Olfaction and cyanide sensitivity in the six-spot burnet mothZygaena filipendulae and the silkmothBombyx mori

Summary Olfactory receptor and standing potentials of maleZygaena filipendulae andBombyx mori were recorded from isolated antennae before and after exposure to hydrogen cyanide. Exposure of two minutes caused a transient decrease and complete recovery of both potentials inZ filipendulae, while an exposure of 0.5 min led to disappearance of these parameters inB. mori. Mechanisms of cyanide resistance are discussed.We thank Dr. E. Priesner for hints concerning biotopes ofZ. filipendulae, Miss I. Block for experimental help, Drs. U. Thurm and R. Villet for helpful criticism, and the Stiftung Volkswagenwerk for financial support.     

Volatiles from the burnet moth Zygaena filipendulae (Lepidoptera) and associated flowers,and their involvement in mating communication

The burnet moth Zygaena filipendulae L. contains the cyanogenic glucosides linamarin and lotaustralin, which can be degraded to the volatiles hydrogen cyanide (HCN), acetone and 2‐butanone. Linamarin and lotaustralin are transferred from the male to female during mating and thus are considered to be involved in mating communication. Because volatile semiochemical cues play a major role in mating communication in many insect species, the emissions of HCN, acetone and 2‐butanone from Z. filipendulae are characterized in the present study, aiming to determine the interplay between the degradation products of cyanogenic glucosides and pheromones. The volatile emissions from Z. filipendulae and flowers inducing mating are measured using headspace solid‐phase micro‐extraction and gas chromatography‐mass spectrometry analysis. All Z. filipendulae life stages emit HCN, acetone and 2‐butanone. Virgin females show higher emissions than mated females, whereas mated males have higher emissions than virgin males. Hydrogen cyanide is only rarely detected in the course of male–female copulation. These observations indicate a role for the cyanogenic glucoside derived volatiles in female calling and male courtship behaviours, although not as a defence during copulation. Males rejected for mating by a female are accepted after injection of linamarin or lotaustralin, demonstrating that cyanogenic glucosides are also important for female assessment of the fitness of the male. Volatiles from flowers occupied during mate calling are also analyzed, and emissions from males and females result in the identification of novel putative pheromones for Z. filipendulae.     

Contrasting movement patterns in two species of chequerspot butterflies, Euphydryas aurinia and Melitaea phoebe, in the same patch network

Abstract.  1. Mark–release–recapture studies were conducted on two species of chequerspot butterfly, Euphydryas aurinia and Melitaea phoebe , in the same habitat patch network in Yanjiaping, a small basin in the Taihang Mountains, north-west of Beijing, China, in 2000.
2. Euphydryas aurinia tended to stay in the habitat patches and to move to neighbouring patches, whereas M. phoebe moved widely among patches in the entire network.
3. The parameters of the virtual migration model showed higher daily emigration propensity in M. phoebe and in E. aurinia males than in E. aurinia females, and significantly greater average daily movement distance in M. phoebe than in E. aurinia .
4. The results are consistent with the previous findings showing genetic structuring among local populations of E. aurinia but not among local populations of M. phoebe .
5. Based on the genetic and ecological results, it was concluded that E. aurinia has a classic metapopulation in the study area, whereas M. phoebe appears to have a source–sink metapopulation.
6. In 2000, when there was an overall increase in the abundance of the two species, the limited mobility of E. aurinia resulted in an increase in the average local population size, whereas the increase in the number of local populations in M. phoebe was due to its high mobility .     

Survival, movement, and resource use of the butterfly Parnassius clodius

Abstract.  1. A mark–recapture study was conducted on the American Apollo butterfly Parnassius clodius Menetries during three field seasons (1998–2000) to examine its movement patterns over the course of a season within a sagebrush meadow in Grand Teton National Park, Wyoming, U.S.A. The study examined how resources affected butterfly distribution patterns and used mark–recapture data to gain insight into movement differences between sexes and over time.
2. The average straight-line movement of P. clodius was 202 m day⁻¹, adjusted for sampling effort at different distances. Movement estimates in all 3 years were highly correlated with the average distance between plots sampled.
3. Butterfly abundance was correlated positively with per cent cover of its host plant Dicentra uniflora , but this relationship decreased in importance during the peak of the flight period when individuals may be more interested in finding mates. There was a weak, positive correlation between butterfly abundance and the abundance of its primary nectar source, Eriogonum umbellatum in 1999, but no relationship in 2000.
4. Survival, recapture, and transition probabilities were estimated using open population, capture–recapture models. Survival and recapture probability decreased over the course of each season, while the probability of moving between plots increased. Recapture probability was significantly lower for females than for males among all 3 years, but there was no difference between the sexes in survival rate.     

Genetic differentiation across North America in the generalist moth Heliothis virescens and the specialist H. subflexa

The two moth species Heliothis virescens (Hv) and H. subflexa (Hs) are closely related, but have vastly different feeding habits. Hv is a generalist and an important pest in many crops in the USA, while Hs is a specialist feeding only on plants in the genus Physalis. In this study, we conducted a comparative population genetic analysis to assess whether and how generalist and specialist life styles are reflected in differences in population structures. In Hv 98% of the total variation occurred within populations. The overall differentiation (F(ST) ) between regions was 0.006 and even lower between years (0.0039) and hosts (0.0028). Analyses of population structure suggest that all individuals form one genetically homogeneous population, except for at most 12 individuals (6%) that diverged from this cluster. Population homogeneity likely results from the high mobility of Hv and its generalist feeding behaviour. Hs exhibited substantially more population structure. Even though 96% of the total variation was attributable to within-population variability, F(ST) -values between Hs populations were 10 times higher than between Hv populations. Hs populations showed significant isolation by distance. Analyses of Hs population structure suggest at least two subpopulations and thus some degree of metapopulation structure. We speculate that the patchy distribution of Physalis- the exclusive food source of Hs - contributes to differences in population structure between these closely related species. The finding that the specialist shows more population differentiation than the generalist corroborates the notion that host specialization is not an evolutionary dead end but a dynamic trait.     

Sex differences but no evidence of quantitative honesty in the warning signals of six‐spot burnet moths (Zygaena filipendulae L.)*

The distinctive black and red wing pattern of six‐spot burnet moths (Zygaena filipendulae, L.) is a classic example of aposematism, advertising their potent cyanide‐based defences. While such warning signals provide a qualitatively honest signal of unprofitability, the evidence for quantitative honesty, whereby variation in visual traits could provide accurate estimates of individual toxicity, is more equivocal. Combining measures of cyanogenic glucoside content and wing color from the perspective of avian predators, we investigate the relationship between coloration and defences in Z. filipendulae, to test signal honesty both within and across populations. There were no significant relationships between mean cyanogenic glucoside concentration and metrics of wing coloration across populations in males, yet in females higher cyanogenic glucoside levels were associated with smaller and lighter red forewing markings. Trends within populations were similarly inconsistent with quantitative honesty, and persistent differences between the sexes were apparent: larger females, carrying a greater total cyanogenic glucoside load, displayed larger but less conspicuous markings than smaller males, according to several color metrics. The overall high aversiveness of cyanogenic glucosides and fluctuations in color and toxin levels during an individual's lifetime may contribute to these results, highlighting generally important reasons why signal honesty should not always be expected in aposematic species.     

Ecological differentiation between the Sardinian endemic Maniola nurag and the pan-European M. jurtina

Recently refined evolutionary theories have highlighted that ecological interactions and environmental gradients can play a major role in speciation. This paper reports on a 3‐year field study, in which the ecology of two congeneric butterfly species was used to explore and compare the environmental factors determining their spatial distribution. These data are discussed in the context of possible speciation scenarios between the Sardinian populations of Maniola nurag and M. jurtina. M. nurag is endemic to the island of Sardinia, while M. jurtina is widespread over Europe. In Sardinia, the two species are locally sympatric. Mark–release–recapture experiments were combined with measures of environmental variables in 15 1‐ha plots, established in areas of potential habitat for the butterflies. Constrained linear models were parameterized from mark–recapture data to estimate both individual (survival and capture probabilities) and population (population size and recruitment) parameters. The two species had similar demography, movement patterns, life history, and behaviour. Population sizes developed in a parabolic fashion from beginning to end of the flight season. Differences included local population size, adult phenology, and habitat requirements. Long‐distance movements larger than 1.5 km were observed, suggesting a substantial amount of gene‐flow between populations of the endemic as well as the widespread species. Multivariate analyses revealed four main environmental gradients responsible for the abundance of the butterflies in an area. Both species responded similarly to environmental variables. However, each species’s abundance was correlated with a different environmental gradient determined by vegetation cover and structure. When sympatric, the two species responded to subtle differences in microhabitat structure. This might originally have induced their divergence. This study is an example of how empirical field data on population dynamics, dispersal, and habitat characteristics of two sympatric congeners can further our understanding of how species differentiate despite existing gene‐flow. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89 , 561–574.     

Movement of two grassland butterflies in the same habitat network: the role of adult resources and size of the study area

Abstract. 1.  Movement patterns of two butterfly species (meadow brown Maniola jurtina L. and scarce copper Lycaenae virgaureae L.) were studied in a 172 ha area within a landscape with a high percentage of suitable habitats for mark–release–recapture experiments.
2.  Adult resource density, but not patch size or larval food plant abundance, influenced the numbers and the fractions of residents, emigrants, and immigrants.
3.  Differences between species were observed in movement frequency and maximum distances moved but not in mean distances moved.
4.  The scarce copper showed much greater movement ability than expected from the results of published studies. This is believed to be a result of the comparatively large size of the study area and the high cover of suitable habitat (>50%).
5.  The mean and maximum distances travelled by butterflies reflected differences in the size of the study area.     

Mimicry in the burnet moth Zygaena ephialtes: population studies and evidence of a Batesian—Müllerian situation

Abstract. 1. Population size as well as the absolute and relative frequencies of the phegea and filipendulae mimetic complexes have been estimated in an area of Central Italy selected because it is: (a) little affected by man, (b) representative of many others in the Central Appennines, and (c) containing an abundant population of Zygaena ephialtes.
2. This distasteful polymorphic species belongs, in Central Italy as in other Southern areas, to the phegea complex ( Zygaena ephialtes , yellow ephialtoid form), while in Northern and Central Europe it belongs to the filipendulae complex ( Z.ephialtes , red peucedanoid form).
3. In the Southern areas the phegea complex is much more abundant than the filipendulae one, which gives the yellow ephialtoid form of Z.ephialtes a strong mimetic advantage over the red peucedanoid.
4. In addition, Amata phegea greatly outnumbers the other distasteful members of its complex and emerges first.
5. Throughout its flight period Z.ephialtes , which emerges about 20 days later than A.phegea , never attains a frequency higher than 0.03 in this complex. It is suggested that in such a situation the protection that this species gains as a mimic would be very high regardless of its unpalatability.
6. These findings and others discussed in the present paper, such as the degree of unpalatability, the genetics of the mimetic form as well as the polymorphism for colour and pattern, are examined according to the hypothesis of divergent evolutionary strategies in Batesian and Müllerian mimicry. It is suggested that these are meaningful concepts but that individual mimics, of which Z.ephialtes is one, can fall between them.     

Molecular biogeography of the arctic-alpine disjunct burnet moth species Zygaena exulans (Zygaenidae, Lepidoptera) in the Pyrenees and Alps

Aim The phylogeography of ‘southern’ species is relatively well studied in Europe. However, there are few data about ‘northern’ species, and so we studied the population genetic structure of the arctic‐alpine distributed burnet moth Zygaena exulans as an exemplar. Location and methods The allozymes of 209 individuals from seven populations (two from the Pyrenees, five from the Alps) were studied by electrophoresis. Results All 15 analysed loci were polymorphic. The mean genetic diversities were moderately high (A: 1.99; H_e: 11.5; P: 65%). Mean genetic diversities were significantly higher in the Alps than in the Pyrenees in all cases. F_ST was 5.4% and F_IS was 10%. Genetic distances were generally low with a mean of 0.022 between large populations. About 62% of the variance between populations was between the Alps and the Pyrenees. The two samples from the Pyrenees had no significant differentiation, whereas significant differentiation was detected between the populations from the Alps (F_ST = 2.8%, P = 0.02). Main conclusion Zygaena exulans had a continuous distribution between the Alps and the Pyrenees during the last ice age. Most probably, the species was not present in Iberia, and the samples from the Pyrenees are derived from the southern edge of the glacial distribution area and thus became genetically impoverished. Post‐glacial isolation in Alps and Pyrenees has resulted in a weak genetic differentiation between these two disjunct high mountain systems.     

Migration to and dispersal from oilseed rape by the pollen beetle,Meligethes aeneus,in relation to wind direction

1 Host‐plant‐odour‐induced upwind anemotaxis is accepted as the main mechanism by which herbivorous insects find their host plant within an heterogenous environment, but field data supporting this hypothesis are scarce. 2 The flight directions of the pollen beetle Meligethes aeneus to and from a plot of winter oilseed rape and the direction of the wind were recorded concurrently. The beetles were sampled by eight double‐sided window traps encircling the plot. Distal sides of the traps, relative to the plot, sampled the beetles as they flew towards the plot, whereas the proximal sides of the traps sampled them as they flew away from the plot. Paired data on daily catch of beetles in distal or proximal traps and the volume of air impacting each trap were compared. 3 Correlations between daily catch of M. aeneus into distal traps and trap wind volume were negative, indicating that flights by both overwintered‐ and new‐generation insects towards the plot were upwind. 4 Correlations between daily catch of M. aeneus into proximal traps and trap wind volume varied with generation. Catch of overwintered‐generation M. aeneus was negatively correlated with trap wind volume, indicating that flight was upwind. Proximal trap catch of the new‐generation M. aeneus was weakly/modestly positively correlated with trap wind volume, indicating that flights were downwind or crosswind. 5 Understanding the effect of wind direction on flight of M. aeneus holds potential for improving the forecasting of their arrival and spatial distribution on the crop for integrated pest management.     

Comparative analysis of peach and pear fruit volatiles attractive to the oriental fruit moth,Cydia molesta

Cydia molesta is an important pest of peach and pear fruits late in growing season. We identified and quantified volatiles from immature and mature fruits of peach and pear using gas chromatography-mass spectrometry. Antennally, active compounds were identified by gas chromatography-electroantennogram detection and were further tested in the field. Consistent electroantennographic activity was obtained for nine compounds from headspace collections of the mature fruits of two peach and two pear varieties. Esters were the major components, but qualitative and quantitative differences were found among four odor profiles. Blends mimicking pear fruit volatiles were more attractive to both sexes than blends mimicking peach fruit volatiles in both orchards. Our finding indicated that mixtures mimicking peach and pear fruit volatiles attracted both females and males of C. molesta, and these host volatiles may also be involved in the host switch behavior from peach to pear in mid-late season.     

Irradiation does not affect field dispersal ability in the West Indian sweetpotato weevil, Euscepes postfasciatus

The effect of irradiation on the dispersal ability of males and females of the flightless West Indian sweetpotato weevil, Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae), a major sweet potato pest, was examined in the field using mark–release–recapture techniques. To evaluate the dispersal ability of the weevil, we released 7 619 weevils in two replicates (July and August 2007). Each replicate lasted 5 days from release to sampling and consisted of one weevil release and two weevil samplings. Thirty-two traps were placed in lines corresponding to eight compass directions and in four distance classes (8, 12, 16, and 20 m) in each replicate. We captured 709 (9.3%) weevils in the two replicates. Weevils dispersed at least 20 m from the release point in 2 days, regardless of sex or irradiation. Dispersal was strongly affected by wind direction, and in both replicates most weevils were recaptured in upwind directions. The mean dispersal distance for non-irradiated weevils was about 11 m per 2 days. Although there were some differences between sexes in recapture rate and dispersal distance, there was no consistent difference between irradiated and non-irradiated weevils in dispersal distance. We conclude that irradiation does not affect the dispersal ability of flightless E. postfasciatus in the field.     

Experimental Density-Related Emigration in the Cranberry Fritillary Boloria aquilonaris

Studies on sedentary butterflies often show that most individuals cover only short distances and rarely leave their original patch. An important question for metapopulation dynamics and conservation is “When and why do sedentary species disperse?” To imitate high population densities I harassed individuals of the sedentary butterfly Boloria aquilonaris with a dummy and recorded the time that passed until they emigrated from their original bogpatch of 800 m². Harassed individuals emigrated significantly sooner than unharassed ones. Repeated harassment following their return to the marked patch led to an increase in the residence times of these individuals outside the patch. Individuals receiving a potentially detrimental stimulus (e.g., harassment) are more likely to emigrate sooner than other individuals. The results support the hypothesis that emigration is density dependent in this butterfly, such that gene flow and colonization may be episodic.     

The role of group size and environmental factors on survival in a cooperatively breeding tropical passerine

1. Variation in survival, a major determinant of fitness, may be caused by individual or environmental characteristics. Furthermore, interactions between individuals may influence survival through the negative feedback effects of density dependence. Compared to species in temperate regions, we have little knowledge about population processes and variation in fitness in tropical bird species. 2. To investigate whether variation in survival could be explained by population size or climatic variables we used capture-recapture models in conjunction with a long-term data set from an island population of the territorial, cooperatively breeding Seychelles warbler (Acrocephalus sechellensis). The lack of migration out of the study population means that our results are not confounded by dispersal. 3. Annual survival was high, both for adults (84%) and juveniles (61%), and did not differ between the sexes. Although there was significant variation in survival between years, this variation could not be explained by overall population size or weather variables. 4. For territorial species, resource competition will work mainly on a local scale. The size of a territory and number of individuals living in it will therefore be a more appropriate measure of density than overall population density. Consequently, both an index of territory quality per individual (food availability) and local density, measured as group size, were included as individual covariates in our analyses. 5. Local density had a negative effect on survival; birds living in larger groups had lower survival probabilities than those living in small groups. Food availability did not affect survival. 6. Our study shows that, in a territorial species, although density-dependent effects might not be detectable at the population level they can be detected at the individual territory level - the scale at which individuals compete. These results will help to provide a better understanding of the small-scale processes involved in the dynamics of a population in general, but in particular in tropical species living in relatively stable environments.     

Volatile compounds from young peach shoots attract males of oriental fruit moth in the field

Abstract

The oriental fruit moth (OFM) is one of the most serious pests of commercial fruit orchards worldwide. Newly planted peach trees in particular, can be very attractive for mated OFM females for oviposition. Samples of airborne host plant volatiles from intact young peach shoot tips and old leaves of the same potted plants were collected and analyzed with a gas chromatograph-mass spectrometer. Chemicals present in young shoot tips, but not in old leaves, were used for field trials. Moth capture by traps with the synthetic chemicals was compared to that of the standard terpinyl acetate (TA) food trap. The TA food trap caught OFM males and mated females, but tested synthetic chemicals trapped only OFM males. We observed that the mixture of (Z)-3-hexenyl acetate: (E)-β-ocimene: (E)-β-farnesene in proportion 1:2:2 attracted OFM males. Further, 1 mg of (E)-β-ocimene, and that of (E)-β-farnesene also attracted OFM males.     

Repeated larval diapause and diapause-free development in geographic strains of the burnet moth Zygaena trifolii Esp. (Insecta,Lepidoptera)

Summary Zygaena trifolii is a long-day insect with temperature-dependent photoperiodic responses. All larval instars are sensitive to photoperiod; however, diapause may occur at the third larval stage or any subsequent larval instars. There were quantitative differences within populations in the threshold photoperiod for diapause induction. The diapause response was polymorphic, so that larvae might enter diapause at different instars under the same culture conditions. Furthermore, decreasing photoperiods below a critical daylength shifted the diapausing instar towards earlier stages. Geographic strains of Z. trifolii showed discontinuous clinal variation. Near the northern edge of the distribution [Cologne (Köln), FRG], there is first an obligatory diapause, mainly during early instars, and additional facultative (repeat) diapauses during later larval instars in subsequent years. In the southern part of its distribution, this burnet moth is partially bivoltine in the field with a facultative first developmental arrest and a decreased capacity for repeated diapause (Valencia, Spain; Marseille, France). Further experiments indicated that the photoperiodically controlled diapause reaction is also influenced by the number of photoperiodic cycles experienced during the period spent in each larval instar, which depends on temperature. The adaptive significance of obligatory and facultative repeated diapause, varying even among the offspring of a single female, may be to buffer the populations against the more extreme and, from year to year, unpredictable fluctuations in climatic conditions at the northern edge of the distribution.Abbreviations L₃ feeding 3rd larval instar - L₄D diapausing 4th larval instar - L₅D2 repeat-diapausing larval instar with second diapause at the 5th larval stage - LD light-dark cycle - KT shortday conditions (e.g. LD 8:16) - LT long-day conditions (e.g. LD 16:8)     

Determining the density dependence of immigration and emigration of benthic stream invertebrates: theoretical considerations

Simple mathematical models are formulated to describe density independent and density dependent dispersal. These models clarify hypotheses of density dependence and may be manipulated easily to suit particular applications. The models demonstrate that the initial composition of a species aggregate must be controlled before valid conclusions can be drawn about the density dependency of the aggregate's dispersal. Stochastic models of emigration are derived to assess the power of particular experimental designs and statistical techniques to discriminate a known form of density dependent emigration. Contribution No. 369, Great Lakes Research Division, University of Michigan Contribution No. 369, Great Lakes Research Division, University of Michigan     

梨小食心虫化学通信中的信息物质

梨小食心虫Grapholitha molesta Busck是我国北方果区发生的重要害虫。当前对该虫的防治主要依赖化学农药, 但引起的害虫抗药性、 杀伤天敌和环境污染等问题非常严重。食心虫自身的信息素、 寄主/非寄主的他感化学物质对于调节其配偶选择和寄主定位起着至关重要的作用, 基于信息化学物质的害虫管理策略为果园食心虫的治理提供了新的途径。本文综述了国内外有关梨小食心虫化学信息物质研究进展, 包括雌蛾释放的性信息素组分及对雄、 雌两性的引诱, 雄蛾释放的信息素, 利用性信息素的迷向研究, 寄主植物挥发性引诱物质的鉴定和筛选, 以及梨小食心虫寄主转换机制等方面的研究现状和存在的问题。具体来说, 雌蛾的性信息素包括顺-8-十二碳烯醋酸酯、反-8-十二碳烯醋酸酯、顺-8-十二碳烯-1-醇和十二碳-1-醇, 各个组分的比例在地理种群间存在变异。在室内, 通过行为试验证实两种醋酸酯对雄蛾的引诱是必不可少的, 微量的顺-8-十二碳烯-1-醇对二元组分起到增效作用。在田间, 上述3种物质组成的诱芯具有较强的活性; 由此开发的性信息素迷向技术（人工迷向丝、 可喷施的微胶囊和蜡滴）被用于梨小食心虫交配干扰, 取得了很好的效果。梨小食心虫最主要的寄主植物桃梢挥发物包括22种化合物, 其中绿叶挥发物占到50%, 行为生测证实6~8个碳原子的物质是主要的活性化合物。顺-3-己烯丁酸酯、顺-3-己烯醇、反-2-己烯醛、苯甲醛和苯甲腈的五组分混合物, 其引诱力与天然桃梢挥发物相当。通过钙成像试验证实, 尽管苯甲腈在桃梢天然挥发物中仅占0.14%, 但雌蛾对含有该物质的混合物有显著趋性, 该物质对梨小食心虫成功识别寄主具有重要意义。最后对梨小食心虫信息化学物质下一步的研究和应用前景进行了探讨。