Pararge Aegeria Research Articles

Flight during oviposition reduces maternal egg provisioning and influences offspring development inPararge aegeria(L.)

As a result of increased habitat fragmentation in anthropogenic landscapes, flying insects may be required to travel over larger distances in search of resources such as suitable host plants for oviposition. The oogenesis-flight syndrome hypothesis predicts that physiological constraints caused by an overlap in the resources used by thoracic muscles during flight and during oogenesis (e.g. carbohydrates, lipids and water) result in a resource trade-off, with any resources used during flight no longer available for reproduction. Increased flight costs could therefore potentially result in a decrease in maternal provisioning of eggs. In the present study, the speckled wood butterfly Pararge aegeria (L.) is used to investigate whether increased flight during oviposition results in changes in maternal investment in eggs and whether this contributes to variation in the development of offspring in subsequent life stages. Forcing females to fly during oviposition directly influences egg size and embryonic development time, and indirectly influences (through changes in egg size) egg hatching success and larval development time. These effects are mediated through 'selfish maternal effects', with mothers forced to fly maximizing their fecundity at the expense of investment to individual egg size. The present study demonstrates that a change in maternal provisioning as a result of increased flight during oviposition has the potential to exert nongenetic cross-generational fitness effects in P. aegeria. This could have important consequences for population dynamics, particularly in fragmented anthropogenic landscapes.

Visual mate detection and mate flight pursuit in relation to sunspot size in a woodland territorial butterfly

Territory residency is associated with considerable benefits. In the speckled wood butterfly, Pararge aegeria , males fight over ownership of large sunspots in open forest habitats ; winners become sunspot residents, and losers become nonterritorial and sit and wait for females in small sunspots. A previous study has shown that residents have higher mating success than nonterritorial males, although females are not more attracted to territorial males or sunspot territories per se. Here we tested the hypotheses (1) that the higher success of resident males is caused by visual mate detection being more efficient in a large than in a small sunspot, and (2) that only sunspots above a certain size are defended as territories. Field assessment of territorial sunspot size showed that defended sunspots were significantly larger than ‘average sunspots’ on the forest floor. Experimental tests of male ability to detect visually a model butterfly passing through a sunspot showed that males were more successful in pursuing and intercepting a passing model when flown a longer distance in the sunspot. Hence, we conclude that light conditions and associated visual mate detection and ability to complete mate flight pursuit can explain why P. aegeria males defend territories in large sunspots in forest habitats.

Larval food stress differentially affects flight morphology in male and female speckled woods (Pararge aegeria)

Abstract 1. Adaptive plasticity in flight morphology can be of great importance for organisms, in order to deal with changing environments. When different demands are imposed to this morphology, different responses to environmental changes can be expected.2. The aim of this study is to examine whether males and females of Pararge aegeria, which show different flight behaviours, respond differently to larval food stress.3. In a food‐stress experiment, larvae of 35 families were reared on host plants subjected to a drought‐stress treatment with three groups: a control group, a low‐stress group and a high‐stress group.4. Individuals from stress treatments significantly differed in wing morphology; they had lower wing loadings, and stressed females tended to have more pointed wings than females of the control group.5. The difference in phenotypic response to food stress between both sexes may indicate that males and females benefit from different changes in morphology. In females, an increase in dispersal capacity may entail fitness benefits, whereas male morphology is mainly shaped by mate‐location strategy.

TAKE-OFF FLIGHT PERFORMANCE IN THE BUTTERFLYPARARGE AEGERIARELATIVE TO SEX AND MORPHOLOGY: A QUANTITATIVE GENETIC ASSESSMENT

Adult fitness components may strongly depend on variation in locomotory performance such as flight; this variation can be sex specific. Fast take-off to intercept females and competing males is an essential behavioral component of the territorial perching behavior in male speckled wood butterflies (Pararge aegeria L.). Females on the other hand avoid frequent take-offs particularly under suboptimal temperatures, typically showing fewer but longer flights than males. We estimated the heritability of take-off acceleration performance under suboptimal body temperatures by a restricted maximum-likelihood model. We calculated genetic correlations between this performance and a selection of morphological traits: size (body mass), flight muscle investment (relative thorax mass), and wing shape (forewing aspect ratio). Our results show significant additive genetic variation for mean acceleration performance and a similar but nonsignificant trend (P= 0.08) for maximal acceleration performance during take-off in males (h(2)= 0.15). In females, heritability was not significantly different from zero for either of the acceleration performance measures. Morphological traits and take-off performance were genetically linked in a sex-specific way. In males, relative thorax mass and forewing aspect ratio were positively genetically correlated with acceleration performance. In females, there was a negative genetic correlation between acceleration performance and abdomen mass, but not with residual abdomen mass (i.e., regressed on total body mass). To fully understand the evolution of sexual differences in flight performances and morphology, several other flight performances will have to be included. This multifunctional nature of flight and its consequences for the evolutionary study of flight has not yet been fully appreciated in the literature.

Environmental stress and quantitative genetic variation in butterfly wing characteristics

Butterfly wing characteristics are extensively used as model system in studies of development, quantitative genetics and phenotypic plasticity. In spite of its evolutionary relevance, however, the effect of stress on the expression of genetic variation itself has only rarely been studied. In this paper, we explore genetic variation of wing characteristics of the Speckled wood Pararge aegeria along a host plant drought stress gradient. Forewing area, basal and distal degree of melanization and the area of five yellow wing spots were measured. We found an increase in (additive) genetic variation in degree of melanization at higher drought stress, and a similar, yet non-significant, relationship for forewing size. As a result, both the upper limits of the narrow-sense heritability and the coefficient of additive genetic variation of wing size and melanization increased with drought stress. Patterns for the different yellow wing spots were less consistent, suggesting trait-specificity in the relationships between genetic variation and environmental stress.

Maternal body size as a morphological constraint on egg size and fecundity in butterflies

Summary It is a widespread notion that in arthropods female reproductive output is strongly affected by female size. In butterflies egg size scales positively with female size across species, suggesting a constraint imposed by maternal size. However, in intraspecific comparisons body size often explains only a minor part of the variation in progeny size. We here include representatives of various butterfly families to test the generality of this phenomenon across butterflies. Phenotypic correlations between egg and maternal body size were inconsistent across species: correlations were non-significant for Pararge aegeria and Lycaena tityrus , significantly positive for Papilio machaon , significantly negative for Araschnia levana , and contradictory for Pieris napi . Thus, there was no general pattern linking egg size to maternal size, e.g., caused by an allometric relationship. Consequently, there was at best limited evidence for maternal size acting as a morphological constraint on egg size within butterfly species. Realized fecundity depended on maternal size in P. napi and A. levana , but not in P. aegeria , suggesting that maternal size may affect egg number more strongly than egg size. Yet, variation in fecundity was primarily explained by variation in longevity as is expected for income breeders. Heritability estimates across species were rather similar for pupal mass (ranging between 0.14 and 0.19), but more variable for egg size (0.17–0.31).

Phenotypic and genetic variations and correlations in multitrait developmental instability: a multivariate Bayesian model applied to Speckled Wood butterfly (Pararge aegeria) wing measurements

Phenotypic and genetic variation and covariation in developmental instability (DI) have been the subject of many debates. In this paper we develop and apply a statistical model in a Bayesian context to analyse different traits simultaneously in a multivariate model of DI. We apply the model to measurements of yellow spots on the front wing of the Speckled Wood butterfly (Pararge aegeria L.) in a full-sib breeding experiment. We estimated the posterior distribution of the broad-sense heritability of DI averaged across the five yellow spots, which had a median of 0.19 and a 95% credibility interval ranging between 0.04 and 0.64. Phenotypic and genetic correlations in DI could not be estimated accurately with the present sample size. Yellow spots 4 and 5 appeared to show some degree of developmental integration. The importance of this model and its possible extensions are discussed.

Integration of wings and their eyespots in the speckled wood butterfly Pararge aegeria

We investigated both the phenotypic and developmental integration of eyespots on the fore- and hindwings of speckled wood butterflies Pararge aegeria. Eyespots develop within a framework of wing veins, which may not only separate eyespots developmentally, but may at the same time also integrate them by virtue of being both signalling sources and barriers during eyespot development. We therefore specifically investigated the interaction between wing venation patterns and eyespot integration. Phenotypic covariation among eyespots was very high, but only eyespots in neighbouring wing cells and in homologous wing cells on different wing surfaces were developmentally integrated. This can be explained by the fact that the wing cells of these eyespots share one or more wing veins. The wing venation patterns of fore- and hindwings were highly integrated, both phenotypically and developmentally. This did not affect overall developmental integration of the eyespots. The adaptive significance of integration patterns is discussed and more specifically we stress the need to conduct studies on phenotypic plasticity of integration.

Contributions to the Knowledge of Papilionoidea (Lepidoptera: Rhopalocera) Fauna from Northern Cyprus

The local distribution of the superfamily Papilionoidea was investigated in Northern Cyprus. As a result of this study, a total of 45 species belonging to families Papilionidae, Pieridae, Lycaenidae, Satyridae and Nymphalidae was identified; out of which 19 species are new records for the study area. The Nymphalidae had the greatest number of species following by Pieridae, Satyridae, Lycaenidae and Papilionidae. The most abundant species in the study area were Pieris brassicae (L.) (19% of the catches), Polyommatus icarus (Rott.) (11%), Euchloe ausonia (Hbn.) (5%) and Pararge aegeria (L.) (5%).

Mating success of resident versus non-resident males in a territorial butterfly

Male–male competition over territorial ownership suggests that winning is associated with considerable benefits. In the speckled wood butterfly, Pararge aegeria, males fight over sunspot territories on the forest floor; winners gain sole residency of a sunspot, whereas losers patrol the forest in search of females. It is currently not known whether residents experience greater mating success than non-residents, or whether mating success is contingent on environmental conditions. Here we performed an experiment in which virgin females of P. aegeria were allowed to choose between a resident and a non-resident male in a large enclosure containing one territorial sunspot. Resident males achieved approximately twice as many matings as non-residents, primarily because matings were most often preceded by a female being discovered when flying through a sunspot. There was no evidence that territorial residents were more attractive per se, with females seen to reject them as often as non-residents. Furthermore, in the cases where females were discovered outside of the sunspot, they were just as likely to mate with non-residents as residents. We hypothesize that the proximate advantage of territory ownership is that light conditions in a large sunspot greatly increase the male's ability to detect and intercept passing receptive females.

Thermal flight budget of a woodland butterfly in woodland versus agricultural landscapes: An experimental assessment

We assessed the impact of vegetation structure and type of landscape on microclimatic conditions for flight activity, using the heliothermic butterfly Pararge aegeria as a model. Dummies (i.e., dead specimens with spread wings, on a hypodermic needle with a thermocouple) that initially were warmed up to similar thorax temperatures were able to cover larger distances, and they heated up faster, with higher ambient temperature and with lower wind speed. Convective cooling decreased with increasing degree of shelter, and with lower flight height. Simulated flight bouts bridged 140±23 m under unsheltered, 81±13 m under partly sheltered and 29±6 m under sheltered conditions. Ambient temperature and wind force were higher in highly fragmented, agricultural landscapes than in continuous woodland landscapes. Males, and not females, were furrier in agricultural landscapes. This sexual difference is interpreted in terms of differences in life history among the sexes.

What Keeps Insects Small? Time Limitation during Oviposition Reduces the Fecundity Benefit of Female Size in a Butterfly

Laboratory studies of insects suggest that female fecundity may increase very rapidly with adult size and that mass may often increase close to exponentially with time during larval growth. These relationships make it difficult to see how realistic levels of larval mortality can outweigh the fecundity benefit of prolonged growth. Hence, it is unclear why many insects do not become bigger. In this study, we experimentally explore the relationship between female size and fecundity in the butterfly Pararge aegeria and show that thermally induced time limitation during oviposition may substantially reduce the fecundity benefit of larger females. We model time-limited oviposition under natural temperature conditions and show that fecundity is also likely to increase asymptotically with female size in the field. With realistic estimates of juvenile mortality, the model predicts optimal body sizes within the observed range even when larvae grow exponentially. We conclude that one important reason for why insects with a high capacity of larval growth do not evolve toward larger sizes may be that the fecundity benefit is in fact relatively limited under natural conditions. If so, these results may help resolve some of the inconsistencies between theory and empirical patterns in explaining optimal size in insects.

What Keeps Insects Small? Time Limitation during Oviposition Reduces the Fecundity Benefit of Female Size in a Butterfly

Laboratory studies of insects suggest that female fecundity may increase very rapidly with adult size and that mass may often increase close to exponentially with time during larval growth. These relationships make it difficult to see how realistic levels of larval mortality can outweigh the fecundity benefit of prolonged growth. Hence, it is unclear why many insects do not become bigger. In this study, we experimentally explore the relationship between female size and fecundity in the butterfly Pararge aegeria and show that thermally induced time limitation during oviposition may substantially reduce the fecundity benefit of larger females. We model time‐limited oviposition under natural temperature conditions and show that fecundity is also likely to increase asymptotically with female size in the field. With realistic estimates of juvenile mortality, the model predicts optimal body sizes within the observed range even when larvae grow exponentially. We conclude that one important reason for why insects with a high capacity of larval growth do not evolve toward larger sizes may be that the fecundity benefit is in fact relatively limited under natural conditions. If so, these results may help resolve some of the inconsistencies between theory and empirical patterns in explaining optimal size in insects.

On the sex-specific mechanisms of butterfly flight: flight performance relative to flight morphology, wing kinematics, and sex in Pararge aegeria

Many evolutionary ecological studies have documented sexual dimorphism in morphology or behaviour. However, to what extent a sex-specific morphology is used differently to realize a certain level of behavioural performance is only rarely tested. We experimentally quantified flight performance and wing kinematics (wing beat frequency and wing stroke amplitude) and flight morphology (thorax mass, body mass, forewing aspect ratio, and distance to centre of forewing area) in the butterfly Pararge aegeria (L.) using a tethered tarsal reflex induced flight set-up under laboratory conditions. On average, females showed higher flight performance than males, but frequency and amplitude did not differ. In both sexes, higher flight performance was partly determined by wing beat frequency but not by wing stroke amplitude. Dry body mass, thorax mass, and distance to centre of forewing area were negatively related to wing beat frequency. The relationship between aspect ratio and wing stroke amplitude was sex-specific: females with narrower wings produced higher amplitude whereas males show the opposite pattern. The results are discussed in relation to sexual differences in flight behaviour. (c) 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89, 675-687.

Occurrence pattern of Pararge aegeria (Lepidoptera: Nymphalidae) with respect to local habitat suitability, climate and landscape structure

Distribution patterns of wild species are affected by environmental variables, such as climate, anthropogenic land use or habitat quality, which act simultaneously at different scales. To examine the relative importance of particular factors and scales on population response we investigated the speckled wood butterfly Pararge aegeria (L.) as a model organism occupying semi-natural habitats. Its distribution was recorded in 23 study sites (5×5 km) over a 2 year study period. The sites were located in agricultural landscapes within seven Temperate European countries. Environmental predictors were mapped at a local and a regional scale. Logistic regression models were then developed to represent humid (beneficial) and dry (adverse) weather conditions during larval development. The humid year model predicted that P. aegeria is equally but generally not very dependent on local and regional factors, resulting in generally high occurrence probabilities. In contrast, the dry year model predicted severe restrictions of P. aegeria to both high quality patches and landscapes with beneficial structural and climatic preconditions. As both models resulted in entirely different predictions, our study showed that the sensitivity of P. aegeria to local and landscape features might change, and that factors of less importance could easily become limiting factors. The results stress that high quality landscape is important at both the local and regional scale even for species that are considered relatively robust. They also sound a note of caution when predictions about population response for management purposes are based on just a single or a few year(s) of observation.

Sex‐ and landscape‐related differences in flight ability under suboptimal temperatures in a woodland butterfly

1 Active time budgets of flying ectotherms depend on body temperature. Knowledge of flight ability and performance is currently heavily biased towards the range of temperatures favourable for voluntary flight activity. Flight under suboptimal temperature conditions may also be important, however, in terms of fitness. 2 Here we study the ability of organisms to fly at body temperatures colder than those at which flight is initiated voluntarily, and the duration of such flights. Laboratory-reared Pararge aegeria (L.) butterflies that originated from woodland and agricultural landscape were tested at five temperatures (range: 10–21 °C) in a common-garden set-up. 3 We predict that males are able to fly at lower temperatures than females as males have lower wing loading (i.e. body mass/wing area). Since woodland is on average cooler than agricultural landscape we also predict that flight ability at low temperature is better developed in woodland individuals. 4 Individuals showed an ability to fly at all tested temperatures and flights were longer with increasing temperature. Males flew for longer than females. There was no difference in flight duration between sexes at the lowest temperature, but an increasing difference with increasing temperature. We showed that woodland butterflies flew for longer and had higher wing loading than agricultural landscape butterflies. 5 Our results shed new light on the thermal ecology of flight at suboptimal temperatures and are discussed from both proximate and ultimate points of view.

Landscape structure and phenotypic plasticity in flight morphology in the butterflyPararge aegeria

In evolutionary time, varying environments may lead to different morphs as a result of genetic adaptation and divergence or phenotypic plasticity. Landscapes that differ in the extent of habitat fragmentation may provide different selection regimes for dispersal, but also for other ecological functions. Several studies on flying insects have shown differences in flight morphology between landscapes, but whether such differences result from plastic responses have rarely been tested. We did a reciprocal transplant experiment with offspring of speckled wood butterfly females (Parargeaegeria) from three types of landscape differing in fragmentation: woodland landscape, landscape with woodland fragments and agricultural landscape with only hedgerows. Young caterpillars were allowed to grow individually on potted host grasses in small enclosures under the three landscape conditions (split‐brood design). Mortality in caterpillars was much higher in agricultural landscape compared to the other landscapes. Additive to the effect of landscape of development, landscape of origin also affected mortality rate in a similar way. Flight morphology of the adults resulting from the experiment differed significantly with landscape. Independent of the landscape of origin, males and females that developed in agricultural landscape were the heaviest and had the greatest wing loadings. Females that developed in agricultural landscape had higher relative thorax mass (i.e. greater flight muscle allocation) in line with adaptive predictions on altered dispersal behaviour with type of landscape. In males, relative thorax mass did not respond significantly relative to landscape of development, but males originating from landscape with woodland fragments allocated more into their thorax compared to males from the other types. We found significant G×E interactions for total dry mass and wing loading. Our results suggest the existence of phenotypic plasticity in butterfly flight morphology associated with landscape structure.

Life History Effects Upon Contest Behaviour: Age as a Predictor of Territorial Contest Dynamics in Two Populations of the Speckled Wood Butterfly, Pararge aegeria L.

AbstractAlthough empirical studies of life history effects upon sexually selected phenomena have largely overlooked contest behaviour, recent research suggests that territorial contest participation in butterflies may be mediated by ageing per se. Verbal and mathematical arguments predict lifetime increases in the expression of risky male reproductive behaviours, such as fighting, under a range of ecological conditions. Here we explored the relevance of ageing per se to contest dynamics in two phenologically distinct populations of the speckled wood butterfly, Pararge aegeria. We established 160 experimental like‐population contests among naïve south Swedish and Madeiran dyads, 112 of which we varied the age difference between combatants by 3–4 d. Although this age asymmetry did not influence contest outcome in either population, we found weak positive covariance between the loser's age and contest duration amongst Madeiran males. This is consistent with a slight lifetime increase in aggression because the duration of these aerial persistence contests is a sensitive measure of the losing male's level of aggressive intent. However, the size of this effect (semi‐partial correlation = 0.281) suggests age is not as strongly relevant to contest behaviour in P. aegeria as in other territorial butterflies. We discuss the ways in which ecological differences between butterfly species, particularly with respect to predation risk, may have influenced the evolution of lifetime aggressive strategies in this group.

The cost of mating with a non-virgin male in a monandrous butterfly: experimental evidence from the speckled wood, Pararge aegeria

Contrary to vertebrates, sperm production in insects may bear considerable costs for males. This is especially true in species that donate spermatophores containing sperm and nutrient-rich accessory gland products like in butterflies. Hence, spermatophores at first and subsequent copulations can differ in a quantitative and qualitative way. Such effects have particularly been shown in polyandrous species providing large spermatophores. Here we experimentally tested the effect of male mating status (virgin male vs recently mated male) on copulation duration, spermatophore size and females' fitness components in a monandrous butterfly Pararge aegeria that typically donates small spermatophores. Copulations with non-virgin males lasted on average five times longer than that with virgin males and resulted in a spermatophore which was on average three times smaller. Number of eggs laid and female life span were not affected by the mating status treatment, but there was a significant effect on the number of living caterpillars a female produced, as copulations with virgin males resulted in higher numbers of larval offspring. Interestingly, the difference in spermatophore mass at the first and the second copulation increased with male body size. This suggests differential spermatophore allocation decisions among males of different size. Consequences for females and potential mechanisms influencing female fitness components are discussed. Given the small absolute size of spermatophores in P. aegeria, components other than consumable nutrients (perhaps hormones) should cause the observed effects.

Multiple host-plant use may arise from gender-specific fitness effects

Ovipositing females are predicted to select host-plants that will maximise offspring survival and fitness. Yet hosts often differ in the component of larval fitness affected so host-selection often involves a trade-off between short development times and large size and high fecundity of offspring. If host-species can directly affect development rates and body size, and if there are gender differences in resource allocation during development, there can be different sex-specific selection pressures associated with different hosts. Using a Madeiran population of the speckled wood butterfly Pararge aegeria (L.) as the model species gender differences in larval development and size were examined in response to the hosts Brachypodium sylvaticum, Holcus lanatus and Poa annua. It was observed that male and female P. aegeria larvae differed, with their responses dependent on the host species. These results would suggest that oviposition behavior is a complex process, and use of multiple hosts may have evolved to balance the conflicting needs of male and female larvae. Co-evolution of host selection and oviposition behaviors may help to balance the differing performance needs of offspring.