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1. Climate change is causing shifts in the distribution of many species and populations inhabiting mountain tops are particularly vulnerable to these threats because they are constrained in altitudinal shifts. Apennines are a relatively narrow and low mountain chain located in Southern Europe, which hosts many isolated populations of mountain species. The butterfly Erebia pandrose was recorded for the last time in the Apennines in 1977, on the top of a single massif (Monti della Laga). 2. We confirmed the presence of a small, isolated population of E. pandrose in the Apennines, at a distance of more than 400¿km to any other known populations. Then, we examined the cytochrome c oxidase subunit 1 mitochondrial DNA marker of this species across the Palaearctic area and estimated the potential decline over the Alps and the Apennines due to future climatic changes. 3. The Apennine population represents an endemic lineage characterised by eight mutations over the 658¿bp analysed (1.2%). In the Alps and Apennines, this species has shifted uphill more than 3 m per year since the end of the 19th century and more than 22 m per year since 1995. Species distribution models suggested that these mountain populations will experience a generalised loss of climatic suitability, which, according to our projections, could lead to the extinction of the Apennine population in a few decades. 4. Erebia pandrose has the potential to become a flagship species for advertising the risk of losing unique fractions of genetic diversity for mountain species., funded by the project: ‘Ricerca e conservazione sui lepidotteri diurni di sei Parchi Nazionali dell'Appennino centro-settentrionale’. Support for this research was also provided by ‘La Caixa’ Foundation (ID 100010434) to M.M. (grant LCF/BQ/DR20/11790020), by the Academy of Finland to V.D. (Academy Research Fellow, decision no. 328895) and by project PID2019-107078GB-I00/AEI/10.13039/501100011033 to R.V., 1 Introduction 2 Materials and methods 2.1 Data collection 2.2 COI analyses 2.3 Species distribution modelling 2.4 Elevational shifts of E. pandrose records through time 3 Results 3.1 COI structure 3.2 Climate change effects 4 Discussion 4.1 The population of Monte Gorzano 4.2 What is the future for E. pandrose on the Apennines? 4.3 Erebia pandrose in the context of preserving Apennine butterflies 5 Conclusions Acknowledgements Conflict of interest

Here we report the strong decline of a population of the endangered species Parnassius mnemosyne (Linnaeus, 1758) (Lepidoptera: Papilionidae) in the National Park of Foreste Casentinesi, Monte Falterona and Campigna (Italy). We compared historical presence data (before 1969) with current data from two years of monitoring (2018–2019) and provided evidence of a drastic reduction in the number of sites inhabited by the species. A preliminary assessment suggested that the population of P. mmemosyne occurring in the Park is limited to a few individuals. We argue that the population of this iconic and charismatic butterfly is at the verge of extinction in this National Park, probably because of a combination of habitat loss (i.e. decrease in size and number of areas of open grassland on the mountain belt) and climatic changes. Being one of the few populations of P. mnemosyne in the Northern Apennines, the implementation of protection measures is a high conservation priority. Several other butterfly species on the Italian mountains that are facing the same survival challenges would also benefit from the establishment of conservation actions aimed at improving habitat quality for P. mnemosyne. Present article is meant to call for action researchers, stakeholders, and especially decision-makers in order to increase the efforts to upturn the evident decline in abundance of this population. Key words: biodiversity conservation, butterflies, habitat loss, habitat management, Lepidoptera, Parnassinae

The aculeate wasps are one of the most diverse and speciose insect taxa; they are omnipresent across ecosystems and exhibit diverse co-evolutionary and exploitative associations with other organisms. There is widespread conjecture that aculeate wasps are likely to perform essential ecological and economic services of importance to the health, well-being and nutritional needs of our planet. However, the scope and nature of the ecosystem services they provide are not well understood relative to other insect groups (e.g. bees, butterflies, beetles), and appreciation of their value is further tarnished by their public reputation as pointless pests. Here, we conduct the first comprehensive review of how aculeate wasps contribute to the four main areas of ecosystem services: regulatory, provisioning, supporting and cultural services. Uniting data from a large but previously disconnected literature on solitary and social aculeate wasps, we provide a synthesis on how these insects perform important ecosystem services as parasitoids, predators, biological indicators, pollinators, decomposers and seed dispersers; and their additional services as a sustainable alternative to meat for human consumption, and medicinal potential as sources of research leads for anti-microbials and cancer treatments. We highlight how aculeate wasps offer substantial, but largely overlooked, economic benefits through their roles in natural pest management and biological control programs. Accordingly, we provide data-driven arguments for reasons to consider the ecosystem service value of aculeate wasps on a par with other ‘useful’ insects (e.g. bees). Finally, we provide a research roadmap identifying the key areas of research required to capitalise better on the services provided by these important insects.

The abandonment of agricultural land and the afforestation of grassland habitats represent major threats for butterflies in European and Mediterranean areas. A crucial goal for Lepidoptera conservation is to maintain and/or restore habitat quality by targeted management. Nevertheless, there are few experimental studies allowing to derive data-driven strategies to protect butterflies of open grasslands in the Mediterranean region. We developed a habitat management strategy for the conservation of the Italian endemic butterfly Zerynthia cassandra by adopting a three-step procedure: (i) characterising which environmental and host plant features influence oviposition on plants; (ii) identifying and testing the effect of a data-driven habitat management intervention; (iii) understanding which micro-habitat features promote Z. cassandra oviposition in restored places to optimise the intervention. Both patch (areas of 1 m radius hosting Aristolochia shoots) and plant features affect oviposition, with the strongest positive effects showed by high irradiation of the patch and plant quality (high number of flowers and leaves). Accordingly, the management consisted in vegetation cuts to increase irradiation, and 2 years of monitoring demonstrated that this procedure significantly increased oviposition (average increase of about 2 eggs per plant) and larval presence. Micro-habitat sampling demonstrated that the maximum oviposition differed between vegetation structures, highlighting the importance of a local fine-tuning before the intervention. We provided a data-driven, effective, and sustainable management strategy to increase habitat suitability and oviposition for an endemic and endangered Mediterranean butterfly. Our framework can drive management strategies for other species with similar ecological requirements and subjected to similar threats.

Endemics co-occur because they evolved in situ and persist regionally or because they evolved ex situ and later dispersed to shared habitats, generating evolutionary or ecological endemicity centres, respectively. We investigate whether different endemicity centres can intertwine in the region ranging from Alps to Sicily, by studying their butterfly fauna. We gathered an extensive occurrence data set for butterflies of the study area (27,123 records, 269 species, in cells of 0.5 × 0.5 degrees of latitude-longitude). We applied molecular-based delimitation methods (GMYC model) to 26,557 cytochrome c oxidase subunit 1 (COI) sequences of Western Palearctic butterflies. We identified entities based on molecular delimitations and/or the checklist of European butterflies and objectively attributed occurrences to their most probable entity. We obtained a zoogeographic regionalisation based on the 69 endemics of the area. Using phylogenetic ANOVA we tested if endemics from different centres differ from each other and from nonendemics for key ecological traits and divergence time. Endemicity showed high incidence in the Alps and Southern Italy. The regionalisation separated the Alps from the Italian Peninsula and Sicily. The endemics of different centres showed a high turnover and differed in phylogenetic distances, phenology and distribution traits. Endemics are on average younger than nonendemics and the Peninsula-Sicily endemics also have lower variance in divergence than those from the Alps. The observed variation identifies Alpine endemics as paleoendemics, now occupying an ecological centre, and the Peninsula-Sicily ones as neoendemics, that diverged in the region since the Pleistocene. The results challenge the common view of the Alpine-Apennine area as a single “Italian refugium”., Support for this research was provided by ‘La Caixa’ Foundation (ID 100010434) to MM (grant LCF/BQ/DR20/11790020), the Ramón y Cajal program of the Spanish Ministry of Science and Innovation to GT (RYC2018-025335-I), the Academy of Finland to VD (Academy Research Fellow, decision no. 328895), by project PID2019-107078GB-I00/AEI/10.13039/501100011033 to RV and by the projects "Ricerca e conservazione sui lepidotteri diurni di sei Parchi Nazionali dell’Appennino centro-settentrionale”, “Ricerca e conservazione sugli Impollinatori dell’Arcipelago Toscano e divulgazione sui Lepidotteri del parco” and “Servizio di attuazione delle azioni per la protezione degli impollinatori e diffusione dell’entomofauna del Parco Nazionale dell'Alta Murgia” to LD.

Eusocial insect colonies represent some of the most extreme examples of specialized division of labor. Ageing in workers is often associated with a temporal polyethism in the tasks performed both i...

Reproduction in cooperative animal groups is often dominated by one or a few individuals, with the remaining group members relegated to nonreproductive helping roles. This reproductive skew can evolve if helpers receive fitness benefits such as potential future inheritance of the breeding position, but the mechanisms by which inheritance is determined are not well resolved. Polistes paper wasps form highly reproductively skewed groups and inheritance of the breeding position is likely to play a key role in the maintenance of this social structure, making them excellent models for the processes by which simple societies are maintained. Reproductive succession is thought to be determined via an age-based convention in some Polistes species, but there is also evidence for contest-based succession systems in which the replacement queen uses physical aggression to overpower and thereby subordinate her nestmates. Here, we provide evidence that queen succession in colonies of the European paper wasp Polistes dominula is determined via convention rather than contest, with little disruption to the colony’s social functioning. We use queen removal experiments and fine-scale behavioral analyses to confirm that age is a strong predictor of succession, and that behavioral responses to queen removal are restricted to the oldest individuals rather than being experienced equally across the group. We provide the most comprehensive and detailed experimental analysis on the dynamics of breeder succession in a cooperatively breeding invertebrate to date, thereby shedding light on the mechanisms by which animal societies are able to maintain cohesion in the face of within-group conflict.

1. Invasive alien species are a major threat to biodiversity. In addition to predation and parasitism, native species might suffer from competition when invasive alien species occupy a similar ecological niche. 2. This study focused on the potential interspecific interaction between two hornets: the Asian yellow‐legged hornet, Vespa velutina, a high‐concern invasive alien species recently arrived in Europe; and the native European hornet, Vespa crabro. The two species share a similar ecological niche and V. velutina is rapidly expanding across Europe, which suggests that V. crabro might suffer from competition. 3. Under laboratory‐controlled conditions, two life‐history traits that might cause the two species to compete were investigated: (i) the ability of workers to find food sources and their flexibility in exploiting them (through individual food item choice tests and exploration assays); and (ii) the worker resistance to pathogens (through immune challenge tests). 4. The results show that trophic preference of both species highly overlaps, with a marked dietary preference for honeybees compared with other insect prey and non‐prey protein items. No differences were observed in the exploratory behaviour of both species. Finally, constitutive antibacterial activity was greater in workers of the native species than in workers of the invasive hornet. 5. This laboratory study provides a first assessment under controlled conditions of the factors affecting competition between workers of two hornet species and proposes a framework to assess, in wild contexts, the magnitude of the competition and the impact of the introduced V. velutina on the native V. crabro.

Nestmate recognition, i.e., the ability to discriminate nestmates from foreign individuals, is a crucial feature of insect societies, and it has been traditionally considered to be predominantly based on chemical cues. Recent empirical evidence, however, suggests a relevant plasticity in the use of different communication channels according to cue availability and reliability in different contexts. In particular, visual cues have been shown to influence various types of social recognition in several social insects, but their role in nestmate recognition is still under-investigated. We tested the hypothesis of plasticity in the use of visual and chemical recognition cues in the primitively eusocial wasp Polistes dominula, in which the availability and reliability of recognition cues vary across the colony cycle. Indeed, before the emergence of workers, P. dominula colonies are rather small (one to few individuals), and the variability in the facial pattern might allow resident wasps to use visual cues for nestmate recognition. After workers' emergence, the increase in the number of colony members reduces the reliability of visual cues, thus leaving chemical cues as the most reliable nestmate recognition cues. We thus predict a differential use of chemical and visual cues along colony life. We experimentally separated visual and chemical cues of nestmates and non-nestmates and presented them alone or in combination (with coherent or mismatched cues) to resident wasps to test which communication channel was used in the two stages and, in case, how visual and chemical cues interacted. Our results show, for the first time in a social insect, the differential use of visual and chemical cues for nestmate recognition in two different phases of colony, which supports the hypothesis of a plastic, reliability-based use of recognition cues in this species according to the different colonial contexts.

Understanding host plant preference and the relative quality of resource provided by co-occurring host plants is a key step to predict butterfly species abundance and responses to environmental changes, and, consequently, to plan management measures. Zerynthia cassandra is an Italian endemic species strongly dependent on the availability of its host plants, Aristolochia rotunda and Aristolochia lutea. The insular population occurring on Elba island (Tuscan Archipelago) is highly threatened, because of limited host plant distribution, small population size and apparent lack of gene flow with the mainland. In 2017, we carried out field surveys and rearing experiments to (i) identify the characteristics of the host plants (vegetative status) and the site characteristics (aspect, irradiation, distance from other patches) correlated with the number of eggs occurring on individual plants, (ii) compare larval growth, food-conversions rate and larval and adult survivorship on the two host plants species. Egg occurrence depends on patch irradiation, the number of leaves and flowers occurring on individual plants and the occurrence of nearby patches. These findings allowed to identify the optimal Aristolochia patch features for egg laying and development. Laboratory rearing success was higher than 50% and although plant species did not show a significant effect on oviposition, we found that larval and adult survival was higher on A. rotunda. Our results suggest habitat management aimed at increasing resource availability for Z. cassandra and possible ex-situ conservation actions aimed at recovering the population in case of potential catastrophic events.

In the early stage of the invasion process, alien species may face costs linked to pioneer effect due to genetic bottleneck, drift and the consequential inbreeding depression. Thus, introduced species that show an immediate exponential growth in their invasive population should have some mechanisms to reduce such costs minimising the chance of inbreeding or allowing them to cope with them. The yellow-legged hornet Vespa velutina nigrithorax has been spotted in France in 2004; since then, the species has been invading Europe with a relentless pace. In their native range, males and reproductive females of a Chinese non-invasive sub-species of V. velutina seem to leave their nests to search for unrelated partners. However, previous studies showed a low genetic diversity and a high rate of diploid males in colonies of the invasive population, suggesting that mating could occur inside nests, where males should be able to discriminate between reproductive gynes and sterile workers. Here, we used laboratory behavioural assays to investigate the mating preferences of yellow-legged hornet males from the recent invasive population in Italy. We assessed the importance of nestmateship and female morpho-physiological traits, likely indicators of caste, in determining male attraction towards potential partners. Our results demonstrate that males are more attracted to bigger females with more abundant fat storage, good indicators of female reproductive caste in wasps, regardless of nestmateship. Our findings represent a first step in understanding the reproductive biology of V. velutina nigrithorax in its invasive range, providing a framework for future research in the field to prevent or reduce the species expansion.

Assessing the conservation status of protected species needs quantitative population data, generally obtained using Capture-Mark-Recapture methods (CMR). The exploitation of natural marking (e.g. individual morphological traits) offers an interesting alternative, based on image analyses, which may result in a less manipulation of protected species compared to the typical artificial marking method. In our 2-year CMR study, we tested for the first time in the natural setting the feasibility and the application of the computer-aided photographic identification method of Rosalia alpina using the individual elytral spots as the natural marking. The I3SC software was used for the photographic analysis. Data were collected from populations of two National Parks of central Italy during July–August in 2014 and 2015. We developed a standard procedure in order to optimise the image acquisition in the field and to acquire clear and comparable images, facilitating the I3SC screening process. The results demonstrated that the computer-aided photographic identification of natural markings can be implemented in a CMR population study of R. alpina. Our image processing approach showed that using only the elytral central spot contours made the tracing contour process less time-consuming obtaining reliable results. Furthermore, I3SC output scores were used to identify a threshold value for the identification of new individuals or recaptures, facilitating the final identification proposed by operators. Finally, we assessed the possibility of performing the methodology using a Citizen Science approach.

Cooperative breeding decreases the direct reproductive output of subordinate individuals, but cooperation can be evolutionarily favored when there are challenges or constraints to breeding independently. Environmental factors, including temperature, precipitation, latitude, high seasonality, and environmental harshness have been hypothesized to correlate with the presence of cooperative breeding. However, to test the relationship between cooperation and ecological constraints requires comparative data on the frequency and variation of cooperative breeding across differing environments, ideally replicated across multiple species. Paper wasps are primitively social species, forming colonies composed of reproductively active dominants and foraging subordinates. Adult female wasps, referred to as foundresses, initiate new colonies. Nests can be formed by a single solitary foundress (noncooperative) or by multiple foundress associations (cooperative). Cooperative behavior varies within and among species, making paper wasps species well suited to disentangling ecological correlates of variation in cooperative behavior. This data set reports the frequency and extent of cooperative nest founding for 87 paper wasp species. Data were assembled from more than 170 published sources, previously unpublished field observations, and photographs contributed by citizen scientists to online natural history repositories. The data set includes 25,872 nest observations and reports the cooperative behavioral decisions for 45,297 foundresses. Species names were updated to reflect modern taxonomic revisions. The type of substrate on which the nest was built is also included, when available. A smaller population-level version of this data set found that the presence or absence of cooperative nesting in paper wasps was correlated with temperature stability and environmental harshness, but these variables did not predict the extent of cooperation within species. This expanded data set contains details about individual nests and further increases the power to address the relationship between the environment and the presence and extent of cooperative breeding. Beyond the ecological drivers of cooperation, these high-resolution data will be useful for future studies examining the evolutionary consequences of variation in social behavior. This data set may be used for research or educational purposes provided that this data paper is cited.

The Rosalia longicorn (Rosalia alpina) is a strictly protected saproxylic beetle, widely distributed in Central and Southern Europe and mainly associated with ancient beech forests. To improve knowledge about the conservation status of R. alpina in Italy, available molecular markers (microsatellites and mitochondrial cytochrome c oxidase I(COI)) were tested for the first time on Italian populations. The study was performed in four sampling sites distributed in two areas placed in Northern (“Foreste Casentinesi” National Park) and Central Apennines (“Abruzzo, Lazio and Molise” National Park) where populational data about Rosalia longicorn were collected in the framework of the European LIFE MIPP Project. The genetic relationship among Apennine and Central/South-eastern European populations was explored by a comparison with mitochondrial DNA (mtDNA) data from literature. Microsatellite markers were only partially informative when applied to R. alpina Italian individuals, although providing some preliminary indication on an extensive gene flow among populations from the Apennines and local ongoing processes of genetic erosion. Genetic data are consistent with previous ecological data suggesting that the maintenance of variability in this species could be related to both habitat continuity and preservation of large senescent or standing dead trees in forests. Finally, a peculiar origin of the Apennine populations of R. alpina from a putative “Glacial Refugium” in Italy was inferred through COI data. The high genetic distance scored among the analysed populations and those from Central and South-eastern Europe indicates that the R. alpina deme from Apennine Mountains might represent a relevant conservation unit in Europe. Further genetic analyses will allow assessing other possible conservation units of R. alpina and, thus, defining large-scale conservation strategies to protect this endangered longhorn beetle in Europe.

The fast tracking of invasion spatial patterns of alien species is crucial for the implementation of preventive and management strategies of those species. Recently, a honeybee pest, the small hive beetle Aethina tumida (hereafter SHB), has been reported in Italy, where it colonized more than 50 apiaries in an area of about 300 km2. SHB is a nest parasite and scavenger of honeybee colonies native of Sub-Saharian Africa. Likely being helped by the globalization of apiculture, SHB underwent several invasions in the last twenty years, causing locally relevant economic impact. While many features of its biology have been addressed, an important knowledge gap concerns the spatial invasion dynamics in invaded areas. In this paper we coupled two spatial analysis techniques (geographic profiling and a density-based spatial clustering algorithm) to uncover the possible invasion pattern of SHB in Italy. We identified the port town of Gioia Tauro as the most likely point from which SHB may have spread and suggested the possible successive axes of diffusion. These putative diffusion paths suggest that the SHB spread in south Italy might have been due to a mix of natural dispersal between close apiaries and longer distance movement through faster, likely human-mediated, communication routes. Keywords: Apis mellifera, DBSCAN, Invasive alien species, Pest species, Spatial dynamics, Small hive beetle, Notifiable disease

The implementation of conservation actions requires a reliable assessment of presence and/or abundance of targeted species. This is particularly difficult for rare and elusive species. In this study the use of bottle traps and the effects of two potential baits in relation to height in the trees were tested to detect presence and assess abundance of stag beetles (Lucanidae) and flower chafers (Scarabaeidae, Cetoniinae), an important component of forest biodiversity. The study was carried out in a flood-plain forest of northern Italy. Forty-eight handcrafted traps were assigned to two height categories (1.5–2 m and 10–20 m) and three kinds of bait: (i) red wine, white wine and sugar, (ii) red wine, beer and mashed banana, (iii) no bait, as control. Fieldwork lasted 8 weeks, with 32 surveys, from May to July. Overall, we recorded 399 captures of the following species: Dorcus parallelipipedus, Lucanus cervus, Cetonia aurata, Protaetia speciosissima, P.affinis, P. morio and P. cuprea. Traps baited with red wine, white wine and sugar showed the highest detection probabilities for all the species. A clear preference for the canopy layer (traps between 10 and 20 m high) was shown by all species except for D. parallelipipedus which was mostly captured between 1.5 and 2 m of height. The study period was long enough to improve ecological knowledge on species phenology, but not enough to include the whole phenology for all of them. The method allowed the assessment of population size only for flower chafers as the number of stag beetles captures was too low.

Copyright Alessandro Campanaro et al. Rosalia alpina (Linnaeus, 1758) is a large longhorn beetle (Coleoptera: Cerambycidae) which is protected by the Habitats Directive and which typically inhabits beech forests characterised by the presence of mature, dead (or moribund) and sun-exposed trees. A revision of the current knowledge on systematics, ecology and conservation of R. alpina is reported. The research was carried out as part of the LIFE MIPP project which aims to find a standard monitoring method for saproxylic beetles protected in Europe. For monitoring this species, different methods were tested and compared in two areas of the Apennines, utilising wild trees, logs and tripods (artificially built with beech woods), all potentially suitable for the reproduction of the species. Even if all methods succeeded in the survey of the target species, these results showed that the use of wild trees outperformed other methods. Indeed, the use of wild trees allowed more adults to be observed and required less intensive labour. However, monitoring the rosalia longicorn on wild trees has the main disadvantage that they can hardly be considered “standard sampling units”, as each tree may be differently attractive to adults. Our results demonstrated that the most important factors influencing the attraction of single trunks were wood volume, sun-exposure and decay stage. Based on the results obtained during the project LIFE MIPP, as well as on a literature review, a standard monitoring method for R. alpina was developed.

Aim We describe fine-scale diversity patterns of the entire butterfly fauna occurring on the Tuscan Archipelago. By assessing the traits associated with population diversification, haplotype uniqueness and extinction, we aim to identify the factors determining the origin and maintenance of genetic diversity, and population vulnerability to environmental changes. Location Tuscan Archipelago, Sardinia, Tuscany (Italy) and Corsica (France). Methods We built a mtDNA dataset (1,303 COI sequences) for the 52 butterfly species reported in the Archipelago, also including specimens from neighbouring areas, and compiled data on 12 species traits and on the apparent extinction of species from the main islands. We calculated indices that measure genetic differentiation, and using phylogenetic regressions we evaluated the relationships between these indices and species traits. Finally, we inferred which traits are associated with disappearance of species on individual islands using phylogenetic regression. Results The overall spatial pattern of genetic diversity corresponded with the proximity of the areas, but strong contrasts were also identified between geographically close areas. Together with the island endemics, several common and widespread species had a high genetic diversification among islands and mainland. Phylogenetic regressions revealed that smaller-sized, more specialized species, with a preference for drier regions, displayed greater genetic structure and/or haplotype uniqueness. Species that disappeared from islands had a higher population diversification. Capraia has experienced a notable loss of diversity, which significantly affected species with shorter flight periods. Main conclusions Tuscan island butterflies are characterized by strong genetic contrasts and species differ in their contribution to the overall genetic diversity. By ranking the species for their contribution to genetic diversity and identifying the traits linked to the emergence and maintenance of diversity, we have developed a valuable tool for prioritizing populations as targets for monitoring and conservation action. The dataset constructed also represents a valuable resource for testing biogeographical hypotheses.

Biological invasions are a leading threat to native wildlife, human health and food production worldwide. Understanding the invasion history helps identifying introduction pathways and organizing integrated management strategies especially aimed at avoiding multiple reintroductions. We coupled a recently developed spatial analysis (Geographic profiling) with trade flows quantification to identify the most likely spreading centre of a recent invader of Europe, the spotted wing drosophila, Drosophila suzukii. This polyphagous vinegar fly recently colonized western countries, where it is heavily threatening fruit production causing severe economic losses. Characterized by a rapid spread and a huge impact, the invasion of this pest has a few precedents and it is becoming a model in invasion biology and pest management. Thanks to our spatial approach based on data presence of D. suzukii in European countries in the very first years of it spread, we update the current knowledge of a first spread in Spain and Italy, suggesting on the contrary that the South of France may be the most likely spreading centre of D. suzukii in Europe. Estimates of propagule pressure (fresh host fruits importation) support this finding as imports from contaminated South East Asian countries are higher in France than in Spain or Italy. Our study provides a first step in the comprehension of invasion history of this pest species and emphasizes geographic profiling as an efficient technique to track down invaders colonization patterns.

Complex insect societies are formed by thousands to millions of individuals which incessantly interact in the sheltered nest space under the constant pressure of different challenges, such as infectious disease and group coordination. Understanding how colony members interact in space and time may therefore help unravelling how different evolutionary pressures have moulded insect societies. We investigated the spatial positions and the association network of workers and the queen in the honeybee (Apis mellifera) society using high-resolution recordings of spatial behaviour and ‘network connections’ at the individual level. Our socio-spatial combined approach highlighted a highly compartmentalised structure inside honeybee colony. Connectivity and spatial overlap are high among same-age workers but low among different-age workers. Foragers are at the periphery of the social network, while the colony core is formed by young bees and the queen. Our study presents the first empirical quantitative description of socio-spatial organisation of a honeybee hive, confirming a highly compartmentalised structure and highlighting how the pressures exerted by ergonomics requirements could have shaped insect societies.

The social biology of the wasp Parischnogaster striatula has been studied in Peninsular Malaysia. This species shows the main characteristics of hover wasps (Vespidae, Stenogastrinae) which set them aside from the other social Vespidae. These include the use of an abdominal secretion in brood rearing, the three-phase egg deposition and the presence of only four larval instars. Colonies are invariably small sized and on average 1.92 females on the nests present developed ovaries. Colony life revolves around a dominant female, which has developed ovaries and spends most of her time on the nest. Since visual cues were recently found in two different species of Liostenogaster (Stenogastrinae), we also looked for a possible badge of status in these wasps but with negative results.

High reproductive skew among colony members is a fundamental feature of insect’s societies, where one or few individuals monopolize reproductive output. However, many social insect species retain a high reproductive plasticity, with all colony members able of developing ovaries and laying eggs. Understanding the partition of reproduction inside these plastic societies is a key step in the comprehension of social evolution. The reproductive status of social insects is usually evaluated by looking at the degree of ovarian development, and a great variety of different methods has been used even in the same species. It is not known, however, to which extent the different methods give comparable results and represent good indexes of individual egg-laying activity. We used the model paper wasp species Polistes dominula to tackle these issues, by compiling the main methods of ovarian development assessment on the same set of individuals and quantifying their egg-laying activity. Our results showed that all examined methods give similar and highly correlated results, but none of them allows reliable estimates of the egg-laying activity. Ovarian indexes (whatever method is used) are good indicators of individual reproductive potential, but can only roughly reflect actual reproductive activity. We thus propose to be very cautious in extrapolating egg-laying activity levels from ovarian index estimations.

Global change imposes rapid assessments to obtain reliable and updated distribution data to implement conservation measures. This task is undoubtedly unaffordable for numerous invertebrate species, both in terms of time and economic resources, because they are often elusive, detectable life stages are present for a restricted time and ecological data are scarce. Citizen science might be able to provide a large number of records and these data might facilitate the evaluation of extinction risks. Large saproxylic beetles represent an ideal group to assess the potential of citizen science to map distributions on a large scale geographic distribution. The data presented were collected during a citizen science program developed within the LIFE Project “Monitoring of Insects with Public Participation”, which used a website and a mobile app to involve citizens. We selected three pan European species protected under the Habitats Directive: Lucanus cervus , Morimus asper / funereus and Rosalia alpina , and we compared the data gathered by the citizen science project with distributional data from the official national species inventory. For all species we found a low overlap of occupied cells and the integration of the two datasets resulted in an increase in the distributional ranges of up to one third. Furthermore, taking into account the time frame of data collection, we found that the extent of occurrence obtained in 10 years of records present in the national dataset was comparable to data collected in only two years of the citizen science data project. Similar results were obtained for the estimates of spatial parameters for the IUCN Red List assessment. Our study shows the potential and the efficacy of citizen science projects as rapid tools to provide reliable distributional data for neglected species of high conservation priority.

The red palm weevil Rhynchophorus ferrugineus, recently spreading from its native range throughout the world, is one of the most dangerous pests for several palm species. The sterile insect technique (SIT) is a promising environmentally friendly approach for the management of this pest. An essential prerequisite of the SIT is that irradiated males released in nature are as sexually competitive as their wild counterparts. Here we evaluated the potential use of the SIT in red palm weevil males by investigating the sexual competitiveness of gamma-irradiated (Cobalt 60 source, dose 80 Gray) males in the laboratory in a natural-like social context with a balanced sex ratio and by testing their efficacy in impairing female reproduction. Our results showed that irradiation did not affect any of the mating behaviour parameters. However, the male reproductive potential was drastically reduced, as fertility was markedly decreased in females mated with irradiated males. Therefore, an 80 Gray irradiation dose is a promising option for the SIT in the control of the red palm weevil, producing sterile but sexually competitive males when tested in a natural-like mating context.

Division of labour is a major feature of insect societies. Behavioural differences can be present also during non-colonial stages of the life cycle, when it is difficult to discriminate between distinct behavioural phenotypes and by-products of differences in overall activity levels. We used the social wasp Polistes dominulus to address this issue. In pre-hibernating aggregations some individuals (helpers) perform external tasks by collecting food and providing it to cluster mates. Such helpers have been so far identified using only a descriptive approach, and their behaviour was not disentangled from a possible higher level of overall activity. Here we provide an operational definition of the helper's trait and we then compare behavioural patterns of helpers and non helpers, verifying that helpers actually represent a peculiar behavioural phenotype. Our result expands knowledge on the caste differentiation issue in Polistes wasps and on the assessment of behavioural phenotypes in a non-colonial context.

Distribution of mobile organisms on near-continent islands is mainly shaped by factors operating over ecological rather geological time. However, the phylogeography of single species has the potential to expose historical factors at work. In the present study, West Mediterranean populations of the butterfly Maniola jurtina are studied using geometric morphometrics. The distribution of the two well established lineages (Maniola jurtina jurtina in the Atlanto–Mediterranean area and Maniola jurtina janira in the Central–Eastern-Mediterranean area) on 12 islands and the adjoining continents are compared. The south-western lineage unexpectedly occurs on islands close to shores occupied by the eastern lineage. We have modelled the distribution of the lineages using three different hypotheses: (1) a contemporary isolation model, which predicts lineage occupancy of islands is linked to relative distances from neighbouring continental areas; (2) a refugial hypothesis, which predicts one lineage to be the ancestral one for the whole region studied, and then successively replaced over part of it; (3) a changing geography hypothesis, which predicts the two lineages to have evolved in their currently occupied areas, continuously sourcing islands subsequent to the Wurm maximum glaciation. Of the three models, the refugial hypothesis is most highly correlated with the observed pattern, suggesting that Mediterranean islands may function as refugia during cold periods, much as the three mainland peninsulas of Iberia, Italy and Greece are known to have done. Thereafter, hybridization on the nearest and smallest islands has occurred, with the entire process supporting the notion of the joint influence of factors in ecological and geological time. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98, 677–692.

The aim of this paper is to examine whether current and/or Pleistocene geography affect the species richness and composition of Tuscan archipelago butterflies. This archipelago is located between Tuscany (Italy) and Corsica (France). Faunistic data was obtained from the literature and surveys. Our data revealed that contemporary geography is the most important factor determining the species richness and faunal composition of Tuscan archipelago butterflies. Indeed, current area and isolation of the islands are the only predictors significantly correlated with species richness. Paleogeographic features of the archipelago are not significantly correlated with species richness. Multidimensional scaling revealed patterns similar to those reported for other living groups. Specifically, Capraia and Montecristo group together with Corsica, while Elba, Giglio, Pianosa, Gorgona, Giannutri and the fossil island of Monte Argentario group with the Tuscan mainland. Recent geography seems to affect the faunal composition. Indeed Mantel test indicates that the similarity in the faunal composition of the Tuscan Archipelago islands is mainly related to present-day island characteristics and their relative distance from Tuscany and Corsica. Our results are similar to those recently obtained for Aegean archipelago butterflies.

The LIFE project “MIPP” - Monitoring of Insects with Public Participation (11 NAT/IT/000252) is focused on selected insect species (five Coleoptera, three Lepidoptera, one Orthoptera), all included in the annexes II and IV of the Habitats Directive (HD) 92/43/EEC. One important aim is a citizen science initiative where every person may become a citizen scientist and collect faunistic data on the above species throughout Italy. Another objective of the project MIPP is the development of standard methods for monitoring the conserva- tion status of the five target beetle species. One innovative method employed is a sniffer-dog (“Osmodog”), trained to find the rare and endangered hermit beetle, Osmoderma eremita, which lives in veteran, hollow trees. The dog detects the strong smell of mature peach produced by adult males and an odor produced by the larvae. Another objective of the project MIPP is the dissemination of topics such as HD, Natura 2000, importance of dead-wood, Life projects, insect monitoring and conservation., Fragmenta Entomologica, Vol. 47 No. 1 (2015)

Invasive alien pest insect species represent a major threat for agriculture and biodiversity. Because chemical treatments employed to contrast such pests elicit serious environmental and human health problems, a great effort is currently directed to develop long term and environmentally friendly biological control strategies. However, the successful application of some promising techniques, such as the Sterile Insect Technique (SIT), requires a deep knowledge of the pest basic biology. Here, we argue that understanding pest sexual biology using a social network approach can significantly improve the performance of control strategies. For example, SIT may benefit from understanding how individuals interact and how males accede to reproduction, in order to target the most reproductively active and polygamic males. In this paper we studied the socio-sexual networks of the Asian red palm weevil (RPW) Rhynchophorus ferrugineus, a worldwide invader which is causing heavy economic impacts on several palm species. We found that the RPW has a highly promiscuous mating system, characterized by forced interruptions of pair copulations by additional males. The social network is highly non-random nor regular: few males almost monopolize reproduction, behaving as key-players in the network of matings. Additionally, males have a stable pattern of sexual behaviour over time. We use RPW social network as a case study to direct the development of management techniques such as SIT strategy.

Nest-mate recognition is fundamental for protecting social insect colonies from intrusion threats such as predators or social parasites. The aggression of resident females towards intruders is mediated by their cuticular semiochemicals. A positive relation between the amount of cues and responses has been widely assumed and often taken for granted, even though direct tests have not been carried out. This hypothesis has important consequences, since it is the basis for the chemical insignificance strategy, the most common explanation for the reduction in the amount of semiochemicals occurring in many social parasites. Here we used the social wasp Polistes dominulus , a model species in animal communication studies and host of three social parasites, to test this hypothesis. We discovered that different amounts of cuticular hydrocarbons (CHC) of a foreign female evoke quantitatively different behavioural reactions in the resident foundress. The relation between CHC quantity and the elicited response supports the idea that a threshold exists in the chemical recognition system of this species. The chemical insignificance hypothesis thus holds in a host–parasite system of Polistes wasps, even though other explanations should not be discarded.

SUMMARY Insect social life is governed by chemicals. A great number of studies have demonstrated that the blend of hydrocarbons present on the cuticle (CHCs) plays a pivotal role in intra- and inter-specific communication. It is not surprising, therefore, that social parasites, specialized in exploiting the costly parental care provided by host workers, exploit the host chemical communication system too. Throughout their life cycle, social parasites intercept and break this CHC-based code. Recently, however, several polar compounds (mainly peptides) have been found in addition to CHCs both on the cuticle and on the comb surface of social insects, and their semiochemical role has been demonstrated in some circumstances. In the present study, we used the paper wasp social parasite–host system Polistes sulcifer (Zimmerman)–Polistes dominulus (Christ) to evaluate the relative importance of the CHCs and polar compounds in two different steps of the host exploitation process: host nest detection by the pre-usurping parasite and parasite chemical integration into the host colony. After separating the polar and apolar fractions of the host nest as well as those of pre- and post-usurpation parasites, we carried out laboratory assays based on the binary choice model. Our results show that nest polar compounds neither are used by the parasite to detect the host's nest nor play a role in parasite chemical integration into the host colony. In contrast, we demonstrate that CHCs are fundamental in both steps, thus confirming their primary role in social insect life and consequently in social parasite–host interactions.

Polistes dominulus are social wasps which are the host of the strepsipteran endoparasite Xenos vesparum. In the hibernating phase, unparasitized and parasitized wasps leave natal nests and aggregate together in sheltered quarters. In aggregations, wasps are socially active, and some individuals perform helping behaviour. Here we investigated if castrated parasitized wasps perform worker tasks in mixed aggregations. Moreover, by gas chromatography and mass spectrometry, we examined the cuticular hydrocarbons of unparasitized and parasitized wasps to evaluate if the infection alters the composition of cuticular waxes that are recognition cues in social insects. In clusters, infected females do not perform helping behaviour and they are less active than unparasitized wasps. Cuticular hydrocarbons are slightly differentiated between unparasitized and parasitized wasps but, generally, unparasitized wasps are more similar to wasps infected by Xenos females compared to wasps infected by Xenos males. Wasps infected by Xenos males do not usually survive the winter. This chemical similarity is probably a consequence of the similar physiological condition of unparasitized and female-affected Polistes wasps. At this stage, it is difficult to affirm whether these modifications are a true parasite manipulation or a consequence of infection.

At a very early age several mammals establish a first dominance hierarchy, which often persists into adulthood. Social wasps offer an excellent opportunity to study such a phenomenon in insects. Indeed, foundresses of several paper wasps meet in clusters to hibernate from September to March. In spring, wasps found new associative nests where linear hierarchies occur. In the first phase of hibernation, clustering Polistes dominulus wasps show most of the social interactions occurring on the nest 6 months later. At the emergence from diapause, some females already show some behavioral and physiological traits typical of dominant individuals. Here, we investigated the significance of the interactions in the autumnal clusters. We demonstrated that in a given pair, it is more likely that the dominant wasp in autumn becomes the alpha female in spring after the nest foundation phase occurred. Moreover, we showed that dominant females in clusters have both larger body size and ovaries. As ovarian development mainly depends on the social context, our findings seem to indicate that social factors affect the tendency to dominate in aggregations. Furthermore, we suggest that some females may reinforce their physiological status by dominating in clusters, thus increasing the probability to become dominant in spring.