Phylogenetic studies of ribosomal RNA variation in higher moths and butterflies (Lepidoptera: Ditrysia).

The selection of exemplars has been shown both theoretically and empirically to affect tree topology, but the importance of the number and nature of taxa used to represent higher taxonomic lineages in molecular studies is rarely stressed. In our rRNA study of higher moths and butterflies (Lepidoptera: Ditrysia), the selection of different exemplars and outgroups caused major tree rearrangements. We also examined the effectiveness with which conserved rRNA regions track the diversification of Lepidoptera. Homoplasy is as prevalent at the few variable sites of conserved regions (18E, 18J, 28F) as at the many variable sites of a more rapidly evolving region (28B). Finally, 28B sequence variation differs qualitatively among lepidopteran superfamilies of presumed comparable age, the Papilionoidea (true butterflies) and Noctuoidea (cutworm moths and relatives).     

Phylogeny of the superfamily Gelechioidea (Lepidoptera: Ditrysia): an exemplar approach

Phylogenetic relationships within the megadiverse lepidopteran superfamily Gelechioidea have been poorly understood and consequently the family level classification has been problematic. An analysis of phylogeny using 193 characters, including 241 informative character states, derived from larval, pupal and adult morphology and larval ecology, was performed to resolve the phylogeny of the Gelechioidea. 143 species representing the diversity of the putative Gelechioidea were included, supplemented with 13 species representing 11 other Ditrysian families. The monophyly of the Gelechioidea was supported, although only with homoplastic characters. The putative position of the Gelechioidea as the sister group of the Apoditrysia was not supported, since the Gelechioidea was nested within this clade. The Gelechioidea was divided into two main lineages: (1) the gelechiid lineage constituting Deoclonidae, Syringopainae, a re‐composed Coleophoridae (including Coelopoetinae and Batrachedrinae as paraphyletic with Stathmopodinae, and Coleophorinae nested within it), Momphidae, Pterolonchidae, Scythrididae, Cosmopterigidae, and Gelechiidae, and (2) the oecophorid lineage constituting the “autostichid” family assemblage (including taxa formerly assigned to Autostichinae, Holcopogoninae, Symmocinae, Glyphidoceridae and Lecithoceridae), Xyloryctidae s.l. (including a paraphyletic Xyloryctidae of authors, some oecophorids of authors, Deuterogoniinae and Blastobasinae), Oecophoridae s.s., Amphisbatidae s.s., Carcinidae, Stenomati[n/d]ae, Chimabachidae and Elachistidae (including Depressariinae s.s., Telechrysis, Ethmiinae, Hypertrophinae s.l., miscellaneous “amphisbatids”sensu authors, Aeolanthinae, Parametriotinae, Agonoxeninae and Elachistinae). Detritivory/fungivory may have evolved only twice within Gelechioidea, though the evolution of larval food substrate use frequently reverses. To avoid an unnecessary further proliferation of names, it is recommended that no further family group names are introduced within the Gelechioidea, unless based on a rigorous analysis of inter‐relationships.     

Evolutionary dynamics of rDNA clusters on chromosomes of moths and butterflies (Lepidoptera)

We examined chromosomal distribution of major ribosomal DNAs (rDNAs), clustered in the nucleolar organizer regions (NORs), in 18 species of moths and butterflies using fluorescence in situ hybridization with a codling moth (Cydia pomonella) 18S rDNA probe. Most species showed one or two rDNA clusters in their haploid karyotype but exceptions with 4–11 clusters also occurred. Our results in a compilation with previous data revealed dynamic evolution of rDNA distribution in Lepidoptera except Noctuoidea, which showed a highly uniform rDNA pattern. In karyotypes with one NOR, interstitial location of rDNA prevailed, whereas two-NOR karyotypes showed mostly terminally located rDNA clusters. A possible origin of the single interstitial NOR by fusion between two NOR-chromosomes with terminal rDNA clusters lacks support in available data. In some species, spreading of rDNA to new, mostly terminal chromosome regions was found. The multiplication of rDNA clusters without alteration of chromosome numbers rules out chromosome fissions as a major mechanism of rDNA expansion. Based on rDNA dynamics in Lepidoptera and considering the role of ordered nuclear architecture in karyotype evolution, we propose ectopic recombination, i.e., homologous recombination between repetitive sequences of non-homologous chromosomes, as a primary motive force in rDNA repatterning.     

Questions of molecular evolution of pheromone communication in caddisflies and lower moths (Insecta: Trichoptera,Lepidoptera)

Recent GS-MS and GC-EAD studies of pheromone production and perception in caddisflies and lower moths have shown that these insects use a rather limited selection of volatiles as attractants. Most of them are alcohols and ketones, although the diversity of chemicals produced by sternal glands of abdominal segment V is much wider, especially in the lower Trichoptera. Sternal pheromone glands produce only short-chain polymers in all Amphiesmenoptera. These glands are part of the ground-plan for the related orders, Trichoptera and Lepidoptera, occurring in both sexes and producing similar but not identical sets of components in males and females. The presence of pheromone volatiles is shown to be restricted to the gland segments (Fig. 1), although some other shortchain polymers do occur in the head of females of Molanna angustata (Molannidae). The pheromone blends of lower Trichoptera (Glossosomatidae, Rhyacophilidae, and Philopotamidae) are multi-component and resemble plant volatiles in composition. A hypothesis of the origin of pheromone communication is proposed postulating basic resemblance of early pheromones and plant volatiles in variety and chemical composition. These pheromones were detected by non-specialized receptors of the amphiesmenopteran ancestor and served as guides for insect aggregation on plants as well as on shores of reservoirs, marking the places suitable for a wide variety of species. The primary aggregation function of pheromones was changed in more advanced communication systems to the species-specific signaling with sex-related asymmetry of signals, although the aggregation significance persisted in some species. Pheromone communication has disappeared in some most advanced lineages (e.g., Leptoceridae) with parallel reduction of glands, secretion, and antennal receptors. The pheromone composition does not show gradual divergent evolution in related species; instead, abrupt transformation of pheromone blends with persistence of major components in remote families seems to be the typical case.     

Ground plan and evolution of pterothoracic musculature of moths and butterflies (lepidoptera)

Investigation of the meso- and metathoracic musculature in 62 species of Lepidoptera revealed 40 topographically different muscles, 37 of which presumably belong to the ground plan. The archaic (homoneuran) taxa demonstrate a higher similarity in the musculature of the meso- and metathorax than more advanced (heteroneuran) taxa do. The number of muscles increases with body size but does not depend on the forewing shape or the surface area ratio of the hind and fore wings. Reduction of many muscles occurred repeatedly thus causing similarity between phylogenetically distinct lineages; development of new muscles is rare and occurs mostly as a result of splitting of a muscle into morphologically different parts. Decrease in the number of muscles is accompanied by an increase in their size, more advanced groups generally having relatively larger muscles, especially in the mesothorax. The thoracic anatomy of Lepidoptera is strongly linked with biomechanics of flight and therefore has a relatively low value for phylogenetic analysis, although some characters may be used in reconstruction of phylogeny at the level of subfamilies to superfamilies.     

Phylogeny and life history evolution of Prodoxus yucca moths (Lepidoptera: Prodoxidae)

Abstract. Yucca moths (Lep., Prodoxidae) are well‐known for their obligate pollination mutualism with yuccas. In addition to the pollinators, yuccas also host many non‐pollinating yucca moths. Here the genus Prodoxus, the non‐pollinating sister group of the pollinators, is revised using morphological and molecular data, their phylogenetic relationships are analysed, and the evolution of host tissue specialization explored. Twenty‐two species are recognized, including nine new species: Prodoxus gypsicolor sp.n. , P. sonorensis sp.n. , P. carnerosanellus sp.n. , P. tamaulipellus sp.n. , P. weethumpi sp.n. , P. tehuacanensis sp.n. , P. californicus sp.n. , P. mapimiensis sp.n. and P. atascosanellus sp.n. Prodoxus y‐inversus Riley, P. coloradensis Riley and P. sordidus Riley are redescribed. The genus Agavenema is synonymized with Prodoxus. Phylogenetic analyses indicated that stalk‐feeding is basal within the group, that there are three separate origins of fruit‐feeding, and one origin of leaf‐mining from a stalk‐feeding ancestor. Although species with different feeding habits often coexist within hosts, the analyses suggest that ecological specialization and diversification within a host only may have occurred within one or possibly two hosts.     

A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution

Background

Tortricidae, one of the largest families of microlepidopterans, comprise about 10,000 described species worldwide, including important pests, biological control agents and experimental models. Understanding of tortricid phylogeny, the basis for a predictive classification, is currently provisional. We present the first detailed molecular estimate of relationships across the tribes and subfamilies of Tortricidae, assess its concordance with previous morphological evidence, and re-examine postulated evolutionary trends in host plant use and biogeography.

Methodology/Principal Findings

We sequenced up to five nuclear genes (6,633 bp) in each of 52 tortricids spanning all three subfamilies and 19 of the 22 tribes, plus up to 14 additional genes, for a total of 14,826 bp, in 29 of those taxa plus all 14 outgroup taxa. Maximum likelihood analyses yield trees that, within Tortricidae, differ little among data sets and character treatments and are nearly always strongly supported at all levels of divergence. Support for several nodes was greatly increased by the additional 14 genes sequenced in just 29 of 52 tortricids, with no evidence of phylogenetic artifacts from deliberately incomplete gene sampling. There is strong support for the monophyly of Tortricinae and of Olethreutinae, and for grouping of these to the exclusion of Chlidanotinae. Relationships among tribes are robustly resolved in Tortricinae and mostly so in Olethreutinae. Feeding habit (internal versus external) is strongly conserved on the phylogeny. Within Tortricinae, a clade characterized by eggs being deposited in large clusters, in contrast to singly or in small batches, has markedly elevated incidence of polyphagous species. The five earliest-branching tortricid lineages are all species-poor tribes with mainly southern/tropical distributions, consistent with a hypothesized Gondwanan origin for the family.

Conclusions/Significance

We present the first robustly supported phylogeny for Tortricidae, and a revised classification in which all of the sampled tribes are now monophyletic.     

Morphological character evolution in hepialid moths (Lepidoptera: Hepialidae) from New Zealand

Mapping morphological characters on a molecular-based phytogeny enabled examination of character evolution and an historical perspective into evolutionary processes, both of which are important aspects of systematic research and comparative biology. In this study, 63 morphological characters from hepialid moths in New Zealand were mapped on a phylogenetic tree reconstructed from mitochondrial DNA COI & II sequence data. Morphological characters hypothesized to be synapomorphies for the New Zealand 'Oxycanus' lineages and 'Oxycanus' lineage s.s. were confirmed to be homologous when mapped on the COI & II phytogeny. The direction of character state transformation was determined for five characters, with members of the Aenetus and Aoraia lineages exhibiting hypothesized ancestral states. Male genitalic characters were less homoplasious than other character partitions and covaried significandy with phytogeny.     

Linnaeus's butterflies (Lepidoptera: Papilionoidea and Hesperioidea)

The names of all the species of butterflies described by Linnaeus under "Papilio" are researched. Of the 305 names treated, 243 (c. 80%) are currently valid as specific (241) or subspecific (2), 29 are junior synonyms, 14 are invalid (one of these applying to a fake), and for 13 the identity is unknown or uncertain. Six species of moths misidentified by Linnaeus as butterflies are cited in the study, but details are not included. One hundred and fifty-two lectotypes have been designated, representing about 56% of the sum of the valid names and junior synonyms. Of these, 99 were selected from specimens in the Linnean Society of London, 52 from Queen Ludovica Ulrica's collection, Uppsala, and one lectotype is a Petiver specimen from the collection of Sir Hans Sloane. Linnaeus described at least five species, possibly eight, from the literature alone. All Linnaean material examined is documented, as are 'subsequent' specimens that are associated with Linnaean material. Synonymy and homonymy are discussed and presented, as are the identities of type localities. Care has been taken to achieve a practical balance between Linnaean and current species identities. Linnaean material studied included specimens from The Linnean Society of London, Museum Ludovicae Ulricae (Uppsala University), the Clerck and De Geer collections in the Naturhistoriska Riksmuseet, Stockholm, and the collection of James Petiver, now part of the collection of Sir Hans Sloane housed in The Natural History Museum, London.     

基于线粒体ND1和16S rRNA基因序列探讨中国蝴蝶12科的系统发育关系(鳞翅目: 双孔次亚目: 蝶类)

对中国12科共32种代表蝶类的ND1基因和16S rRNA 基因进行了序列测定（包括新测30种ND1基因和9种16S rRNA基因）和比较分析, 同时采用邻接法、最大似然法和贝叶斯法构建了12科蝶类的系统发育树, 探讨了其高级分类群的系统发育关系。序列分析的结果显示: 经比对处理后的两个基因总长度为869 bp, 其中保守位点373个, 可变位点496个, 简约信息位点375个; A+T的平均含量为80.2％, 明显高于C+G的平均含量19.8％。分子系统树表明: 蛱蝶科不是单系群; 珍蝶类、斑蝶类和喙蝶类位于蛱蝶科内; 粉蝶科和凤蝶科具有共同祖先。据此建议: 绢蝶科应归入凤蝶科; 蚬蝶科归入灰蝶科; 珍蝶类、斑蝶类和喙蝶类作为蛱蝶科中的亚科, 眼蝶类从蛱蝶科中分离出来独立成科。另外, 环蝶类的系统分类地位还有待于进一步研究。     

The evolution of chemical defences and mating systems in tiger moths (Lepidoptera: Arctiidae)

The origin and evolution of allelochemical sequestration in tiger moths (Arctiidae) is a complex interplay of larval and adult strategies and phylogenetic history. Using a phylogeny of Arctiidae, we examine the acquisition of secondary compounds from larval host plants and the use of secondary compounds and ultrasound in male courtship displays. We note that two sets of defensive signals (secondary chemicals and ultrasound) have been incorporated independently into arctiid courtship displays. Pyrrolizidine alkaloids are used in larval defence, and transformed into male courtship pheromones in several lineages. Phylogenetic inertia best explains the presence of adult collection and use of PAs in the absence of larval sequestration. Ultrasound, an adult defensive display directed at bats and other predators, has also been incorporated into arctiid mating displays. Sensory exploitation appears to underlie this co-option of defence signals for mating purposes.     

苹果园鳞翅目夜蛾科DNA条形码鉴定

为了检验DNA条形码在鳞翅目夜蛾科蛾类鉴定中的可行性,本文对采自北京昌平苹果园内的夜蛾科14种71头蛾类标本分别提取了DNA,并扩增了线粒体cox1及核基因28S,利用系统发育树、遗传距离、阈值等方法进行了鉴定和比较分析。同时,检验了目前BOLD系统的鉴定成功率。实验表明,基于cox1基因和BOLD系统的鉴定成功率达到了100%,而基于28S则很低,为64.8%。用不同方法构建的系统发育树,鉴定结果均相同。93%的种内遗传距离小于1%,94%的种间遗传距离为大于3%,种内种间的遗传距离形成明显的3%阈值现象。     

Comparative study of courtship in twelve phycitine moths (Lepidoptera: Pyralidae)

The courtship behavior of 12 phycitine moths (Lepidoptera: Pyralidae) was studied using frame-by-frame analysis of video recordings. Behavioral transitions during courtship were quantified for selected species and kinematic diagrams of courtship sequences were constructed. Interspecific similarities in courtship behaviors were measured by calculating Euclidean distances between species based on 12 courtship characters and by clustering species according to UPGMA (unweighted pair-group method using arithmetic averages). The resulting phenogram revealed two major behavioral patterns in courtship: (1) interactive and (2) simple. The former was characterized by a complex sequence in which, typically, a male approached a pheromoneemitting female, engaged in a head- to- head posture with the female, and then brought his abdomen over his head and struck the female on the head and thorax. This action brought male abdominal scent structures into close proximity with the female antennae. The male then attempted copulation from the head- to- head position by a dorsolateral thrust of the abdomen toward the female genitalia. Males of these species possessed scent structures located either on the eighth abdominal segment, or in a costal fold of the forewing, or both. Courtship in the second group was much more prosaic. After locating the female by response to her sex pheromone, the male simply attempted copulation by lateral abdominal thrusts under the female wing, without behavioral embellishments. Males of species exhibiting simple courtship had either no scent structures or structures that appeared vestigial. The grouping of species based on courtship characters was poorly correlated with taxonomic relationships, suggesting that the selective pressures governing the evolution and maintenance of courtship and male pheromones were distinct from those involved in the evolution of other morphological characters. While we argue that the primary force molding the evolution of courtship was an adaptive response to interspecific mating mistakes, we do not believe that isolation is brought about by the sequence of courtship behaviors themselves, due to the striking similarity in the sequence across several diverse species. Rather, these behaviors act to deliver more efficiently the male pheromonal message, which mayhave evolved for reproductive isolation.     

Data partitioning in Bayesian analysis: molecular phylogenetics of metalmark moths (Lepidoptera: Choreutidae)

In this study a multilocus phylogenetic analysis of metalmark moths (Lepidoptera: Choreutidae) focused on resolving the higher‐level phylogeny of this group is presented. Through the analysis of this dataset, I explore different data‐partitioning strategies in Bayesian phylogenetic inference, and find that a partitioning strategy can have a large influence on the results of phylogenetic analysis. Depending on how the data are partitioned, there can be significant differences in branch support. I also test for the existence of the Bayesian star tree paradox, and its importance in this dataset, and find that it appears to inflate support for the clade including Rhobonda gaurisana, Hemerophila houttuinialis, H. diva and H. felis, but plays no role in other cases where the differences between maximum‐likelihood bootstraps and Bayesian posterior probabilities are large. The results of all the phylogenetic analyses strongly suggest that including Millieriinae in Choreutidae renders the family polyphyletic. The monophyly of the other two subfamilies, Brenthiinae and Choreutinae, as well as their sister‐group relationship, is strongly supported. Similarly, the monophyly of all the genera examined except Hemerophila is also well supported. To bring the classification of Choreutidae in line with our current understanding of the phylogenetic relationships in the family, I propose to exclude Millieriinae from Choreutidae, elevate it to Millieriidae Heppner, and place it as incertae sedis within Ditrysia.     

Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae)

Background

The moth family Geometridae (inchworms or loopers), with approximately 23 000 described species, is the second most diverse family of the Lepidoptera. Apart from a few recent attempts based on morphology and molecular studies, the phylogeny of these moths has remained largely uninvestigated.

Methodology/Principal Findings

We performed a rigorous and extensive molecular analysis of eight genes to examine the geometrid affinities in a global context, including a search for its potential sister-taxa. Our maximum likelihood analyses included 164 taxa distributed worldwide, of which 150 belong to the Geometridae. The selected taxa represent all previously recognized subfamilies and nearly 90% of recognized tribes, and originate from all over world. We found the Geometridae to be monophyletic with the Sematuridae+Epicopeiidae clade potentially being its sister-taxon. We found all previously recognized subfamilies to be monophyletic, with a few taxa misplaced, except the Oenochrominae+Desmobathrinae complex that is a polyphyletic assemblage of taxa and the Orthostixinae, which was positioned within the Ennominae. The Sterrhinae and Larentiinae were found to be sister to the remaining taxa, followed by Archiearinae, the polyphyletic assemblage of Oenochrominae+Desmobathrinae moths, Geometrinae and Ennominae.

Conclusions/Significance

Our study provides the first comprehensive phylogeny of the Geometridae in a global context. Our results generally agree with the other, more restricted studies, suggesting that the general phylogenetic patterns of the Geometridae are now well-established. Generally the subfamilies, many tribes, and assemblages of tribes were well supported but their interrelationships were often weakly supported by our data. The Eumeleini were particularly difficult to place in the current system, and several tribes were found to be para- or polyphyletic.     

Systematics and evolution of the cutworm moths (Lepidoptera: Noctuidae): evidence from two protein-coding nuclear genes

Abstract. A broad molecular systematic survey of Noctuidae was undertaken to test recent hypotheses on the problematic definitions and relationships of the subfamilies, with special emphasis on the ‘trifines.’ An initial hypothesis of noctuid classification to the subtribal level was synthesized from recent reviews, and then sampled as broadly as possible. Concatenated sequences for the nuclear genes elongation factor‐1α (EF‐1α; 1200 bp) and dopa decarboxylase (DDC; 700–1100 bp) were analysed for a total of 146 exemplar species, twice that of a previous study. Trees were estimated using likelihood, distance, and both equally weighted and ‘six‐parameter’ parsimony. Of the 144 possible nodes, bootstrap support (BP) was ≥ 50% for ～80, and ≥ 80% for ～60. There was very strong support (BP ≥ 90%) for an ‘L.A.Q.’ clade encompassing nearly all quadrifine noctuids plus Arctiidae and Lymantriidae, decisively rendering Noctuidae paraphyletic. We present a new classification for Noctuoidea in which Noctuidae sensu stricto is restricted to trifines; most quadrifine subfamilies are raised to full families. Within the ‘L.A.Q.’ clade, Aganainae and Herminiinae were strongly grouped, but other relationships were weakly supported, probably due to limited taxon sampling. Nolidae and Euteliinae + Stictopterinae are generally grouped with the ‘L.A.Q.’ clade, but with weak support. All analyses favour the broadest definitions proposed for the trifine clade (our Noctuidae sensu stricto) although support is not strong, except that the exemplar of Eustrotiinae: Eublemmini is placed securely in the ‘L.A.Q.’ clade. Numerous recent proposals for dismantling and recombining the ‘Hampsonian’ traditional trifine subfamilies are strongly supported, most notably a broadly defined ‘true cutworm’ clade (Noctuinae s.l.), encompassing the greater part of the traditional subfamilies Amphipyrinae, Cuculliinae, Hadeninae and Noctuinae s.s. (BP ≥ 95%). Within this clade there is strong support for Apameini s.s.+ Xylenini s.l. and for Noctuinae s.s. and divisions thereof, but little support for monophyly or subdivision of Hadeninae. Noctuinae s.l. invariably are allied with Heliothinae, scattered remnants of the traditional Amphipyrinae, and several smaller groups in a broader ‘pest clade’, albeit with weak support. Relationships among the remaining ‘lower’ trifines are not strongly resolved. Mapping of a preliminary synopsis of species diversities, host use patterns and latitudinal distributions on the phylogeny suggests that the diversification of trifines may have been promoted, to a degree unique among Macrolepidoptera, by the Tertiary expansion of seasonal, open habitats and their associated herbaceous floras.     

The fossil record and taphonomy of butterflies and moths (Insecta,Lepidoptera): implications for evolutionary diversity and divergence-time estimates

Background

It is conventionally accepted that the lepidopteran fossil record is significantly incomplete when compared to the fossil records of other, very diverse, extant insect orders. Such an assumption, however, has been based on cumulative diversity data rather than using alternative statistical approaches from actual specimen counts.

Results

We reviewed documented specimens of the lepidopteran fossil record, currently consisting of 4,593 known specimens that are comprised of 4,262 body fossils and 331 trace fossils. The temporal distribution of the lepidopteran fossil record shows significant bias towards the late Paleocene to middle Eocene time interval. Lepidopteran fossils also record major shifts in preservational style and number of represented localities at the Mesozoic stage and Cenozoic epoch level of temporal resolution. Only 985 of the total known fossil specimens (21.4%) were assigned to 23 of the 40 extant lepidopteran superfamilies. Absolute numbers and proportions of preservation types for identified fossils varied significantly across superfamilies. The secular increase of lepidopteran family-level diversity through geologic time significantly deviates from the general pattern of other hyperdiverse, ordinal-level lineages.

Conclusion

Our statistical analyses of the lepidopteran fossil record show extreme biases in preservation type, age, and taxonomic composition. We highlight the scarcity of identified lepidopteran fossils and provide a correspondence between the latest lepidopteran divergence-time estimates and relevant fossil occurrences at the superfamily level. These findings provide caution in interpreting the lepidopteran fossil record through the modeling of evolutionary diversification and in determination of divergence time estimates.

Electronic supplementary material

The online version of this article (doi:10.1186/s12862-015-0290-8) contains supplementary material, which is available to authorized users.     

A molecular phylogenetic analysis of the vampire moths and their fruit-piercing relatives (Lepidoptera: Erebidae: Calpinae)

Within butterflies and moths, adult hematophagy is limited to species within the vampire moth genus Calyptra. These moths are placed within the subfamily Calpinae, whose other members are known to exhibit a broad range of feeding behaviors including those that can be considered 'piercers' of fruits or other hosts and 'tear feeders'. Here, we reconstruct a phylogenetic hypothesis of Calpinae using molecular data to test whether hematophagy in Calyptra arose from plant or animal-related behaviors. We use a Bayesian method of ancestral state reconstruction to determine the most likely feeding behaviors for the subtribes and genera within this lineage.     

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.     

Hairstreak butterflies (Lepidoptera,Lycaenidae) and evolution of their male secondary sexual organs

Hairstreak butterflies in the Atlides Section of the Eumaeini are biologically notable for a diverse array of male secondary sexual organs. A “species recognition” hypothesis postulates that females use these organs to choose between conspecific and non-conspecific males, thereby promoting reproductive isolation. Alternately, a “sexual selection” hypothesis posits that females use these organs to choose among conspecific males. These hypotheses need not be mutually exclusive but make different predictions about the evolutionary gain and loss of male secondary sexual organs. We analysed most of the Atlides Section (Theclinae, Eumaeini) phylogenetically. Sister lineages were sympatric at 22 of 37 nodes. Nine evolutionary gains occurred in lineages that were sympatric with their phylogenetic sister, and one occurred in a lineage that was allopatric/parapatric with its sister. By contrast, seven of ten evolutionary losses occurred in lineages that were allopatric/parapatric with their sisters. These results are significantly different from those predicted by a sexual selection hypothesis. We conclude that male secondary sexual organs in the Atlides Section function primarily for species recognition and thereby promote sympatric diversification.