Geometric morphometrics reveal male genitalia differences in the Lasiommata megera/paramegaera complex (Lepidoptera, Nymphalidae) and the lack of a predicted hybridization area in the Tuscan Archipelago

The recognition of specific demarcation between allopatric populations, as for island endemics, is particularly difficult according to the Biological Concept of Species. However, the recognition of the specific status of island taxa is decisive in evaluating important areas of endemism for conservation purposes. The recent taxonomic inflation of European butterfly species calls into question the validity and objectivity of taxonomic practices, and of databases dependent on them, which are used in biogeography and conservation. In this paper I applied rigorous morphometric analyses instead of visual evaluation to solve the long debated question of whether Lasiommata paramegaera butterfly from Sardinia and Corsica should be considered as a separated entity from the widespread Lasiommata megera . First, I carried out analyses of male genitalia shape comparing the populations from Corsica and Tuscany. Thereafter, I also searched for possible hybridization evidence in the sole area where it is strongly predicted: the Tuscan Archipelago. These islands are located between Tuscany and Corsica and there is large evidence of a continuous immigration of butterflies from these two main sources. I found that Corsican L. paramegaera and Tuscan L. megera can be clearly distinguished on the basis of genitalia shape. Furthermore, the Lasiommata population of each islet clearly maintains the characteristics of one of the two species without evidence of intermediate individuals. In conclusion, I suggest that the two entities should be considered as separate species maintaining their homogeneity by a reduced gene flow across sea, mating avoidance and/or by some depletion of F1 hybrids in developmental or mating success.     

Effects of large-scale cattle grazing on populations of Coenonympha glycerion and Lasiommata megera (Lepidoptera:Satyridae)

The effects of large-scale cattle grazing by herdsmen (Allmende) on the populations of two butterfly species, Coenonympha glycerion and Lasiommata megera, were analysed on a study plot in the foothills of the Eastern Carpathians, Republic of Ukraine, in the summer and autumn of 1995. In all, 280 individuals of C. glycerion (115 females) and 230 individuals of L. megera (61 females) were individually marked. At recapture rates of 35 to 39%, maximum population size was calculated at 410 and 434 individuals. C. glycerion reacted to high densities by shifting to smaller neighbouring habitats. An increased emigration rate was recorded for L. megera when the population maximum was reached.Both species essentially require habitats formed by the activity of cattle. C. glycerion significantly preferred south facing woodland margins, close to moderately and infrequently grazed grassland. Heavily grazed areas were avoided. L. megera mainly occupied well-trod cattle paths or landslips with rather sparse vegetation, induced by cattle. Preferred habitats are characterized by linear structures for patrolling, steep slope areas and a wide range of nectar resources. Smaller, sporadically used cattle paths did not show these features and were occupied mainly by males at times of high population densities. Both species depend on patchy habitat mosaics which will be lost by intensification as well as by a complete cessation of grazing. The low-intensity, large-scale system utilized in the Eastern Carpathians is therefore favourable to both species.     

黄杨绢野螟雌蛾性信息素的鉴定及诱蛾活性

【目的】研究旨在深入探讨中国黄杨绢野Diaphania perspectalis 的雌蛾性信息素组成及诱蛾活性。【方法】利用气质联用仪（GC-MS）对黄杨绢野螟正常型性成熟雌蛾的性腺体提取物和合成标样比较分析,并用反-11-十六碳烯醛(E11-16:Ald) 、顺-11-十六碳烯醛（Z-11-16:Ald）、顺-9-十六碳烯醛（Z-9-16:Ald)、顺-11-十六碳烯醇（Z-11-16:OH)等物质进行触角的电生理测定,最后开展田间诱集比较试验以筛选出最佳性信息素混合物。【结果】Z-11-16:Ald为中国黄杨绢野螟正常型性信息素主要组分,E-11-16:Ald的含量极低,Z-11-16:OH未检测到。正常型雄性黄杨绢野螟触角对Z-11-16:Ald, E-11-16:Ald, Z-9-16:Ald和Z-11-16:OH产生强烈的EAG反应,并随着浓度的提高而显著增加;而对Z-11-16:Ac和E-11-16:Ac的嗅觉反应较弱,低于对植物绿叶气味顺3-己烯乙酸酯（Z-3-6:Ac）的反应。单一Z-1-16:Ald对正常型雄性黄杨绢野螟具有强烈的诱集效果,加入E-11-16:Ald有一定的增效作用,但在统计上则不显著。单一Z-11-16:Ald组分对黑化型雄性黄杨绢野螟无引诱活性,必需加入一定比例的E-11-16:Ald才显示诱蛾活性。Z-11-16:Ald:E-11-16:Ald的比例为250 μg:250 μg时诱集到的黑化型雄性黄杨绢野螟数量最多,而Z-11-16:Ald:E-11-16:Ald的比例为429 μg:71 μg时则诱集到的正常型雄性黄杨绢野螟数量最多。同时,单一Z-11-16:Ald也可引诱大量雄性粘虫Mythimna separata,但E-11-16:Ald抑制其活性。【结论】中国黄杨绢野螟的性信息素主成分是Z-11-16:Ald,单一组分即可在田间强烈引诱雄蛾,E-11-16:Ald的功能只起到微弱的增效作用,但也可能起种的专一性的作用。正常型与黑化型黄杨绢野螟对性信息素的嗅觉反应存在差异,黑化型黄杨绢野螟的性信息素接近日本种,即性信息素组成为Z-11-16:Ald和E-11-16:Ald 的混合物,其比例为1:1,且E-11-16:Ald为必需。     

Environmental sex reversal,Trojan sex genes,and sex ratio adjustment: conditions and population consequences

The great diversity of sex determination mechanisms in animals and plants ranges from genetic sex determination (GSD, e.g. mammals, birds, and most dioecious plants) to environmental sex determination (ESD, e.g. many reptiles) and includes a mixture of both, for example when an individual’s genetically determined sex is environmentally reversed during ontogeny (ESR, environmental sex reversal, e.g. many fish and amphibia). ESD and ESR can lead to widely varying and unstable population sex ratios. Populations exposed to conditions such as endocrine‐active substances or temperature shifts may decline over time due to skewed sex ratios, a scenario that may become increasingly relevant with greater anthropogenic interference on watercourses. Continuous exposure of populations to factors causing ESR could lead to the extinction of genetic sex factors and may render a population dependent on the environmental factors that induce the sex change. However, ESR also presents opportunities for population management, especially if the Y or W chromosome is not, or not severely, degenerated. This seems to be the case in many amphibians and fish. Population growth or decline in such species can potentially be controlled through the introduction of so‐called Trojan sex genes carriers, individuals that possess sex chromosomes or genes opposite from what their phenotype predicts. Here, we review the conditions for ESR, its prevalence in natural populations, the resulting physiological and reproductive consequences, and how these may become instrumental for population management.     

稻纵卷叶螟性信息素在其种群监测上的应用

为探索性信息素在稻纵卷叶螟Cnaphalocrocis medinalis（鳞翅目: 螟蛾科）预测预报上的应用技术, 2009年8月15日-9月20日在浙江省海宁市, 对不同性信息素化合物配制诱芯的田间诱捕效果, 以及性信息素群集诱捕、灯光诱捕和赶蛾3种不同方法监测成虫消长规律等进行研究。结果表明: 在浙江地区以顺11-十八碳烯醛、顺13-十八碳烯醛、顺11-十八碳烯醇和顺13-十八碳烯醇以60 μg∶500 μg∶60 μg∶120 μg配制的PVC毛细管诱芯对稻纵卷叶螟的诱集效果最佳。通过性信息素诱捕、灯光诱捕和赶蛾3种方法的比较试验, 初步结果显示性诱测报可取代传统的系统赶蛾法用于稻纵卷叶螟的预测预报。     

Effects of changing climate on species diversification in tropical forest butterflies of the genus Cymothoe (Lepidoptera: Nymphalidae)

Extant clades may differ greatly in their species richness, suggesting differential rates of species diversification. Based on phylogenetic trees, it is possible to identify potential correlates of such differences. Here, we examine species diversification in a clade of 82 tropical African forest butterfly species (Cymothoe), together with its monotypic sister genus Harma. Our aim was to test whether the diversification of the Harma–Cymothoe clade correlates with end‐Miocene global cooling and desiccation, or with Pleistocene habitat range oscillations, both postulated to have led to habitat fragmentation. We first generated a species‐level phylogenetic tree for Harma and Cymothoe, calibrated within an absolute time scale, and then identified temporal and phylogenetic shifts in species diversification. Finally, we assessed correlations between species diversification and reconstructed global temperatures. Results show that, after the divergence of Harma and Cymothoe in the Miocene (15 Mya), net species diversification was low during the first 7 Myr. Coinciding with the onset of diversification of Cymothoe around 7.5 Mya, there was a sharp and significant increase in diversification rate, suggesting a rapid radiation, and correlating with a reconstructed period of global cooling and desiccation in the late Miocene, rather than with Pleistocene oscillations. Our estimated age of 4 Myr for a clade of montane species corresponds well with the uplift of the Eastern Arc Mountains where they occur. We conclude that forest fragmentation caused by changing climate in the late Miocene as well as the Eastern Arc Mountain uplift are both likely to have promoted species diversification in the Harma–Cymothoe clade. Cymothoe colonized Madagascar much later than most other insect lineages and, consequently, had less time available for diversification on the island. We consider the diversification of Cymothoe to be a special case compared with other butterfly clades studied so far, both in terms of its abrupt diversification rate increase and its recent occurrence (7 Myr). It is clear that larval host plant shift(s) cannot explain the difference in diversification between Cymothoe and Harma; however, such a shift(s) may have triggered differential diversification rates within Cymothoe. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, ●● , ●●–●●.     

The species of the genus Zethera Felder (Lepidoptera: Nymphalidae,Satyrinae)

The butterfly genus Zethera Felder, endemic to the Philippines and northern Sulawesi, is shown to include six species, four of which comprise the Z. pimplea (Erichson) superspecies. Type data is given for all nominal taxa, and a key to the species recognised is presented, together with data on their variation and distribution. Attention is drawn to the zoogeographic subregions of the Philippine islands, as originally described by Semper, and illustrated by the Zethera species. Their mimicry is also noted, with particular reference to its sex-limitation in the Z. pimplea superspecies, and non-limitation in Z. incerta (Hewitson) and Z. hestioides C. & R. Felder. The need to demonstrate whether or not Zethera represents a true monophyletic unit is briefly discussed.     

稻纵卷叶螟对性信息素和花香的嗅觉行为反应及在其种群监测中的应用

[目的]明确稻纵卷叶螟Cnaphalocrocis medinalis生理状态对性信息素和花香气味的嗅觉行为反应的影响,以及在田间性信息素和花香气味诱蛾量的动态.[方法]于2020年在广西、湖南、浙江、四川5个试验点,通过田间网捕、花香气味和性信息素诱捕稻纵卷叶螟成虫,结合解剖卵巢和精巢,比较性信息素和花香诱捕的基本迁...     

Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae): classification, patterns of larval hostplant colonization and diversification

We present a higher‐level phylogenetic hypothesis for the diverse neotropical butterfly subfamily Ithomiinae, inferred from one of the largest non‐molecular Lepidoptera data sets to date, including 106 species (105 ingroup) and 353 characters (306 informative) from adult and immature stage morphology and ecology. Initial analyses resulted in 1716 most parsimonious trees, which were reduced to a single tree after successive approximations character weighting. The inferred phylogeny was broadly consistent with other past and current work. Although some deeper relationships are uncertain, tribal‐level clades were generally strongly supported, with two changes required to existing classification. The tribe Melinaeini is polyphyletic and Athesis + Patricia require a new tribe. Methona should be removed from Mechanitini into the restored tribe Methonini. Dircennini was paraphyletic in analyses of all data but monophyletic based on adult morphology alone, and its status remains to be confirmed. Hypothyris, Episcada, Godyris, Hypoleria and Greta are paraphyletic. A simulation analysis showed that relatively basal branches tended to have higher partitioned Bremer support for immature stage characters. Larval hostplant records were optimized on to a reduced, generic‐level phylogeny and indicate that ithomiines moved from Apocynaceae to Solanaceae twice, or that Tithoreini re‐colonized Apocynaceae after a basal shift to Solanaceae. Ithomiine clades have specialized on particular plant clades suggesting repeated colonization of novel hostplant niches consistent with adaptive radiation. The shift to Solanum, comprising 70% of neotropical Solanaceae, occurs at the base of a clade containing 89% of all ithomiines, and is interpreted as the major event in the evolution of ithomiine larval hostplant relationships. © The Willi Hennig Society 2006.     

Mechanisms that influence sex ratio variation in the invasive hymenopteran Sirex noctilio in South Africa

Sirex noctilio is an economically important invasive pest of commercial pine forestry in the Southern Hemisphere. Newly established invasive populations of this woodwasp are characterized by highly male‐biased sex ratios that subsequently revert to those seen in the native range. This trend was not observed in the population of S. noctilio from the summer rainfall regions in South Africa, which remained highly male‐biased for almost a decade. The aim of this study was to determine the cause of this persistent male bias. As an explanation for this pattern, we test hypotheses related to mating success, female investment in male versus female offspring, and genetic diversity affecting diploid male production due to complementary sex determination. We found that 61% of females in a newly established S. noctilio population were mated. Microsatellite data analysis showed that populations of S. noctilio from the summer rainfall regions in South Africa are far less genetically diverse than those from the winter rainfall region, with mean Nei's unbiased gene diversity indexes of 0.056 and 0.273, respectively. These data also identified diploid males at low frequencies in both the winter (5%) and summer (2%) rainfall regions. The results suggest the presence of a complementary sex determination mechanism in S. noctilio, but imply that reduced genetic diversity is not the main driver of the male bias observed in the summer rainfall region. Among all the factors considered, selective investment in sons appears to have the most significant influence on male bias in S. noctilio populations. Why this investment remains different in frontier or early invasive populations is not clear but could be influenced by females laying unfertilized eggs to avoid diploid male production in populations with a high genetic relatedness.     

Identification of two components of the female sex pheromone of the sugarcane‐borer Diatraea flavipennella (Lepidoptera: Crambidae)

Analysis by gas chromatography with electroantennographic detection of extracts of pheromone glands derived from calling females of the sugarcane‐borer Diatraea flavipennella revealed two antennally active compounds. These components were identified as (Z)‐9‐hexadecenal (Z9–16:Ald) and (Z)‐11‐hexadecenal (Z11–16:Ald) by comparison of the retention times of the natural compounds and the synthetic compounds supported by two‐dimensional gas chromatography – time‐of‐flight mass spectrometric analysis and the positions of the double bounds in the chains were confirmed from the mass spectral fragmentation patterns of their dimethyldisulphide adducts. The analysis indicated that Z9–16:Ald and Z11–16:Ald were present in the sex pheromone in the proportions 25 : 75. Trace amounts of tetradecanal, hexadecanal, (Z)‐7‐hexadecenal (Z7–16:Ald), (Z)‐9‐hexadecen‐1‐ol and (Z)‐11‐hexadecen‐1‐ol were also found in the extract, but of these only Z9–16:Ald and Z11–16:Ald appeared to be antennally active. Behavioural bioassays demonstrated that a binary blend composed of Z9–16:Ald and Z11–16:Ald in the ratio of 25 : 75 induced a response in D. flavipennella virgin males similar to that elicited by live virgin females or by an hexane extract of the pheromone glands of calling females. Z9–16:Ald and Z11–16:Ald are, therefore, considered to be the major constituents of the female sex pheromone of D. flavipennella.     

Spring remigration of the monarch butterfly, Danaus plexippus (Lepidoptera: Nymphalidae) in north-central Florida: estimating population parameters using mark-recapture

Monarch butterflies [Danaus plexippus) of the eastern North American population migrate each fall from die northern U.S.A. and southern Canada to overwintering sites in Mexico and return the following spring to the southeastern U.S.A. where they lay eggs and then die. The spring remigration is the least studied phase in the annual migration cycle. We therefore conducted a mark-recapture study and examined population recolonization dynamics and residence time in a north-central Florida pasture where the monarch's milkweed host plant (Asclepias humistrata) was abundant. Beginning in late March 1995 two waves of monarchs arrived, their numbers peaked at 71 individuals by mid-April, and the butterflies disappeared in early May. After arriving, the adults remained for 3–5 days, laid eggs and then continued to migrate. We also compared population sizes and arrival times in 1994 and 1996. We found no evidence of a second spring generation, which was also consistent with the deteriorating quality of the A. humistrata plants. Individuals of the new spring generation disappear shordy after eclosion. The arriving population was approximately nine times greater in 1995 than in 1996. Our findings support two recent hypodieses: (1) the bird-like migration of the monarch butterfly in North America evolved with the northward expansion and phenology of milkweeds; and (2) monarchs appear to be migratory throughout their annual cycle of several generations. By lingering for only a short time at each milkweed patch they encounter, the old monarchs returning from Mexico locate the resurgent milkweed flora over an extensive area in the southern states. Then, within less than a month, their fresh offspring continue the migration and exploit the unfolding cornucopia of milkweeds as the spring advances northward. The more we discover about the biology of this insect, the more remarkable is its annual migratory, breeding and overwintering cycle.     

Within population variation in the demography of Speyeria mormonia (Lepidoptera: Nymphalidae)

The adult demography of a population of Speyeria mormonia (Lepidoptera: Nymphalidae) near Gothic, Colorado USA was studied for four years. Values of demographic parameters, including survival, dispersal and sex ratio, varied yearly, without major changes in density. Shift in sex ratio with constant population numbers entails variation in effective population size among years. Flight period, catchability, relationship between body size and eclosion date, and significance of correlation between weather and daily survival rates differed between the sexes. Sexual dimorphism in correlation of wheather with survival rate may partially explain the lack of yearly variation in total population size. There was no significant sexual dimorphism in large scale habitat use, or in dispersal, except in 1982.
The demography of this insect is compared to that of related species of Argynnis and heliconiines, from the old world temperate zone and new world tropics, respectively. While some differences are observed which can be ascribed to differences in length of the growing season, many patterns are similar. This implies conservation of demographic traits, even across broad habitat and geographic areas.     

Phylogeny and diversification of the cloud forest Morpho sulkowskyi group (Lepidoptera,Nymphalidae) in the evolving Andes

The monophyletic Morpho sulkowskyi butterfly group, endemic of Andean cloud forests, was studied to test the respective contributions of Mio‐Pliocene intense uplift period and Pleistocene glacial cycles on Andean biodiversity. We sampled nine taxa covering the whole geographical range of the group. Two mitochondrial and two nuclear genes were analysed using a Bayesian method. We established a dated phylogeny of the group using a relaxed clock method and a wide‐outgroup approach. To discriminate between two hypotheses, we used a biogeographical probabilistic method. Results suggest that the ancestor of the M. sulkowskyi group originated during the Middle–Late Miocene uplift of the Eastern Cordillera in northern Peru. Biogeographical inference suggests that the M. sulkowskyi and Morpho lympharis clades diverged in the northern Peruvian Andes. The subsequent divergences, from the Late Miocene to the Late Pliocene, should have resulted from a dispersal towards the Northern Andes (M. sulkowskyi clade), after the closure of the West Andean Portal separating the Central and Northern Andes, and a southwards dispersal along the Peruvian and Bolivian Eastern Cordilleras (M. lympharis clade). Only a few divergences occurred at the very end of the Pliocene or during the Pleistocene, a period when the more recent uplifts interfered with Pleistocene glacial cycles.     

Host-plant influence on the population ecology of the jack pine budworm, Choristoneura pinus (Lepidoptera: Tortricidae)

Abstract.

• 1 Newly-emerged, second-instar jack pine budworm (Choristoneura pinus Freeman) establish spring feeding sites preferentially in the pollen cones of their host tree, Pinus banksiana Lamb.
• 2 Laboratory studies showed that the rate of establishment and survival of jack pine budworm on pollen cones was high throughout the entire spring emergence period of the insect.
• 3 In contrast, the rate of establishment and survival of jack pine budworm on vegetative buds was very poor early in the spring. Vegetative buds were only acceptable as feeding sites to the jack pine budworm for a relatively brief period in late spring.
• 4 Field studies showed that the change in population density of jack pine budworm during the spring emergence stage, as expressed by k-values, was a function of the abundance of pollen cones in the stand. Population reduction was greatest in those stands with the fewest pollen cones.
• 5 Direct measurement of spring dispersal by jack pine budworm showed that dispersal and consequent losses to the budworm population were greatest in stands with the fewest pollen cones.
• 6 We conclude that changes in the density of jack pine budworm are strongly influenced by production of pollen cones in the host stand. Because pollen cone production is related to previous years of defoliation by the jack pine budworm, we propose that pollen cones act as a density-dependent factor governing the density of early-stage jack pine budworm.
• 7 The resulting dynamics are compared to those of other budworm species and used to explain observed regional and temporal patterns of jack pine budworm outbreaks.

     

Male‐killer dynamics in the tropical butterfly,Acraea encedana (Lepidoptera: Nymphalidae)

Sex ratio distortion in the tropical butterfly Acraea encedana is caused by infection with a male‐killing bacterium of the genus Wolbachia. Previous research on this species has reported extreme female bias, high bacterial prevalences, and full sex role reversal. In this paper, we provide an assessment for the dynamics of the male‐killer, based on a survey for sex ratios and Wolbachia prevalences among wild populations of A. encedana in Uganda. The study reveals that Wolbachia infection showed considerable variation over both spatial and temporal scales.     

香梨优斑螟性信息素化学合成及雄蛾数量动态监测

香梨优斑螟是一种严重危害库尔勒香梨的重要害虫之一,利用昆虫性信息素防控和监测香梨优斑螟,可为香梨生产提供科学的绿色防控依据。本文通过化学合成方法得到香梨优斑螟性信息素活性组分:顺,反-9,12-十四碳二烯-1-醇(Z9E12-14∶OH),并利用标准化合物研究了香梨优斑螟雄蛾的野外发生规律。结果表明:香梨优斑螟性信息素活性组分对雄蛾有较好的引诱活性,诱蛾高峰十分明显,此虫在阿克苏地区包括越冬代在内一年发生3代,分别是4月中下旬、6月上中旬和7月中下旬。因此,该性引诱剂可为香梨优斑螟的综合防控提供技术支持。     

苹果蠹蛾性信息素的研究和应用进展

苹果蠹蛾Cydia pomonella (L.)是我国重要的检疫害虫, 在我国仅分布于新疆和甘肃以西局部地区, 但一直保持向我国东部扩张的趋势。在国际上, 利用性信息素监测和迷向防治苹果蠹蛾已经成为一种切实可行并广泛应用的害虫管理技术。本文综述了苹果蠹蛾性信息素的成分鉴定、人工合成和应用情况的研究进展, 指出了目前存在的问题和应用前景, 以期为我国苹果蠹蛾的防控策略的研究和应用提供参考。     

The effect of hybridization on the periodicity of sex pheromone release of Euxoa species (Lepidoptera: Noctuidae)

ABSTRACT. The periodicities of sex pheromone release by reciprocal F₁ hybrid females obtained from laboratory matings between Euxoa declarata (Walker) and E. rockburnei Hardwick were found to be dependent upon the parental cross. The period of sex pheromone release by females from the E. declarata (female) × E. rockburnei (male) cross paralleled that of the maternal parent species, E. declarata. However, female hybrid E. rockburnei (female) × E. declarata (male) had a prolonged calling period which intersected the pheromone release periods of both parental species and peaked later than that of E. declarata. These results indicate that the circadian periodicity of calling in these species is controlled by autosomal genes and a matroklinal inheritance factor.     

Varying rates of diversification in the genus Melitaea (Lepidoptera: Nymphalidae) during the past 20 million years

The influence of Quarternary glacial cycles on the extant diversity of Holarctic species has been intensively studied. It has been hypothesized that palaeoclimatic changes are responsible for divergence events in lineages. A constant improvement in DNA sequencing and modeling methods, as well as palaeoclimatic reconstruction, permit a deeper exploration of general causes of speciation in geological time. In the present study, we sampled, as exhaustively as possible, the butterflies belonging to the genus Melitaea (Lepidoptera: Nymphalidae), which are widely spread in the Palaearctic region. We conducted analyses to assess the phylogeny of the genus and estimated the timing of divergence and the most likely distribution of ancestral populations. The results obtained indicate that the systematics of the genus is in need of revision and that the diversity of the genus has been profoundly shaped by palaeoenvironmental changes during its evolutionary history. The present study also emphasizes that, when employed with caveats, major palaeoenvironmental events could represent very powerful tools for the calibration of the dating of divergences using molecular data.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 346–361.