now, neither similar glacial refugia, nor comparable secondary disjunctions in late Würm, are reported for any other animal or plant species.     

利用线粒体CO II基因序列对中国尾蛱蝶属系统分化的研究

利用线粒体COII基因序列分析法,研究了我国尾蛱蝶属5种蝴蝶的系统分化。结果表明,在尾蛱蝶属5个种的12个样品中,405 bp长的COII片段有11.4%的位点为多态性位点,大部分的碱基改变是转换。各物种内不同个体间的差异明显小于不同物种间的差异,种内个体间的差异一般为0.5%～1.5%,各物种间的差异绝大多数在4%以上。利用最大似然性法构建的尾蛱蝶属聚类关系图显示,尾蛱蝶属蝴蝶分为两大分支,一支包括大二尾蛱蝶、二尾蛱蝶和忘忧尾蛱蝶,另外一个分支包括窄斑凤尾蛱蝶和黑凤尾蛱蝶聚在一起。在大二尾蛱蝶、二尾蛱蝶和忘忧尾蛱蝶这一分支中,大二尾蛱蝶和忘忧尾蛱蝶的亲缘关系较近,而二尾蛱蝶较远。这些分子系统学的结果均与形态学的结果相一致,是对形态分类的有力支持。     

Some butterflies do not care much about topography: a single genetic lineage of Erebia euryale (Nymphalidae) along the northern Iberian mountains

The phylogeography of montane species often reveals strong genetic differentiation among mountain ranges. Both classic morphological and genetic studies have indicated distinctiveness of Pyrenean populations of the butterfly Erebia euryale. This hypothesis remained inconclusive until data from the westernmost populations of the distribution area (Cantabrian Mountains) were analysed. In the present study, we set out to describe the population structure of Erebia euryale in western Cantabria, where the species occurs in scattered localities. For this goal, we estimate the genetic diversity and differentiation found in 218 individuals from six Cantabrian (North Spain) localities genotyped by 17 allozyme loci. We also sequence 816 bp of the cytochrome oxidase subunit I mitochondrial gene in 49 individuals from Cantabrian localities and 41 specimens from five other European sites. Mitochondrial data support the recognition of four major genetic groups previously suggested for the European populations based on allozyme polymorphisms. Both mitochondrial and nuclear markers reveal genetic distinctiveness of a single Pyrenean–Cantabrian lineage of E. euryale. The lack of geographical structure and the star‐like topology displayed by the mitochondrial haplotypes indicate a pattern of demographic expansion in northern Iberia, probably related to Upper Pleistocene climatic oscillations. By contrast, within the Pyrenean–Cantabrian lineage, Cantabrian samples are genetically structured in nuclear datasets. In particular, San Isidro is significantly differentiated from the other five populations, which cluster into two groups. We recognize an evolutionary significant unit for Pyrenean–Cantabrian populations of Erebia euryale. Our results also illustrate that the evolutionary history of a species may be shaped by processes undetectable by using mtDNA alone.     

Molecular phylogeny of the Erebia tyndarus (Lepidoptera, Rhopalocera, Nymphalidae, Satyrinae) species group combining CoxII and ND5 mitochondrial genes: a case study of a recent radiation

The genus Erebia is one of the more controversial groups among the Holarctic Lepidoptera. In this genus, composed of around 100 mountain species, the Erebia tyndarus complex has certainly been a major source of debate for more than a century. According to the literature, between 2 and 11 species, and at least 45 subspecies or semi-species have been recognized. For the first time, we provide a complete molecular study, based on the ND5 and CoxII mitochondrial gene sequences, including all the recognized species and 60% of the subspecies of the entire group, with European, Caucasian, Siberian and North-American taxa. Our analyses led us to recognize 11 species even though there was some poor discrimination between Western Europe species. A complete study of the events of speciation is presented in the light of glacial and interglacial cycles during the Pleistocene.     

大卫绢蛱蝶线粒体基因组全序列测定和分析

目前有关蝶类线粒体基因组全序列及其分子进化的研究报道还不多见。本文利用long PCR和引物步移法得到大卫绢蛱蝶Calinaga davidis的线粒体基因组全序列, 同时就其基因组成和结构特点作了初步分析。结果显示： 其基因组全长为15 267 bp (GenBank登录号为HQ658143), 包括13个蛋白质编码基因(ATP6, ATP8, COI-III, ND1-6, ND4L, Cytb)、22个tRNA基因、2个rRNA基因(16S和12S)以及非编码的控制区。与其他鳞翅目昆虫相一致, 其基因组未出现基因重排现象。基因组共包含11个基因间隔区,总长度为130 bp, 间隔长度1~46 bp, 最大间隔在tRNA^Gln与ND2基因之间; 基因间共存在13处重叠, 总长度为66 bp, 重叠碱基数1~35 bp, 最长的重叠区位于COII与tRNA^Lys基因。lrRNA和srRNA基因长度分别为1 337 bp和773 bp; 除tRNA^Ser(AGN)缺少二氢尿嘧啶臂(DHU stem), 在相应的位置上只形成一个简单环外, 其余的tRNA基因都能形成典型的三叶草结构。13个蛋白编码基因总长度为11 247 bp, 共有3 737个密码子, 它们的碱基组成和密码子的使用具有明显的偏倚性; 除COI外(起始密码子TTG), 其余的12个蛋白质编码基因都以标准的ATN作为起始密码子; COI基因终止密码子为不完全T, ND4基因终止密码子为不完全TA, 其余基因都以TAA为终止密码子。A+T丰富区全长为389 bp, A+T含量高达92.0%, 其中存在2段类似微卫星的重复序列(TA)₆和(AAT)₄。本文的研究结果为探讨绢蛱蝶亚科在蛱蝶科中的系统学地位及其与其他亚科间的系统发生关系等问题提供了重要的分子生物学数据。     

Biogeography and pattern of gene flow among Barbus species (Teleostei: Cyprinidae) inhabiting the Italian Peninsula and neighbouring Adriatic drainages as revealed by allozyme and mitochondrial sequence data

Genetic relationships among 24 Italian and Adriatic populations of barbs (genus Barbus ) were assessed using electrophoretic analysis of allozymes and mitochondrial DNA sequences of the cytochrome b gene. Results obtained with both markers were concordant, but they were not congruent with the current morphology-based systematics and taxonomy. Populations assigned to the same nominal taxa ( B. caninus , B. petenyi and B. rebeli ), were very divergent in both allozymes and mtDNA, indicating that these populations deserve recognition as different units for conservation and management. On the other hand, the two fluvio-lacustrine taxa considered as distinct species (i.e. B. plebejus and B. tyberinus ) are genetically very close to each other, showing no clear differences at either allozymes or mtDNA. The population of B. caninus from Pellice River carried allozyme alleles and mtDNA specific for B. plebejus , indicating a genetic introgression towards the former species. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 83–99.     

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 and morphological divergence in the butterfly genus Lycaeides (Lepidoptera: Lycaenidae) in North America: evidence of recent speciation

Male genital morphology, allozyme allele frequencies and mtDNA sequence variation were surveyed in the butterfly species Lycaeides idas and L. melissa from across much of their range in North America. Despite clear differences in male genital morphology, wing colour patterns and habitat characteristics, genetic variation was not taxonomically or geographically structured and the species were not identifiable by either genetic data set. Genetic distances (Nei's D=0.002–0.078, calculated from allozyme data) between all populations of both species were within the range commonly observed for conspecific populations of other butterflies. The most frequent mtDNA haplotype was present in individuals of both species in populations from southern California to Wisconsin. We conclude that speciation has probably happened recently and the lack of genetic differentiation between the species is the product of either (1) recent or ongoing gene flow at neutral loci, and/or (2) an insufficiency of time for lineage sorting. The evolution of male genital morphology, wing colour patterns and ecological characteristics has proceeded more rapidly than allozyme or mtDNA evolution.     

Phylogeography of the Marsh Fritillary Euphydryas aurinia (Lepidoptera: Nymphalidae) in the UK

We sampled extant (and extinct) populations of Euphydryas aurinia to examine the phylogeography of the species in the UK. We were interested in whether the genetic structuring of populations reflects anything other than a single recent post-glacial colonization event. Four hundred base pairs of the mitochondrial Cytochrome b gene were sequenced from individuals from populations throughout the UK, as well as populations in France and Portugal and seven polymorphic allozyme loci resolved. The mean number of allozyme alleles per locus was 4 and isolation by distance was shown not to be a factor in the geographic structuring of genetic diversity either with or without the inclusion of the French data (Mantel statistic Z = 0.015 and 0.112 respectively, P >0.5). Cytochrome b nucleotide diversity (average number of nucleotide differences per site between two sequences) was low overall (0.003, n = 63) but mean cytochrome b gene diversity over all populations was 0.77. The presence of the Portuguese and French haplotype in Scottish populations indicates that the Iberian peninsula was likely to be one glacial refugium for E. aurinia populations. The pattern of mitochondrial DNA found in the UK could be interpreted in one of two ways: (1) two separate colonization events or (2) a single slow colonization event. Allele frequency distributions followed a similar geographical pattern as mitochondrial DNA haplotypes. An AMOVA assigned just 2.68% of allozyme genetic variation to the grouping tested, providing more support for the single colonization event theory.     

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。     

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.     

Characterization of microsatellite loci in Coenonympha hero (Lepidoptera: Nymphalidae)

Microsatellite markers are suitable tools for studying dispersal pattern among local populations. I report on the characterization of seven polymorphic microsatellite loci in the scarce heath butterfly (Coenonympha hero), from one unenriched and one enriched genomic library. Number of alleles ranged from two to 20 when 108 individuals from seven populations were screened. H_O ranged from 0.140 to 0.889. Primers were also tested on three other butterfly species. Amplification was successful for all loci in Erebia triaria, while only one gave products in Maculinea alcon and Maculinea rebeli. To my knowledge, these are the first microsatellite markers published in the Nymphalidae subfamily Satyrinae.     

The complete mitogenome of Parapoynx crisonalis (Walker, 1859) (Lepidoptera: Crambidae), with phylogenetic relationships amongst three acentropine larval forms

The larvae of Parapoynx crisonalis (Walker, 1859) is an important pest in China, causing significant damage on the aquatic crops. To improve the management of P. crisonalis and demonstrate the phylogenetic relationships amongst Pyraloidea, the complete mitochondrial genome (mitogenome) sequences of four aquatic pyralid moths, which belong to three acentropine larval forms – Parapoynx-type, Nymphula-type and Potamomusa-type, and twenty-three terricolous pyralids were compared. The complete mitogenome P. crisonalis, a typical parapoynx larval species, was 15,374 base pairs in length and included 13 protein-coding genes, 22 transfer RNA genes, 2 ribosomal RNA genes and an A + T-rich region. The phylogenetic analysis revealed that all the aquatic acentropine moths formed a monophyletic clade with strong phylogenetic support via maximum likelihood and Bayesian methods with a bootstrap value of 100% and a posterior probability of 1.00. The Potamomusa-type was more closely related to the Nymphula-type than to the Parapoynx-type.     

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. ).     

斐豹蛱蝶线粒体基因组全序列的测定和分析(英文)

该研究对斐豹蛱蝶(Argyreus hyperbius)(鳞翅目:蛱蝶科)线粒体基因组全序列进行了测定和初步分析。结果表明:斐豹蛱蝶线粒体基因全序列全长为15156bp,包含13个蛋白质编码基因、22个tRNA和2个rRNA基因以及1个非编码的A+T富集区,基因排列顺序与其它鳞翅目种类一致;线粒体全序列核苷酸组成和密码子使用显示出明显的A+T偏好(80.8%)和轻微的AT偏移(AT skew,?0.019)。基因组中共存在11个2～52bp不等的基因间隔区,总长96bp;以及14个1～8bp不等的基因重叠区,总长34bp。除COI以CGA作为起始密码子外,13个蛋白质编码基因中的其余12个基因是以ATN作为起始密码子。除COI和COII基因是以单独的一个T为终止密码子,其余11个蛋白质编码基因都是以TAA结尾的。除了缺少DHU臂的tRNASer(AGN),其余的tRNA基因都显示典型的三叶草结构。tRNA(AGN)和ND1之间的基因间隔区包含一个ATACTAA结构域,这个结构域在鳞翅目中是保守的。A+T富集区没有较大的多拷贝重复序列,但是包含一些微小重复结构:ATAGA结构域下游的20bp poly-T结构,ATTTA结构域后的(AT)9重复,以及位于tRNAMet上游的5bp poly-A结构等。这项研究所揭示的斐豹蛱蝶的线粒体基因组特征,不仅为认识蛱蝶科的遗传多样性贡献数据,而且对于该物种的保护生物学、群体遗传学、谱系地理及演化研究等具有重要意义。