An integumental anatomy for the butterfly Glaucopsyche lygdamus (Lepidoptera:Lycaenidae):a morphological terminology and homology

An integumental anatomy for the lycaenid butterfly Glaucopsyche lygdamus is presented. Comparisons with other lepidopteran taxa are made to rectify the homology of parts and contrast anatomical divergences within the Lycaenidae. A general terminology based on Snodgrass is given, to replace many of the specialized and often synonymous terms restricted to the Lepidoptera. Many common anatomical svnonyms are also given. Several reinterpretations of the anatomy and homology of various integumental regions are discussed. A previously unreported cuticular anomaly on abdominal tergum 2 of male Polyommatinae (Downey's area) is described. The following new or newly combined terms are used:postgenal-occipital area, postgenal-occipital protuberance, dorsal temporal sulci, postantennal projections, pronotal projection, infraepisternal-basisternal plate, paracoxal-marginopleural sulci, dorsal epimeral sulci, ventral epimeral sulci, secondary coxal sulci, ventral subcostal-radial process, lateral secondary sclerite and Downey's area.     

Analysis of museum specimens suggests extreme genetic drift in the adonis blue butterfly (Polyommatus bellargus)

We amplified microsatellite DNA from museum specimens over 100 years old of the adonis blue butterfly, Polyommatus bellargus. These results were compared with butterfly samples taken from the same site near Folkestone in southern UK in 1998/9, 200 generations later, and with samples from other extant UK populations. Dramatic changes in allele frequencies have occurred over time, which is indicative of substantial genetic drift or extinction/recolonization. Patterns of heterozygosity in the 1998/9 sample are indicative of a past bottleneck, and one was known to have occurred in the late 1970s in this and many other UK populations. One allele present at high frequency in 1896 was not detected in any extant UK population, suggesting that it may have been lost from the UK (a 'ghost' allele), although the allele may well persist elsewhere within the range of the species. Although the present study is relatively small in scale (20 museum specimens from one site), it serves to reinforce the enormous potential of museum specimens in well represented taxa such as butterflies for examining the effects of demographic events spanning many years.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 447–452.     

酢浆灰蝶生物学特性研究

通过野外观察、室内饲养等方法研究了酢浆灰蝶Pseudozizeeria maha(Kollar)的生活史及生物学特性。在南京地区,该蝶幼虫以酢浆草科Oxalidaceae的黄花酢浆草(Oxalis corniculataL.)为食,1年5代,世代交替发生。酢浆灰蝶在10月末以蛹在枯枝落叶或土壤表层浅洞中越冬,越冬代成虫次年5月始见、中旬为高峰期,5月中旬始见第1代卵及幼虫,6—10月间各月各虫态均同时发生,期间每月中下旬为成虫活动高峰期。幼虫共4龄,野外2～4龄幼虫具迁移、避热与适蚁行为。在室内饲养条件下,卵期为3～4d,幼虫期为22～26d,蛹期为6～7d。     

Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity

Lions were the most widespread carnivores in the late Pleistocene, ranging from southern Africa to the southern USA, but little is known about the evolutionary relationships among these Pleistocene populations or the dynamics that led to their extinction. Using ancient DNA techniques, we obtained mitochondrial sequences from 52 individuals sampled across the present and former range of lions. Phylogenetic analysis revealed three distinct clusters: (i) modern lions, Panthera leo ; (ii) extinct Pleistocene cave lions, which formed a homogeneous population extending from Europe across Beringia (Siberia, Alaska and western Canada); and (iii) extinct American lions, which formed a separate population south of the Pleistocene ice sheets. The American lion appears to have become genetically isolated around 340 000 years ago, despite the apparent lack of significant barriers to gene flow with Beringian populations through much of the late Pleistocene. We found potential evidence of a severe population bottleneck in the cave lion during the previous interstadial, sometime after 48 000 years, adding to evidence from bison, mammoths, horses and brown bears that megafaunal populations underwent major genetic alterations throughout the last interstadial, potentially presaging the processes involved in the subsequent end-Pleistocene mass extinctions.     

Reinstatement of a New Zealand copper butterfly,Lycaena rauparaha (Fereday, 1877)

Abstract

Previous confusion over the nomenclature and status of the New Zealand copper butterflies is reviewed. Lycaena rauparaha n. comb., which has been regarded as a variety of L. salustius in recent literature, is reinstated and adult, larval, and pupal characters are described. Its distinctions from L. salustius and L. feredayi are tabulated and a key to this species complex is given.     

A mirage of cryptic species: Genomics uncover striking mitonuclear discordance in the butterfly Thymelicus sylvestris

Mitochondrial DNA (mtDNA) sequencing has led to an unprecedented rise in the identification of cryptic species. However, it is widely acknowledged that nuclear DNA (nuDNA) sequence data are also necessary to properly define species boundaries. Next generation sequencing techniques provide a wealth of nuclear genomic data, which can be used to ascertain both the evolutionary history and taxonomic status of putative cryptic species. Here, we focus on the intriguing case of the butterfly Thymelicus sylvestris (Lepidoptera: Hesperiidae). We identified six deeply diverged mitochondrial lineages; three distributed all across Europe and found in sympatry, suggesting a potential case of cryptic species. We then sequenced these six lineages using double‐digest restriction‐site associated DNA sequencing (ddRADseq). Nuclear genomic loci contradicted mtDNA patterns and genotypes generally clustered according to geography, i.e., a pattern expected under the assumption of postglacial recolonization from different refugia. Further analyses indicated that this strong mtDNA/nuDNA discrepancy cannot be explained by incomplete lineage sorting, sex‐biased asymmetries, NUMTs, natural selection, introgression or Wolbachia‐mediated genetic sweeps. We suggest that this mitonuclear discordance was caused by long periods of geographic isolation followed by range expansions, homogenizing the nuclear but not the mitochondrial genome. These results highlight T. sylvestris as a potential case of multiple despeciation and/or lineage fusion events. We finally argue, since mtDNA and nuDNA do not necessarily follow the same mechanisms of evolution, their respective evolutionary history reflects complementary aspects of past demographic and biogeographic events.     

Reconstructing past species assemblages reveals the changing patterns and drivers of extinction through time

Predicting future species extinctions from patterns of past extinctions or current threat status relies on the assumption that the taxonomic and biological selectivity of extinction is consistent through time. If the driving forces of extinction change through time, this assumption may be unrealistic. Testing the consistency of extinction patterns between the past and the present has been difficult, because the phylogenetically explicit methods used to model present-day extinction risk typically cannot be applied to the data from the fossil record. However, the detailed historical and fossil records of the New Zealand avifauna provide a unique opportunity to reconstruct a complete, large faunal assemblage for different periods in the past. Using the first complete phylogeny of all known native New Zealand bird species, both extant and extinct, we show how the taxonomic and phylogenetic selectivity of extinction, and biological correlates of extinction, change from the pre-human period through Polynesian and European occupation, to the present. These changes can be explained both by changes in primary threatening processes, and by the operation of extinction filter effects. The variable patterns of extinction through time may confound attempts to identify risk factors that apply across time periods, and to infer future species declines from past extinction patterns and current threat status.     

Using genetics to inform re-introduction strategies for the Chequered Skipper butterfly (Carterocephalus palaemon, Pallas) in England

The Chequered Skipper butterfly (Carterocephalus palaemon) is extinct in England but extant in Scotland and Continental Europe. The possibility of re-introducing the species is under active consideration by conservation bodies, but ecological differences between Scottish and English populations raise the question of which populations should donate individuals, Continental European, or Scottish? We used mitochondrial DNA sequences (CO I, CO II and Cyt b) from potential donor populations to test the hypothesis that ecological differences could have arisen as a result of differing routes of post-glacial colonisation from separate refugia and subsequent isolation of UK populations. Shared haplotypes between populations in Belgium, Norway, Scotland and England provides no evidence to support the hypothesis that populations in Scotland result from an alternative post-glacial colonisation route. As the genetic evidence remains equivocal we suggest that choice of donor stock for a re-introduction to England should be made primarily on ecological grounds.     

Phylogeny and palaeoecology of Polyommatus blue butterflies show Beringia was a climate-regulated gateway to the New World

Transcontinental dispersals by organisms usually represent improbable events that constitute a major challenge for biogeographers. By integrating molecular phylogeny, historical biogeography and palaeoecology, we test a bold hypothesis proposed by Vladimir Nabokov regarding the origin of Neotropical Polyommatus blue butterflies, and show that Beringia has served as a biological corridor for the dispersal of these insects from Asia into the New World. We present a novel method to estimate ancestral temperature tolerances using distribution range limits of extant organisms, and find that climatic conditions in Beringia acted as a decisive filter in determining which taxa crossed into the New World during five separate invasions over the past 11 Myr. Our results reveal a marked effect of the Miocene-Pleistocene global cooling, and demonstrate that palaeoclimatic conditions left a strong signal on the ecology of present-day taxa in the New World. The phylogenetic conservatism in thermal tolerances that we have identified may permit the reconstruction of the palaeoecology of ancestral organisms, especially mobile taxa that can easily escape from hostile environments rather than adapt to them.     

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.     

Paleogenomics reveals independent and hybrid origins of two morphologically distinct wolf lineages endemic to Japan

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Modelling the impacts of two exotic invasive species on a native butterfly: top-down vs. bottom-up effects

1. Exotic invasive species can influence population dynamics of native species through top-down or bottom-up forces. The present study examined separate and interactive effects of multiple exotic species invasions on the native mustard white butterfly, Pieris napi oleracea Harris (Lepidoptera: Pieridae), using a stochastic simulation model. 2. P. n. oleracea populations in North America have decreased regionally since the 1860s. Competition with an exotic congener (P. rapae L.), loss of native host plants and parasitism by the introduced broconid wasp (Cotesia glomerata L.), have been suggested to be independently responsible for its decline. The present study examined these hypotheses, as well as an alternative, invasion by an exotic crucifer, garlic mustard (Alliaria petiolata[Bieb.] Cavara & Grande). 3. A stochastic simulation model of P. n. oleracea population dynamics revealed that decreasing the number of host plants available for oviposition and larval development (i.e. habitat loss), sharply reduced the probability of populations persistence and decreased population size for those that persisted. 4. Simulated invasion by garlic mustard also substantially decreased both probability of persistence (= 0 at approximately 50% cover) and mean population size. Persistence probability never reached zero under any C. glomerata scenarios, even when larval mortality in the second generation due to parasitism was 100%. The impact of garlic mustard was intensified by the addition of C. glomerata parasitism. 5. Results suggest that bottom-up forces, loss of host plants through forest understorey loss and/or garlic mustard invasion are the most important forces driving P. n. oleracea population decline. Parasitism by C. glomerata may interact to reduce P. n. oleracea populations more rapidly, but appears insufficient alone to cause local extinction.     

Application of the unified species concept reveals distinct lineages for disjunct endemics of the Brassica repanda (Brassicaceae) complex

Species delimitation is an important issue in terms of conservation priorities, especially for narrow endemics under threat of extinction. The Alpine endemics Brassica repanda subsp. glabrescens and subsp. baldensis belong to a highly polymorphic species complex, although their disjunct distribution suggests favourable conditions for independent evolution. In the present study, we applied the unified species concept to test whether the endemics form distinct evolutionary lineages, both from one another and also from the remainder of the complex. Compliance with the criteria of monophyly, diagnosability, and genotypic clustering was examined, primarily by making use of amplified fragment length polymorphism data. Both endemics were indicated as monophyletic by phylogenetic analyses, and diagnostic characters were found for both taxa. Population structure analyses showed clear genetic discontinuity for each of the endemics, with little admixture among the clusters. This evidence indicates that the endemics have acquired multiple properties that satisfy each of the species criteria considered. Hence, we suggest the taxonomic recognition of B. baldensis and B. glabrescens as separate species. Comparative population genetics analyses show the lack of marked genetic structuring within either taxon, as well as low levels of heterozygosity. Conclusions on the status of threat and on recommended conservation actions are drawn. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 482–497.     

Bacterial communities within Phengaris (Maculinea) alcon caterpillars are shifted following transition from solitary living to social parasitism of Myrmica ant colonies

Bacterial symbionts are known to facilitate a wide range of physiological processes and ecological interactions for their hosts. In spite of this, caterpillars with highly diverse life histories appear to lack resident microbiota. Gut physiology, endogenous digestive enzymes, and limited social interactions may contribute to this pattern, but the consequences of shifts in social activity and diet on caterpillar microbiota are largely unknown. Phengaris alcon caterpillars undergo particularly dramatic social and dietary shifts when they parasitize Myrmica ant colonies, rapidly transitioning from solitary herbivory to ant tending (i.e., receiving protein‐rich regurgitations through trophallaxis). This unique life history provides a model for studying interactions between social living, diet, and caterpillar microbiota. Here, we characterized and compared bacterial communities within P. alcon caterpillars before and after their association with ants, using 16S rRNA amplicon sequencing and quantitative PCR. After being adopted by ants, bacterial communities within P. alcon caterpillars shifted substantially, with a significant increase in alpha diversity and greater consistency in bacterial community composition in terms of beta dissimilarity. We also characterized the bacterial communities within their host ants (Myrmica schencki), food plant (Gentiana cruciata), and soil from ant nest chambers. These data indicated that the aforementioned patterns were influenced by bacteria derived from caterpillars’ surrounding environments, rather than through transfers from ants. Thus, while bacterial communities are substantially reorganized over the life cycle of P. alcon caterpillars, it appears that they do not rely on transfers of bacteria from host ants to complete their development.     

Development and characterization of polymorphic microsatellite loci in the endangered Miami blue butterfly (Cyclargus thomasi bethunebakeri)

The Miami blue butterfly (Cyclargus thomasi bethunebakeri) is a state-endangered taxon in Florida and a candidate for federal listing. Here we report 12 polymorphic microsatellite loci appropriate for use in population and conservation studies. We genotyped 114 individuals sampled from a metapopulation in the lower Florida Keys over a 2-year period (2005-2006). These results show 4-14 alleles per locus, and ranges of observed and expected heterozygosities are 0.02679-0.79630 and 0.06154-0.69565, respectively. Large deviations from Hardy-Weinberg equilibrium (HWE) are observed across the whole sample set. When a single breeding population is analysed alone, seven of the loci are in HWE.     

Body length of bony fishes was not a selective factor during the biggest mass extinction of all time

The Permo‐Triassic mass extinction devastated life on land and in the sea, but it is not clear why some species survived and others went extinct. One explanation is that lineage loss during mass extinctions is a random process in which luck determines which species survive. Alternatively, a phylogenetic signal in extinction may indicate a selection process operating on phenotypic traits. Large body size has often emerged as an extinction risk factor in studies of modern extinction risk, but this is not so commonly the case for mass extinctions in deep time. Here, we explore the evolution of non‐teleostean Actinopterygii (bony fishes) from the Devonian to the present day, and we concentrate on the Permo‐Triassic mass extinction. We apply a variety of time‐scaling metrics to date the phylogeny, and show that diversity peaked in the latest Permian and declined severely during the Early Triassic. In line with previous evidence, we find the phylogenetic signal of extinction increases across the mass extinction boundary: extinction of species in the earliest Triassic is more clustered across phylogeny compared to the more randomly distributed extinction signal in the late Permian. However, body length plays no role in differential survival or extinction of taxa across the boundary. In the case of fishes, size did not determine which species survived and which went extinct, but phylogenetic signal indicates that the mass extinction was not a random field of bullets.     

Parallel tagged amplicon sequencing reveals major lineages and phylogenetic structure in the North American tiger salamander (Ambystoma tigrinum) species complex

Modern analytical methods for population genetics and phylogenetics are expected to provide more accurate results when data from multiple genome‐wide loci are analysed. We present the results of an initial application of parallel tagged sequencing (PTS) on a next‐generation platform to sequence thousands of barcoded PCR amplicons generated from 95 nuclear loci and 93 individuals sampled across the range of the tiger salamander (Ambystoma tigrinum) species complex. To manage the bioinformatic processing of this large data set (344 330 reads), we developed a pipeline that sorts PTS data by barcode and locus, identifies high‐quality variable nucleotides and yields phased haplotype sequences for each individual at each locus. Our sequencing and bioinformatic strategy resulted in a genome‐wide data set with relatively low levels of missing data and a wide range of nucleotide variation. structure analyses of these data in a genotypic format resulted in strongly supported assignments for the majority of individuals into nine geographically defined genetic clusters. Species tree analyses of the most variable loci using a multi‐species coalescent model resulted in strong support for most branches in the species tree; however, analyses including more than 50 loci produced parameter sampling trends that indicated a lack of convergence on the posterior distribution. Overall, these results demonstrate the potential for amplicon‐based PTS to rapidly generate large‐scale data for population genetic and phylogenetic‐based research.