Patch dynamics ofGlaucopsyche lygdamus (Lycaenidae): correlations between butterfly density and host species diversity

Glaucopsyche lygdamus egg densities were surveyed over a 2000-m section of Gold Creek and at 30 different isolated patches in the Gold Basin drainage in Colorado. Host plant numbers and diversity were quantified, as well as other variables potentially influencing butterfly population size, such as patch size and isolation. Egg densities correlated significantly only with measures of host species diversity. Patches consisting of a single host species, no matter how large, did not support high butterfly densities, but patches of multiple, equitably distributed host species did. The most likely explantation, in light of oviposition preference and larval performance data accumulated for this butterfly species, is that host species diversity is necessary for the persistence ofG. lygdamus populations, because alternative host species buffer population losses during poor or unusual years. The dependence of both ovipositing butterflies and developing larvae on the ephemeral, young, host plant flowers make the butterfly especially vulnerable to year-to-year variation in host plant availability and quality.     

Museum genomics reveals the Xerces blue butterfly (Glaucopsyche xerces) was a distinct species driven to extinction

The last Xerces blue butterfly was seen in the early 1940s, and its extinction is credited to human urban development. This butterfly has become a North American icon for insect conservation, but some have questioned whether it was truly a distinct species, or simply an isolated population of another living species. To address this question, we leveraged next-generation sequencing using a 93-year-old museum specimen. We applied a genome skimming strategy that aimed for the organellar genome and high-copy fractions of the nuclear genome by a shallow sequencing approach. From these data, we were able to recover over 200 million nucleotides, which assembled into several phylogenetically informative markers and the near-complete mitochondrial genome. From our phylogenetic analyses and haplotype network analysis we conclude that the Xerces blue butterfly was a distinct species driven to extinction.     

The setae of parasitic Liphyra brassolis butterfly larvae form a flexible armour for resisting attack by their ant hosts (Lycaenidae: Lepidoptera)

Caterpillars of the lycaenid butterfly, Liphyra brassolis, live inside the nests of arboreal weaver ants, Oecophylla smaragdina, and eat their brood. Observations of mature larvae suggest that they are impervious to relentless ant molestation, yet they lack sclerotized cuticular plates. We document a novel form of integumental defence that imparts protection from ant attack whilst maintaining the flexibility necessary to walk with a hydraulic skeleton. Analysis of the trunk integument and cuticular structures of early and late instars of L. brassolis using light microscopy, scanning electron microscopy, and histology revealed three new setae types (disc, clavate, and lanceolate), as well as three new cuticular structures (pored sockets, cuticular pores, and cuticular domes). The unique cuticle is covered with lanceolate setae, which act as endocuticular struts, and overlapping scale‐like sockets, which form a hard, flexible integument. The imperfect armour of the early‐instar larvae suggests that abundant, putatively secretory pores are likely to be homologous to pore cupola organs (PCOs) found in other lycaenid larvae and thus may exude semiochemicals to allay ant aggression. The importance of these pores presumably wanes as structural (setal) cuticular defenses are reinforced in later instars, when adult ants have been observed attacking caterpillars to no avail. The caterpillar's antennae are unusual and seem to be involved in manipulating ant larvae into the caterpillar's mouth. Behavioural observations indicate that the dexterity of these structures is associated with eating ants (myrmecophagy).     

New Australian butterfly genus Jameela gen. nov. (Lepidoptera: Lycaenidae: Polyommatinae: Polyommatini) revealed by morphological, ecological and molecular data

This paper proposes to split the Australian Polyommatini genus Erysichton Fruhstorfer, 1916 into two genera on the basis of molecular, morphological and early stage behavioral attributes. The utility of reflective eye pattern as a taxonomic character is discussed and a neotype of Lycaena lineata Murray, 1874 is designated.     

A common terminology for the external anatomy of centipedes (Chilopoda)

A common terminology for the external morphological characters of centipedes (Chilopoda) is proposed. Terms are selected from the alternatives used in the English literature, preferring those most frequently used or those that have been introduced explicitly. A total of 330 terms are defined and illustrated, and another ca. 500 alternatives are listed.     

The differences of butterfly (Lepidoptera, Papilionoidea) communities in habitats with various degrees of disturbance and altitudes in tropical forests of Vietnam

A survey was conducted on the species composition, richness and abundance of Papilionoidea (excluding Lycaenidae) butterfly fauna in habitats with various degrees of disturbance and altitudes in tropical forests at Tam Dao National Park, northern Vietnam in 2001. The transect method was used to collect data in the survey. Six transects representing different habitat types at two sites, one site located at a low elevation of 200–250 m a.s.l., and the other located at a high elevation of 950–1000 m a.s.l., were chosen: three transects for each site, with a length of 500 m for each transect. A total of 3594 individuals of 127 species in 240 sets of data were recorded from various habitats. The differences in butterfly composition, species richness, abundance and diversity in different habitat types and altitudes were analyzed. The results showed significant differences of butterfly diversity among the different habitat types and between the low and high altitude sites. The butterfly diversity, species richness and species abundance in the low elevation habitats were higher than in the high elevation habitats. The highest diversity of butterflies occurred in the mixed habitats of agriculture, scrub and clearing lands of high disturbance. However, butterflies most important for conservation are associated with undisturbed or moderately disturbed forests only.     

基于线粒体 COI 和 Cytb 基因的中国灰蝶三亚科主要类群间的系统发育关系分析

【目的】针对中国灰蝶科中亲缘关系较近的3个主要亚科［灰蝶亚科(Lycaeninae)、线灰蝶亚科(Theclinae)以及眼灰蝶亚科(Polyommatina)］,基于线粒体基因序列数据研究它们主要类群间的系统发育关系。【方法】对3亚科共53种灰蝶的线粒体 COI 和 Cytb 基因进行序列测定和序列变异分析,同时,基于最大似然法(maximum likelihood, ML)和贝叶斯法(bayesian inference, BI)等建树方法重建53种灰蝶的系统发育树。【结果】串联的2个基因共1 431 bp,其中保守位点855个,可变位点576个,简约信息位点488个;A+T的平均含量为74.5%,明显高于G+C的平均含量(25.5%)。系统树显示,灰蝶亚科以及眼灰蝶亚科均是单系发生,线灰蝶亚科则为并系群。全部灰蝶物种共分为三大支系：灰蝶亚科为第1支系;眼灰蝶亚科与线灰蝶亚科中的旖灰蝶族(Hypolycaenini)、富丽灰蝶族(Aphnaeini)分别构成单系群并互为姊妹群,它们共同构成第2支系;线灰蝶亚科中的美灰蝶族(Eumaeini)、玳灰蝶族(Deudorigini)、娆灰蝶族(Arhopalini)和线灰蝶族(Theclini)构成第3支系,其亲缘关系为：（（（线灰蝶族+娆灰蝶族）+玳灰蝶族）+美灰蝶族）。【结论】本研究涉及的3个灰蝶亚科中,灰蝶亚科是一个独立的支系,眼灰蝶亚科与线灰蝶亚科之间有较近的亲缘关系,但它们内部主要类群间的系统发育关系还需要进一步的研究。     

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.     

Functional analysis of the myrmecophilous relationships between ants (Hymenoptera: Formicidae) and lycaenids (Lepidoptera: Lycaenidae)

Summary Mature larvae of the myrmecophilous lycaenid Polyommatus coridon produce an average of 30.9 droplets of a honeydew-like secretion per hour. They occur in population densities of about 20/m². The volume of secretion over the whole larval period is estimated to be 22–44 l with an energy content of 55–110 J. Thus, P. coridon larvae produce carbohydrate secretions with an energy equivalent of 1.1–2.2 kJ/m². Using data from the literature on ant metabolism, it is shown that these carbohydrate secretions may contribute significantly to the nutrition of attending ants. The myrmecophilous relationship between the larvae of P. coridon and ants should therefore be regarded as a mutualistic symbiosis.     

Incongruence between morphological and molecular markers in the butterfly genus Zizina (Lepidoptera: Lycaenidae) in New Zealand

The butterfly genus Zizina in New Zealand has a complex taxonomic history due to the presence of morphological intermediates between the two species, the endemic Z. oxleyi and the introduced Z. labradus, in a putative hybrid zone on the east coast of the South Island of New Zealand. This makes species identification in the field problematic, particularly as the presence of hybrids has not been confirmed. We address this uncertainty through morphological and molecular analyses. Specimens were collected from a range of locations in New Zealand, as well as from Australia, and measurements were made of male genitalia and ventral wing coloration. Two mitochondrial genes (COI, ND5) and three nuclear gene fragments (28S, ITS2 and wingless) were also sequenced for a selection of individuals, and the presence of Wolbachia species in genomic DNA was tested. The two species were separable in morphological space, although there was some overlap, and the contact zone appeared to be around Kaikoura on the east coast of the South Island. Furthermore, specimens from the putative hybrid zone could be classified as Z. oxleyi using morphological characters individually, but not when these were used in a principal component analysis. Molecular analysis showed that there was a mean sequence divergence of 2.0% between two clades for COI, and 4.1% for ND5, but suggested that the contact zone between them was in the north‐west of the South Island. However, there was only a single clade for each of the three nuclear markers. It is thought that this incongruence between morphological and molecular markers is indicative of hybridization which is more extensive than previously thought. However, the possibility that recent speciation has occurred or is occurring is not ruled out.     

Comparative study of the morphology of stridulatory organs of the Iberian lycaenid butterfly pupae (Lepidoptera)

We compared the pupal stridulatory organs of 35 species and one subspecies of Iberian Lycaenidae using scanning electron microscopy. The studied species belong to the tribes Theclini, Eumaeini, Lycaenini, and Polyommatini. Nine species do not show stridulatory organs on the pupae but all other species possess them. Stridulatory organs are formed by a stridulatory plate (pars stridens) placed on the fifth abdominal segment and a file (plectron) in the sixth abdominal segment. The plate has tubercles in the Theclini and Lycaenini, tubercles, ridges, or undulations in the Eumaenini, and tubercles, teeth, or unspecialized structures in the Polyommatini. Morphological differences can be found in the files of the different tribes, regarding the number of teeth, their form and size. Cuticular formations of the organs were studied on a surface of 2,500 µm² and the average of ridges, tubercles, and teeth was measured searching for relevant taxonomic information. Stridulatory organs were thought to be an adaptation to myrmecophily but we show that they are present both in myrmecophilous and nonmyrmecophilous species; therefore, we suggest that this trait probably did not evolve in relation with myrmecophily, but may be used to enhance relationships with ants. J. Morphol. 275:414–430, 2014. © 2013 Wiley Periodicals, Inc.     

Specializations and polyphagy of Plebejus argus (Lepidoptera: Lycaenidae) in North Wales

Abstract. 1. The polyphagous butterfly, Plebejus argus L., was found to have specialized requirements: eggs are laid along vegetation/bare ground margins, larvae specialize on tender meristematic and reproductive growth, larvae and pupae are tended by Lasius ants, and warm microclimates are favoured.
2. These requirements are met in early successional habitats within several bio topes. These habitats are locally distributed, as is P. argus.
3. Populations on limestone, heathland and mossland biotopes did not differ greatly in ecologies. Given a choice of foodplants in captivity, larvae from heathland and limestone did show slight differences in preference, selecting some of the hostplants they normally encounter in the wild. Regardless of biotope, all these larvae were plant taxa generalists (on three plant families), but specialists on plant parts.
4. The decline of P. argus in Britain is attributed to loss of biotopes and to declining traditional management of them. Fewer habitats are available, and the distances between them are greater than formerly. As some of these habitats are short lived, and P. argus rarely flies far, the butterfly's decline is a compound effect of reduced areas of habitat and inability to colonize them.     

Muscular anatomy of a whipspider, Phrynus longipes (Pocock) (Arachnida: Amblypygi), and its evolutionary significance

Skeletal muscles in the whipspider Phrynus longipes are surveyed and compared with those of other chelicerates to clarify the evolutionary morphology and phylogenetic relationships of the arachnids. Representatives of 115 muscle groups are described and illustrated, and their possible functions are proposed. Principal results of this analysis include new functional models for the operation of the pharyngeal and sternocoxal mechanisms in Amblypygi and a greatly expanded list of apparently unique synapomorphies supporting the monophyly of Pedipalpi (= Amblypygi, Schizomida, Thelyphonida).     

World travellers: phylogeny and biogeography of the butterfly genus Leptotes (Lepidoptera: Lycaenidae)

Leptotina butterflies (Lycaenidae, Polyommatiinae) are found mostly in tropical and subtropical areas around the globe, marginally penetrating into temperate regions. Here, we investigated phylogenetic and biogeographical relationships of most representatives of the subtribe, using both likelihood and Bayesian approaches. We also estimated the timing of their diversification. And lastly, we studied phylogeographic patterns of the most widespread species, Leptotes pirithous. DNA sequences from two mitochondrial (COI, COII) and two nuclear genes (wingless, Ef1α) were analysed for 13 species of the genus Leptotes Scudder and one species of the genus Cyclyrius Butler. Both genera together form a monophyletic clade, and Cyclyrius is rooted deep inside Leptotes. Therefore, we designate Cyclyrius to be a junior synonym of Leptotes. According to our study, the genus Leptotes originated between the late Eocene and early Oligocene (35–31 Ma). During the Miocene it dispersed to the rest of the southern hemisphere, with further speciation events within the Indo‐Australian region, and separate radiations in the Americas and the Afrotropics. Leptotes webbianus from the Canary Islands turned out to be sister to the American clade from which it split c. 12 Ma. Leptotes pirithous originated in Madagascar c. 4 Ma and invaded the whole of Africa and southern Europe, including numerous surrounding islands. Populations of L. pirithous from Mauritius and Madagascar turned out to represent a distinct species (Leptotes durrelli sp.n. ) and the same applies to the Australasian populations of Leptotes plinius (Leptotes lybas stat. rev. ). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:20308930‐988B‐4327‐A35F‐CC983D46263B .     

The larval head anatomy of Rhyacophila (Rhyacophilidae) with discussion on mouthpart homology and the groundplan of Trichoptera

The external and internal features of the larval head of Rhyacophila fasciata (Trichoptera: Rhyacophilidae) were described in detail. Anatomical examinations were carried out using a multimethod approach including histology, scanning electron microscopy, confocal laser‐scanning microscopy, microcomputed tomography, and computer‐based three‐dimensional reconstructions. Additionally, the information on the larval head of Limnephilus flavicornis (Limnephilidae) and Hydropsyche angustipennis (Hydropsychidae) available in the literature were reinvestigated. These anatomical data were used to address major questions of homology and terminology, that is, the ventral closure of the head capsule, the sclerites, and appendages of labium and maxilla and their muscles. These topics were discussed by summarizing the main hypotheses present in the literature and a critical inclusion of new findings. Consequently, the inner lobe of the maxilla very likely represents the galea. The distal maxillary sclerite (palpifer) is an anatomical composite formation at least including dististipes and lacinia. Based on these homology hypotheses several potential groundplan features of the larval head of Trichoptera were reconstructed. The head of Rhyacophila shows several presumably plesiomorphic features as for instance the prognath orientation of the mouthparts, the well‐developed hypocranial bridge, the triangular submentum and eyes composed of seven stemmata. Derived features of Rhyacophila are the reduced antennae, the anterior directing of three stemmata and the shift of the tentorio‐stipital muscle to the mentum. J. Morphol. 276:1505–1524, 2015. © 2015 Wiley Periodicals, Inc.     

Early comers occupy popular territories in a lycaenid butterfly,Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae)

Territorial activities of a lycaenid butterfly, Chrysozephyrus smaragdinus, were observed throughout its adult season to investigate the territorial dynamics. The territories occupied earlier in the season were abandoned later than later‐occupied territories, indicating that the site preference of male C. smaragdinus was largely maintained during the adult season. Three indexes of territory quality based on animal preference, occupation duration (OD), initial occupation date (IOD) and the number of contests (NOC), were highly correlated. Therefore, these preference‐based indexes are consistent and reliable, although they are indirect estimates of benefits obtained from territories. Males arriving at territory sites early in the season were more likely to hold territories of better quality when NOC was used as the index of territory quality. The positive relationship between territory quality and male arrival date is congruent with the result of a previous study showing that early comers tend to hold a territory whereas late comers do not. However, male arrival date was not related to territory quality when OD or IOD was used as an index of territory quality. One possible reason for this apparent discrepancy is that the coefficient of variation of NOC is larger than those of OD and IOD, and therefore NOC could evaluate territory quality more quantitatively than the other two indexes. To date, studies of territoriality in butterflies have not considered the quality of territories. More studies considering the quality of territories will provide new insights into butterfly territoriality.     

Skeletomuscular anatomy of the harvestman Leiobunum aldrichi (Weed, 1893) (Arachnida: Opiliones: Palpatores) and its evolutionary significance

Skeletal muscles of the North American harvestman Leiobunum aldrichi are exhaustively surveyed and compared with other chelicerates to clarify the evolutionary morphology and phylogenetic relationships of arachnids. Representatives of 104 muscle groups are described and illustrated, and their possible functions are proposed. Comparisons of the feeding apparatus of L. aldrichi with that of other opilions, especially Sim (Cyphophthalmi) and Acromares (Laniatores), and two scorpion genera ( Centruroides, Pandinus ) indicate that the pharyngeal apparatus in L. aldrichi is derived and that its ability to accommodate large food particles is a secondary rather than primitive condition. Comparisons reveal several possible synapomorphies between Opiliones and Scorpiones suggesting that these orders may be sister groups. Apparently unique synapomorphies include an extrinsic cheliceral muscle that arises from the carapace and inserts on the second cheliceral article (deutomerite); an epistome divided into distal and proximal parts by a transverse sulcus; pharyngeal dilator muscles supported by a peripharyngeal skeleton formed by one dorsomedial and two ventrolateral epistomal processes, the latter also with muscular attachments to the endosternite; a specialized preoral chamber (stomodieca) derived from extensions (coxapophyses) of the coxae of the pedipalp and first two leg pairs; internal processes associated with the coxapophyses that serve, in part, as an attachment for muscles operating the coxa-trochanter joints, and lateral endosternal suspensor muscles that insert on the arthrodial membrane between the leg coxae. These are the first observations providing explicit support for an Opiliones-Scorpiones clade.     

Structure of the food web including the endangered lycaenid butterfly Shijimiaeoides divinus asonis (Lepidoptera: Lycaenidae)

The structure of the food web including the endangered lycaenid butterfly Shijimiaeoides divinus asonis (Matsumura) was analyzed to identify species contributing most to maintaining the equilibrium of the food web. Twenty‐seven species belonging to 17 families fed on Sophora flavescens Aiton, the host‐plant of S. divinus asonis: 15 species were leaf and stem feeders, seven (including S. divinus asonis) fed on flower buds, four were flower feeders and one fed on the seeds of So. flavescens. Of these 27 species, four were omnivores. The natural enemies of S. divinus asonis comprised six insect species, 11 spider species and one entomopathogenic fungus species, including six new predator records. The linkage density, total number of trophic links, connectance, average chain length and predator–prey ratio were 1.617, 97, 0.0548, 2.267 and 0.694, respectively. Exclusion of any of the 15 species with four or more trophic links reduced the connectance of the food web. These 15 species included facultative mutualistic attendant ants and predators of S. divinus asonis, herbivores to So. flavescens, an omnivore feeding on S. divinus asonis and So. flavescens, and prey insects. Therefore, future studies should monitor these 15 species.     

Ultrastructure and cuticle formation of the carapace in the myodocopan ostracod exemplified by Euphilomedes japonica (Crustacea: Ostracoda)

The ultrastructure and formation of the cuticle of a myodocopan ostracod, Euphilomedes japonica, are investigated utilizing scanning and transmission electron microscopy. The outer lamella cuticle consists of four layers; epicuticle, exocuticle, endocuticle, and membranous layer like in the cuticle of other arthropods. The exocuticle and endocuticle are well-calcified and the organic matrix develops within the both cuticles. The outermost layer of new cuticle (epicuticle) is secreted first and the inner layers (exocuticle, endocuticle and membranous layer) are added proximally in the pre-, and postmoult stages. The calcification takes place in the whole area of carapace at the same time together with the synthesis of organic matrix within the endocuticle. This study demonstrates that the ultrastructure and formation of the cuticle in myodocopans are different from those in podocopans, and that the myodocopan carapaces have achieved a structural diversity for adaptation to different lifestyles.