Asymmetric dispersal and survival indicate population sources for grassland butterflies in agricultural landscapes

We tested the hypothesis that populations in small habitat fragments remaining in agricultural landscapes are maintained by repeated immigration, using three grassland butterflies ( Aphantopus hyperantus , Coenonympha pamphilus and Maniola jurtina ). Transect counts in 12 matched sets of semi-natural pastures, and linear habitat elements proximate and isolated from the pastures showed that population densities of M.  jurtina and C.  pamphilus were significantly higher in pastures and in linear habitats adjacent to these than in isolated linear elements. A mark-recapture study in a 2×2 km landscape indicated that individuals of all three species are able to reach even the isolated linear elements situated at least 1 km from the grasslands. For two of the species, A.  hyperantus and C.  pamphilus , analysis of the mark-recapture data revealed higher daily local survival rates in the semi-natural pastures and more individuals dispersing from pastures to linear habitat elements. The proportion of old compared to young individuals of C.  pamphilus and M.  jurtina were significantly higher in linear elements than in semi-natural pastures, which suggests that butterflies emerging in pastures subsequently dispersed to the linear elements. In combination, these results suggest that semi-natural pastures act as population sources, from which adult butterflies disperse to surrounding linear elements. Hence, preservation of the remaining fragments of semi-natural grassland is necessary to keep the present butterfly abundance in the surrounding agricultural landscape.     

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.     

Egg weight variation in relation to egg mortality and starvation endurance of newly hatched larvae in some satyrid butterflies

Abstract. 1. Egg weights decrease over the oviposition period in all of the five satyrid butterflies that have been studied, viz Parage aegeria L., Lasiommata petropolitana F., L.maera L., L.megera L. and Lopingaachine Scop. However, within each species, no correlation was found between variation in egg weight and egg mortality in 30% relative humidity or the ability of newly hatched larvae to survive without food for 1–4 days. 2. Although egg weights of Papilio machaon L. were similar to the highest satyrid egg weights, newly hatched larvae of P.machaon showed significantly higher mortality after 1 day of starvation when compared to all of the satyrid larvae. This indicates that the capability of newly hatched larvae to endure starvation is not necessarily correlated with egg weight, and consequently that this character trait of satyrid larvae has been selected for (i.e. should be regarded as an adaptation). It is noteworthy that this ability of satyrid larvae to endure starvation is coupled with the habit of many satyrid butterflies not to deposit their eggs directly on the larval host plants. 3. In 30% relative humidity, egg mortality of L.achine was 100% and that of P.aegeria 29.2%, whereas that of the three Lasiommata spp. was significantly lower. At 100% relative humidity egg mortality in L.achine dropped to 8%. Since egg weights are higher in L.achine than in the other four satyrids this indicates that egg resistance to desiccation is not necessarily correlated with egg size (as might be expected due to the area to volume ratio which is negatively correlated with egg size). Thus the resistance to desiccation which is found in the three Lasiommata spp. should be regarded as an adaptation to the habit of these three butterflies to deposit their eggs in relatively dry microhabitats. This resistance to desiccation is not found in eggs of the two forest-dwelling satyrids P.aegeria and L.achine which lay their eggs in microhabitats where the relative humidity is high.     

Enhancing offspring quality or quantity? Different ways for using nectar amino acids in female butterflies

Butterfly-pollinated flowers offer nectar with higher amino acid concentrations than most flowers pollinated by other animals, and female butterflies of some species prefer to consume amino acid-rich nectar. However, for over 30 years, there has been an ongoing discussion about whether nectar amino acids benefit butterfly fitness. A clear positive effect was only shown for the nectar-feeding Araschnia levana, and females of the fruit-feeding Bicyclus anynana also increased offspring quality when they were fed amino acids as adults. Thus, severe doubts remain about the general significance of these single positive results. We therefore tested a further species from a phylogenetically different butterfly subfamily, the small heath (Coenonympha pamphilus L., Satyrinae), taking into account feeding conditions over the whole life cycle of this species. C. pamphilus females receiving nectar amino acids as adults, irrespective of larval food quality, produced heavier larvae and also increased the hatching success of their eggs over the oviposition period. Furthermore, females raised under nitrogen-poor larval conditions tended to use nectar amino acids to increase the number of eggs laid. Thus, C. pamphilus females used nectar amino acids primarily to increase their offspring quality, and secondly tended to increase offspring quantity, if larval resources were scarce, showing a resource allocation pattern differing from both B. anynana and A. levana. Our study supports the old postulate that nectar amino acids generally enhance butterfly fitness.     

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.     

Cuticular lipid diversification in Lasiommata megera and Lasiommata paramegaera: the influence of species, sex, and population (Lepidoptera: Nymphalidae)

Numerous studies have investigated the presence and the effectiveness of volatile pheromones in Lepidoptera. Conversely, very few studies have focused on the composition and the perception of the relatively low volatile components of cuticular mixtures. Yet, cuticular lipids are implied in the recognition processes of several solitary and social insects. In the present study, the cuticular signatures of the satyrid butterflies Lasiommata megera and Lasiommata paramegaera were examined by gas chromatography. General linear model and discriminant analyses on chemical data clearly revealed large differences between sexes, which showed the same diversification pattern in both species. Moreover, a strong diversification between the two species was found, as were differences among populations. These results represent a first step in demonstrating the communicative function of cuticular compounds in the L. megera / paramegaera complex. Moreover, the discrimination among different species and populations on the basis of cuticular mixtures could represent a platform for studying chemotaxonomy and chemical biogeography in butterflies, as already found in several other insect groups.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 703–710.     

Changes in egg colour, egg weight and oviposition rate with the number of eggs laid by wild females of the small heath butterfly, Coenonympha pamphilus

ABSTRACT. 1. The colour of eggs laid by individual Coenonympha pamphilus (L.) (Lep., Satyridae) females changed over their lifespan. The first laid eggs were green, but after having laid about 100 eggs, females laid only yellow eggs.
2. By following females in the field, or by capturing wild females, and noting the colour of their eggs, it was established that younger females showed higher oviposition rates and laid heavier eggs than older females.
3. Hence, wild C.pamphilus females are unable to sustain a constant egg production by extracting amino acids or any other nutrients from nectar. We hypothesize that this is a general phenomenon in nectar feeding butterflies.
4. The egg colour did not seem to match the colour of the substrate on which the female chose to deposit the egg.     

Dynamics of mate-searching behaviour in a hilltopping butterfly, Lasiommata megera (L.): the effects of weather and male density

By studying a hilltopping population of the satyrine Lasiommata megera in southern Sweden, the effects of density and meteorological conditions on mate-searching behaviour were investigated. Lasiommata megera males switched between stationary and vagrant behaviour, and the behaviour adopted was correlated to meteorological conditions; more males were stationary at low temperatures, low irradiation levels and high wind speeds than vice versa. Body temperature measurements indicated that these factors were important in deciding the body temperature and thereby the flight activity of males. Male density did not significantly influence mate-locating behaviour. The tendency of males to hilltop and their distribution around the hill were strongly influenced by weather. The hilltopping behaviour of this species did not adhere to an all-or-none pattern, but instead males, as well as the stations they used, were found at increasingly higher elevations on the hill with increasing temperatures and decreasing wind speeds. Moreover, males tended to use the lee side and sun-exposed side of the hill. Females, of which the majority were already mated, also showed this distribution around the hill, but preferred lower elevations than males. Release experiments did not reveal any propensity of mated females to fly uphill. Such a propensity was shown by released virgin females.     

The effects of island area, isolation and source population size on the presence of the grayling butterfly Hipparchia semele (L.) (Lepidoptera: Satyrinae) on British and Irish offshore islands

Records of Hipparchia semele on British and Irish islands have been modelled against island area, isolation (sea and land distance) and the size of the nearest potential source populations. All three variables have been found to contribute significantly to the presence or absence of H. semele on the islands. Isolation is a more significant predictor than island area. This result differs from the multiple species case where area was found to be a more important influence than isolation. Records on islands are also shown to depend on the size of populations at the nearest sources; this underpins the relationships identified for the multiple species case, first, between the number of species on islands and at nearest sources and, second, between the incidence of species on islands and at nearest sources. There are clear indications that smaller islands may become increasingly marginalized for H.semele; with ongoing habitat loss, because isolation increases and source populations become sparser, the probability of H. semele recolonizing islands also decreases.     

Individual state controls temperature dependence in a butterfly (Lasiommata maera)

In ectotherms there is typically a strong and positive correlation between growth rate and ambient temperature when food is not limiting. However, the exact relationship between growth rate and temperature varies among populations in many species. As a consequence, it has been suggested that selection for a rapid increase in growth rate with temperature should be stronger in populations experiencing a high degree of time-stress, compared with populations experiencing little time-stress. In the present study we take this adaptive hypothesis further and investigate if variation in time-stress among individuals of a single population may affect the relationship between growth rate and ambient temperature. Time-stress was manipulated by rearing larvae of the butterfly Lasiommata maera in different day-length regimes. The results show that individuals experiencing a higher degree of time-stress increase their growth rates more in higher temperatures compared with individuals under less time-stress. Hence, the adaptive hypothesis was supported and the relationship between growth rate and temperature was highly state dependent. These findings may be of general importance for understanding the evolution of life histories in seasonal environments.     

Parasitism by the mite Trombidium breei on four U.K. butterfly species

Abstract 1.  The incidence of parasitism by larvae of the mite species Trombidium breei was reported in one population of the lycaenid butterfly Polyommatus icarus , four populations of the satyrine butterfly Maniola jurtina , one population of the satyrine butterfly Aphantopus hyperanthus , and two populations of the satyrine butterfly Pyronia tithonus , as well as on one specimen of the dipteran Alophorus hemiptera . A considerable proportion of butterflies (11-50%) was infested in all study populations.
2. The pattern of infestation was examined in detail in M. jurtina . Males had a significantly higher incidence of infestation than females, and middle-aged butterflies had a higher incidence of infestation than old or young butterflies. The incidence of infestation peaked in the middle of the flight season, and this seasonal effect was independent of the effect of butterfly age.
3. Using a model based on capture-recapture data, it was estimated that a hypothetical ideal male M. jurtina that lives exactly the mean expected lifespan of 9-10 days has an approximately 75% chance of becoming infested with mites at least once during its lifetime, a mean time to first infestation of 3-4 days, and an average infestation persistence time of 2-3 days.
4. Capture-recapture data failed to show any effect of mite infestation on the lifespan or within-habitat movement rate of M. jurtina .
5. In experiments in which individual butterflies were taken out of their normal habitat and released, M. jurtina and P. tithonus that were infested with mite larvae did not differ from uninfested individuals in the efficiency with which they returned to suitable habitat. Thus, parasitism by T. breei larvae had no detectable effects on flight performance or orientation ability.
6. The results suggest that trombidiid mite larvae have limited potential in the biological control of insect pests.     

Life history plasticity in the satyrine butterfly Lasiommata petropolitana: investigating an adaptive reaction norm

This study addresses the general hypothesis that insects living in seasonal environments should shorten development times at progressively later dates in the growth season, and that insects living outside equatorial areas should use daylength as a cue to determine the date. Diapause strategies and reaction norms relating the duration of larval development to daylength was investigated in a French population of the butterfly, Lasiommata petropolitana. The results are compared with those of an earlier study of the species in Sweden. Because of the diapausing strategy and phenology of the population, it was expected that an adaptive reaction norm relating larval time to daylength should have a positive slope, i.e. relatively shorter daylengths induce faster growth and development. This prediction was supported, and the reaction norm was qualitatively similar to the one found in Swedish populations. In the French population it was, however, shifted to a range of shorter photoperiods which corresponds to the regime of shorter daylengths in southern Europe. Shorter larval development times and high growth rates were associated with a reduction in pupal size, suggesting a trade off between time and size at pupation. There was no evidence of a trade off between growth rate and starvation endurance. The results suggests that the daylength-dependent decision of what growth trajectory an individual larva will follow, is not made continuously but rather at one or a few occasions during larval development. It is clear that larvae of L. petropolitana make developmental decisions in relation to the daylength they experience during larval growth. The result is a reaction norm that agrees closely to what is predicted by some life history models, suggesting that it is an adaptation for optimising life history traits in a seasonal environment.     

Immature stages of Hipparchia Fabricius and the systematics of the 'Satyrus series>> (Lepidoptera: Nymphalidae: Satyrinae)

Abstract. Characters of potential use from the eggs and larvae of four species of Hipparchia (subgenera Hipparchia, Parahipparchia, Neohipparchia and Pseudotergumia ) are described. These characters, together with previously published information on larval, pupal and adult morphology, are used to construct a seventy-one character matrix for Hipparchia and four other genera of the 'Satyrus series' (Miller, 1968): Arethusana, Kanetisa, Chazara and Satyrus . The cladistic relationships among these genera and the relative contribution of adult and immature data sets are discussed. A list of possible synapomorphies of the immature stages of the 'Satyrus series' is proposed.     

Larvae of lycaenid butterflies that parasitize ant colonies provide exceptions to normal insect growth rules

Fourth instar larvae of Maculinea species of lycaenid butterfly live as social parasites inside Myrmica ant nests. They show highly unusual growth patterns, with small but regular growth in early phytophagous instars, followed by >10 times the growth predicted by extrapolating the early growth rate (following Dyar's rule) during the final carnivorous instar. This produces striking allometry between head and body size in full-grown larvae (ratios of 4–5% compared with 8–10%). Larvae of the Myrmica ant hosts have a similar growth. Data for c. 150 other lycaenid species showed that species with similar life-histories exhibit the same unusual growth pattern (Phengaris spp., Lepidochrysops spp., Niphanda fused); all others have regular growth throughout their larval life, including the carnivorous species that are parasitic on ants from the first instar. It is suggested that Maculinea-type growth pattern has arisen convergently in at least three unrelated lineages of lycaenids. Selection pressures might include the need for reduced early growth to produce late instars that are small enough to be integrated as brood mimics into ant social systems, combined with the need to achieve at least the same adult size as the ancestral species. Trophic pressures that operate on both sedentary ant and butterfly larvae, which must survive long periods of starvation and grow rapidly when food is abundant, may also be involved.     

Geographical variability in, and temperature effects on, the phenology of Maniola jurtina and Pyronia tithonus (Lepidoptera, Satyrinae) in England and Wales

ABSTRACT. 1. Geographical variability in, and temperature effects on, the mean date of adult flight period and the SD about this date are analysed for two univoltine, grassland butterflies in England and Wales from 1976 to 1985. Data were collected on the Butterfly Monitoring Scheme for Maniola jurtina (L.) at twenty-nine sites and Pyronia tithonus (L.) at twenty sites.
2. Substantial variability for mean date and SD occurs between years and between sites. Changes in mean date between years tend to occur consistently at different sites. The species show some parallel in variation between sites, especially for mean date.
3. June maximum temperature accounts for 95% and 75% of the variation in mean date between years in M. jurtina and P. tithonus , respectively ( r =-0.97 and -0.87). Similar relationships occur for temperatures cumulated over the period of post-winter development from March to July or August.
4. Greater geographical variability in phenology, and a generally less synchronized flight period in M.jurtina may be associated with broader habitat preferences than in P. tithonus.
5. The mean date of adult flight period remains at roughly the same date at more northerly latitudes. In M.jurtina the flight period becomes more synchronized, begins later and ends earlier in the north. P. tithonus shows little or no indication of such a response to latitude. This is discussed with regard to changes in season length and factors limiting the species'range.     

The role of the North Atlantic Oscillation in controlling U.K. butterfly population size and phenology

1. The North Atlantic Oscillation (NAO) exerts considerable control on U.K. weather. This study investigates the impact of the NAO on butterfly abundance and phenology using 34 years of data from the U.K. Butterfly Monitoring Scheme (UKBMS).2. The study uses a multi-species indicator to show that the NAO does not affect overall U.K. butterfly population size. However, the abundance of bivoltine butterfly species, which have longer flight seasons, were found to be more likely to respond positively to the NAO compared with univoltine species, which show little or a negative response.3. A positive winter NAO index is associated with warmer weather and earlier flight dates for Anthocharis cardamines (Lepidoptera: Pieridae), Melanargia galathea (Lepidoptera: Nymphalidae), Aphantopus hyperantus (Lepidoptera: Nymphalidae), Pyronia tithonus (Lepidoptera: Nymphalidae), Lasiommata megera (Lepidoptera: Nymphalidae) and Polyommatus icarus (Lepidoptera: Lycaenidae). In bivoltine species, the NAO affects the phenology of the first generation, the timing of which indirectly controls the timing of the second generation.4. The NAO influences the timing of U.K. butterfly flight seasons more strongly than it influences population size.     

Test for latitudinal variation of life history,behavior and mortality in the strictly univoltine damselfly Sympecma fusca (Zygoptera: Lestidae)

A wealth of evidence shows that combinations of ecological stressors interact in shaping life history traits, but little is known about how ecological stressors combine with different seasonal time constraints to shape life history, behavior and mortality across populations. We studied life history, behavior and mortality rate in two latitudinally distant populations of the strictly univoltine, adult‐overwintering damselfly Sympecma fusca. Results from laboratory common‐garden and outdoor experiments indicated countergradient variation of larval development time and growth rate: the more time‐constrained larvae showed faster development and a higher growth rate. This finding led to larger size at emergence in the more time‐constrained individuals. Under conditions of intraspecific interaction (outdoor experiment), northern individuals showed lower survival than southern ones, presumably due to cannibalism. In the absence of intraspecific interactions (laboratory experiment), northern and southern larvae did not differ in survival. Finally, laboratory‐grown northern and southern larvae did not differ in activity level. This is the first time that compensation for seasonal time constraints has been shown in a temperate odonate species that overwinters in the adult stage.     

Diet and growth of a leaf-shredding caddisfly in southern Appalachian streams of contrasting disturbance history

Diet and growth of leaf-shredding caddisfly larvae, Pycnopsyche spp.,were examined in streams draining a reference catchment and a 16-year-oldclear-cut (disturbed) catchment at Coweeta Hydrologic Laboratory insouthwestern North Carolina, USA. The objective was to explain why shredderproduction is higher in the disturbed streams despite the larvae having lessfood (i.e., leaves) available. We predicted larvae would grow faster onfast-decaying leaf material representative of the disturbed streams. Larvaeconsumed mostly leaf detritus in three streams draining each catchment overthree seasons (fall, winter, and spring), which showed larvae did notconsume higher quality foods (e.g., algae and animal material) in disturbedstreams. When fed 2-month-old conditioned black birch (Betula lenta L.) (afast-decaying leaf species) and white oak (Quercus alba L.) (a slow-decayingleaf species) leaves in the laboratory, larvae grew significantly faster onthe birch leaves. However, when larvae were fed the same leaf types after3-months conditioning, larvae grew significantly faster on oak leaves. Afield growth experiment conducted for 42 d using mixed-species leaf dietsrepresentative of each catchment and initially conditioned for 2 monthsfound that Pycnopsyche grew significantly better on the diet representativeof the reference catchment. The reference diet contained more oak leaveswhich apparently became a more acceptable food as the experiment proceeded.High shredder production in the disturbed streams could not be explained byhigh Pycnopsyche growth rates on fast-decaying leaves. Instead, larvae grewbetter on leaves that were apparently conditioned optimally regardless ofconditioning rate.     

Morphology and chaetotaxy of the first instar larvae of six species of the Satyrus (s.l.) series (Lepidoptera: Nymphalidae)

Abstract. The morphology and chaetotaxy of the first instar larvae of six species belonging to the genera Hipparchia, Kanetisa and Chazara are described. Specific characters are stated, drawn mainly from size, setal length and morphology, and the shape of the suranal plate. Several characters, other than chaetotaxy, that are of potential use in nymphalid systematics are discussed. The larval chaetotaxy is briefly compared with that of both heliconiine and danaine first instar larvae.     

Life history plasticity to combined time and biotic constraints in Lestes damselflies from vernal and temporary ponds

Little is known about the ecological correlates and macro-evolution of life history plasticity to time constraints. Here, we compared age and mass at emergence and their plasticity toward combinations of time constraints (delayed larval development), food level and predation risk (caged dragonfly larvae) by rearing two temporary-pond Lestes damselflies ( L. congener and L. forcipatus ) and the derived vernal-pond L. dryas from the eggs until adult emergence in outdoor tubs. Life history plasticity under time constraints, low food and predation risk was as predicted by optimality models. Delayed larvae in all three species accelerated development and showed a lower fat content and a substantially elevated mortality rate. At low food, all species emerged later at a smaller mass. Also under predation risk adults of all species emerged later, and (at high food) at a smaller mass. Unexpectedly, delayed larvae did not show a smaller life history response to predation risk imposed by dragonfly larvae. Compared to the two temporary-pond Lestes studied, the derived species that invaded more ephemeral vernal ponds showed a faster development rate and a lower deceleration of development to low food. It also showed a lower acceleration of development to time constraints, possibly reflecting that it reached development rates near to its physiological maximum. Unexpectedly, the vernal-pond Lestes did not slow its development less under predation risk. Our results stress the importance of evaluating ecological and evolutionary correlates of life history plasticity under as realistic conditions as possible.