Inter-sexual and inter-generation differences in dispersal of a bivoltine butterfly.

In organisms with discrete generations such as most insects, life-history traits including dispersal abilities often vary between generations. In particular, density-dependent differences in dispersal of bi- and multivoltine species may be expected because subsequent generations are usually characterized by a drastic increase in individual abundance. We investigated the inter-sexual and inter-generation differences in dispersal of a bivoltine butterfly, Lycaena helle, testing the following hypotheses: (1) male emigration is higher in spring generation, as males are prone to leave their natal habitat patches when the density of mating partners is low; (2) female emigration is higher in summer generation, when it helps to reduce intraspecific competition between offspring. The outcome of our analyses of dispersal parameters showed that females of the summer generation emigrated from their natal patches considerably more often than those of the spring generation, whereas an opposite trend was detected in males. These findings offer a novel perspective for our understanding of the advantages of voltinism for metapopulation functioning. The spring generation dispersal mainly improves the random mating opportunities favoured by the increase in male emigration. In turn, the dispersal of females of the summer generation appears the key to long-term metapopulation persistence.

Dispersal of Philaenus spumarius (Hemiptera: Aphrophoridae), a Vector of Xylella fastidiosa, in Olive Grove and Meadow Agroecosystems.

The introduction of the Xylella fastidiosa Wells bacterium into Apulia (South Italy) has caused the massive dieback of olive trees, and is threatening olive production throughout the Mediterranean Region. The key vector of X. fastidiosa in Europe is the spittlebug Philaenus spumarius L. The dispersal capabilities of P. spumarius are poorly known, despite being a key parameter for the prediction of the spread of the bacterium. In this study, we have examined the dispersal of P. spumarius adults in two different agroecosystems in Italy: an olive grove in Apulia (Southern Italy) and a meadow in Piedmont (Northern Italy). Insects were marked with albumin and released during seven independent trials over 2 yr. The recapture data were pooled separately for each agroecosystem and used to estimate the dispersal kernels of P. spumarius in the olive grove and in the meadow. The diffusion coefficient estimate for P. spumarius was higher in the meadow than in the olive grove. The median distance from the release point for 1 d of dispersal was 26 m in the olive grove and 35 m in the meadow. On the basis of our model, we estimated that 50% of the spittlebug population remained within 200 m (98% within 400 m) during the 2 mo period of high abundance of the vector on olives in Apulia. The dispersal of P. spumarius is thus limited to some hundreds of meters throughout the whole year, although it can be influenced to a great extent by the structure of the agroecosystem.

Sex-biased topography effects on butterfly dispersal.

BACKGROUND: Metapopulation persistence in fragmented landscapes is assured by dispersal of individuals between local populations. In this scenario the landscape topography, although usually neglected, may have an important role in shaping dispersal throughout the matrix separating habitat patches. Due to inter-sexual differences in optimal reproductive strategies, i.e., males maximizing the number of mating opportunities and females maximizing the offspring survival chances, topography-related constraints are expected to exert a different effect on male and female dispersal behaviour. We tested sex-biased topography effects on butterfly dispersal, with the following hypotheses: (1) females are constrained by topography in their movements and avoid hill crossing; (2) male dispersal is primarily driven by two-dimensional spatial structure of the habitat patches (i.e. their geometric locations and sizes) and little influenced by topography. METHODS: Following intensive mark-recapture surveys of Maculinea (= Phengaris) nausithous and M. teleius within a landscape characterised by an alternation of hills and valleys, we investigated sex-specific patterns in their inter-patch movement probabilities derived with a multi-state recapture model. In particular, we (1) analysed the fit of dispersal kernels based on Euclidean (= straight line) vs. topography-based (= through valley) distances; (2) compared movement probabilities for the pairs of patches separated or not by topographic barriers; and (3) tested the differences in the downward and upward movement probabilities within the pairs of patches. RESULTS: Euclidean distances between patches proved to be a substantially stronger predictor of inter-patch movement probabilities in males, while inter-patch distances measured along valleys performed much better for females, indicating that the latter tend to predominantly follow valleys when dispersing. In addition, there were significantly lower probabilities of movements across hills in females, but not in males. CONCLUSIONS: Both above results provide support for the hypothesis that topography restricts dispersal in females, but not in males. Since the two sexes contribute differently to metapopulation functioning, i.e., only female dispersal can result in successful (re)colonisations of vacant patches, the topography effects exerted on females should be considered with particular attention when landscape management and conservation actions are designed in order to maintain the functional connectivity of metapopulation systems.

A new comprehensive trait database of European and Maghreb butterflies, Papilionoidea.

Trait-based analyses explaining the different responses of species and communities to environmental changes are increasing in frequency. European butterflies are an indicator group that responds rapidly to environmental changes with extensive citizen science contributions to documenting changes of abundance and distribution. Species traits have been used to explain long- and short-term responses to climate, land-use and vegetation changes. Studies are often characterised by limited trait sets being used, with risks that the relative roles of different traits are not fully explored. Butterfly trait information is dispersed amongst various sources and descriptions sometimes differ between sources. We have therefore drawn together multiple information sets to provide a comprehensive trait database covering 542 taxa and 25 traits described by 217 variables and sub-states of the butterflies of Europe and Maghreb (northwest Africa) which should serve for improved trait-based ecological, conservation-related, phylogeographic and evolutionary studies of this group of insects. We provide this data in two forms; the basic data and as processed continuous and multinomial data, to enhance its potential usage.

Intersexual differences in density-dependent dispersal and their evolutionary drivers.

Dispersal is well recognized as a major driver of evolutionary processes in local populations. Nevertheless, dispersal abilities should also be perceived as a life-history trait, being subject to evolutionary changes in response to various drivers. Empirical studies investigating these drivers rarely consider that they may influence male and female dispersal differently. The purpose of our study was to document intersexual differences in density-dependent emigration from local habitat patches. As a model system, we used a metapopulation of Maculinea (Phengaris) teleius butterfly, in which densities of both sexes vary greatly throughout the flying season. Following intensive mark-release-recapture surveys, the parameters and predictors of dispersal were analysed with the Virtual Migration model and the multi-state recapture model. The emigration rate in males was substantially higher in the early season, especially at smaller habitat patches. With the proportion of females increasing with the season progression, males became reluctant to emigrate from their natal patches. In turn, higher female emigration in the later part of the season was most strongly associated with female tendency to reduce intraspecific competition experienced by their offspring. Our findings provide evidence for the impact of reproductive strategies on dispersal in both sexes. The difference in reproductive strategies of males and females explains sex-biased dispersal in different parts of the season, which carries important implications for metapopulation functioning.

Spittlebugs of Mediterranean Olive Groves: Host-Plant Exploitation throughout the Year.

Spittlebugs are the vectors of the bacterium Xylella fastidiosa Wells in Europe, the causal agent of olive dieback epidemic in Apulia, Italy. Selection and distribution of different spittlebug species on host-plants were investigated during field surveys in 2016-2018 in four olive orchards of Apulia and Liguria Regions of Italy. The nymphal population in the herbaceous cover was estimated using quadrat samplings. Adults were collected by sweeping net on three different vegetational components: herbaceous cover, olive canopy, and wild woody plants. Three species of spittlebugs were collected: Philaenus spumarius L., Neophilaenus campestris (Fallén), and Aphrophora alni (L.) (Hemiptera: Aphrophoridae). Philaenus spumarius was the predominant species both in Apulia and Liguria olive groves. Nymphal stages are highly polyphagous, selecting preferentially Asteraceae Fabaceae plant families, in particular some genera, e.g., Picris, Crepis, Sonchus, Bellis, Cichorium, and Medicago. Host-plant preference of nymphs varies according to the Region and through time and nymphal instar. In the monitored sites, adults peak on olive trees earlier in Apulia (i.e., during inflorescence emergence) than in Liguria (i.e., during flowering and beginning of fruit development). Principal alternative woody hosts are Quercus spp. and Pistacia spp. Knowledge concerning plant selection and ecological traits of spittlebugs in different Mediterranean olive production areas is needed to design effective and precise control strategies against X. fastidiosa vectors in olive groves, such as ground cover modifications to reduce populations of spittlebug vectors.

Phenology, seasonal abundance and stage-structure of spittlebug (Hemiptera: Aphrophoridae) populations in olive groves in Italy.

Spittlebugs (Hemiptera: Aphrophoridae) are the dominant xylem-sap feeders in the Mediterranean area and the only proven vectors of Xylella fastidiosa ST53, the causal agent of the olive dieback epidemic in Apulia, Italy. We have investigated the structured population phenology, abundance and seasonal movement between crops and wild plant species of both the nymphal and adult stages of different spittlebug species in olive groves. Field surveys were conducted during the 2016-2018 period in four olive orchards located in coastal and inland areas in the Apulia and Liguria regions in Italy. The nymphal population in the herbaceous cover was estimated using quadrat samplings. Adults were collected through sweep nets on three different vegetational components: herbaceous cover, olive canopy and wild woody plants. Philaenus spumarius was the most abundant species; its nymphs were collected from early March and reached a peak around mid-April, when the 4th instar was prevalent. Spittlebug adults were collected from late April until late autumn. P. spumarius adults were abundant on the herbaceous cover and olive trees in late spring, and they then dispersed to wild woody hosts during the summer and returned to the olive groves in autumn when searching for oviposition sites in the herbaceous cover. A relatively high abundance of P. spumarius was observed on olive trees during summer in the Liguria Region. The present work provides a large amount of data on the life cycle of spittlebugs within an olive agroecosystem that can be used to design effective control programmes against these vectors in infected areas and to assess the risk of the establishment and spread of X. fastidiosa to Xylella-free areas.