The Afrotropical breeding grounds of the Palearctic-African migratory painted lady butterflies (Vanessa cardui).

Migratory insects are key players in ecosystem functioning and services, but their spatiotemporal distributions are typically poorly known. Ecological niche modeling (ENM) may be used to predict species seasonal distributions, but the resulting hypotheses should eventually be validated by field data. The painted lady butterfly (Vanessa cardui) performs multigenerational migrations between Europe and Africa and has become a model species for insect movement ecology. While the annual migration cycle of this species is well understood for Europe and northernmost Africa, it is still unknown where most individuals spend the winter. Through ENM, we previously predicted suitable breeding grounds in the subhumid regions near the tropics between November and February. In this work, we assess the suitability of these predictions through i) extensive field surveys and ii) two-year monitoring in six countries: a large-scale monitoring scheme to study butterfly migration in Africa. We document new breeding locations, year-round phenological information, and hostplant use. Field observations were nearly always predicted with high probability by the previous ENM, and monitoring demonstrated the influence of the precipitation seasonality regime on migratory phenology. Using the updated dataset, we built a refined ENM for the Palearctic-African range of V. cardui. We confirm the relevance of the Afrotropical region and document the missing natural history pieces of the longest migratory cycle described in butterflies.

Two ways to be endemic. Alps and Apennines are different functional refugia during climatic cycles.

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".

How long is 3 km for a butterfly? Ecological constraints and functional traits explain high mitochondrial genetic diversity between Sicily and the Italian Peninsula.

Populations inhabiting Mediterranean islands often show contrasting genetic lineages, even on islands that were connected to the mainland during glacial maxima. This pattern is generated by forces acting in historical and contemporary times. Understanding these phenomena requires comparative studies integrating genetic structure, functional traits and dispersal constraints. Using as a model the butterfly species living across the Messina strait (3 km wide) separating Sicily from the Italian Peninsula, we aimed to unravel the mechanisms limiting the dispersal of matrilines and generating genetic differentiation across a narrow sea strait. We analysed the mitochondrial COI gene of 84 butterfly species out of 90 documented in Sicily and compared them with populations from the neighbouring southern Italian Peninsula (1,398 sequences) and from the entire Palaearctic region (8,093 sequences). For each species, we regressed 13 functional traits and 2 ecological constraints to dispersal (winds experienced at the strait and climatic suitability) against genetic differentiation between Sicily and Italian Peninsula to understand the factors limiting dispersal. More than a third of the species showed different haplogroups across the strait and most of them also represented endemic haplogroups for this island. One fifth of Sicilian populations (and 32.3% of endemic lineages) had their closest relatives in distant areas, instead of the neighbouring Italian Peninsula, which suggests high relictuality. Haplotype diversity was significantly explained by the length of the flight period, an intrinsic phenology trait, while genetic differentiation was explained by both intrinsic traits (wingspan and degree of generalism) and contemporary local constraints (winds experienced at the strait and climatic suitability). A relatively narrow sea strait can produce considerable differentiation among butterfly matrilines and this phenomenon showed a largely deterministic fingerprint. Because of unfavourable winds, populations of the less dispersive Sicilian butterflies tended to differentiate into endemic variants or to maintain relict populations. Understanding these phenomena required the integration of DNA sequences, species traits and physical constraints for a large taxon at continental scale. Future studies may reveal if the patterns here shown for mitochondrial DNA are also reflected in the nuclear genome or, alternatively, are the product of limited female dispersal.

Spatio-temporal ecological niche modelling of multigenerational insect migrations.

Modelling ecological niches of migratory animals requires incorporating a temporal dimension, in addition to space. Here, we introduce an approach to model multigenerational migratory insects using time-partitioned environmental variables (by months and years) and time- and behaviour-partitioned records (breeding records to model reproductive habitat). We apply this methodology to modelling the Palearctic-African migratory cycle of the Painted Lady butterfly (Vanessa cardui), based on data encompassing 36 years (646 breeding sites from 30 countries). Each breeding record is linked to a particular time (month and year), and the associated values of the bioclimatic variables are used for an ensemble modelling strategy, to finally obtain monthly projections. The results show obligated movements, mostly latitudinal, for the species' successive generations across the overall range, and only scattered locations show high probabilities of reproduction year-round. The southernmost reproductive areas estimated for the Palearctic-African migratory pool reach equatorial latitudes from December to February. We thus propose a potential distribution for the winter 'missing generations' that would expand the V. cardui migration cycle to encompass about 15 000 km in latitude, from northernmost Europe to equatorial Africa. In summer, Europe represents the major temporary resource for V. cardui, while January and February show the lowest overall suitability values, and they are potentially the most vulnerable period for the species to suffer yearly bottlenecks. In summary, we demonstrate the potential of the proposed niche modelling strategy to investigate migratory movements of insects.

Ectoparasite sharing among native and invasive birds in a metropolitan area.

Parasite-mediated competition has been reported to be one of the most harmful, although overlooked, impacts that alien species have on native ecosystems. Monk parakeets Myiopsitta monachus are successful invaders in Europe, where they have been introduced from South America. Colonial nests of these parrots may also host other species, e.g. the rock pigeon Columba livia forma domestica. In this work, we analysed the ectoparasite composition of monk parakeets in Barcelona (Spain) and we evaluated their potential role as parasite-mediated competitors, by comparing their parasitic load with that of coexisting rock pigeons. Only two arthropod species were observed on monk parakeets, whereas four species were detected on pigeons. Parakeets were rarely infested by pigeon parasites (prevalence = 0.66%), whereas parakeet mites were recorded more often on pigeons (prevalence = 10.00%). The number of total parasites per bird increased with increasing densities of monk parakeets, both for pigeons and for parakeets. Therefore, overcrowding of birds due to the increasing population of monk parakeets in Barcelona may affect the health status of native pigeons, suggesting a potential role for parasite mediated competition by introduced parakeets. Furthermore, spill-over of alien mites (Ornithonyssus bursa) by monk parakeets to rock pigeons should be monitoring as it may affect human health.

Oscheius tipulae in Italy: Evidence of an Alien Isolate in the Integral Natural Reserve of Montecristo Island (Tuscany).

Montecristo Island is an integral natural reserve of the Tuscan Archipelago National Park (Central Italy), characterized by a peculiar assemblage of flora and fauna, with several endemic taxa, and also with a high number of alien species. During a soil survey, we found an alien Oscheius tipulae Lam & Webster, 1971 isolate, phylogenetically close to others from South America. In this article, we examined the possible pathways of introduction of this nematode. Because of the high number of alien plants in this protected area and the low desiccation survival ability of O. tipulae, we hypothesized that the presence of this alien nematode isolate may be related to the soil of introduced plants, although historical association with plant-associated invertebrates is also possible. Further studies with more populations and marker molecules are necessary to investigate the distribution of O. tipulae and the possible impact on this natural reserve.

Ectoparasite load in the crested porcupine Hystrix cristata Linnaeus, 1758 in Central Italy.

The crested porcupine Hystrix cristata is a large body-sized rodent, occurring in Europe only in the Italian Peninsula, where it may have been introduced in early Medieval times. Its parasite fauna is currently poorly known and limited to few anecdotal observations. We have analyzed the ectoparasite load of 165 crested porcupines from Tuscany and Latium (Central Italy). Both captured and road-killed individuals were checked for fleas and ticks. Overall, only 39 porcupines were infested by four species of ticks and five of fleas. Abundance of ectoparasites was higher in areas with higher habitat richness, with respect to densely wooded areas. The most frequent species was the flea Pulex irritans (25%), whose prevalence peaked in winter probably because of optimal abiotic conditions in the porcupine's den. The remaining species of both hard ticks (Rhipicephalus bursa, Pholeoixodes hexagonus, and Ixodes ventalloi) and fleas (Paraceras melis, Ctenocephalides canis, Dasypsyllus gallinulae, and Hystrichopsylla talpae), all with prevalence lower than 5%, could be due to den sharing with other vertebrates, mainly carnivores such as, e.g., red foxes and badgers. The second most prevalent species was the generalist tick Ixodes ricinus (21%). An adult male-biased parasitism for ticks has been detected, suggesting a possible role of testosterone related immune-depressive effect. The low richness in dominant ectoparasite species, built up by locally acquired generalist taxa, provides support to the allochthonous origin of this rodent in Italy.

Macroparasites of introduced parakeets in Italy: a possible role for parasite-mediated competition.

Alien species are considered a cause of biodiversity loss throughout the world. An important but often overlooked form of competition with native species is the parasite-mediated one. Introduced species may bring their own parasites from their native ranges (spillover) or get native parasites from native species, thus increasing the parasites' spread and transmission risk (spillback). Thus, a complete knowledge of parasites hosted by introduced species is important to assess and to possibly prevent impacts. Ring-necked and monk parakeets have been introduced in many European countries, where they established a number of alien reproductive populations. We sampled 21 ring-necked parakeets and 7 monk parakeets from Italy and identified 35 arthropod ectoparasites belonging to five species. Amongst those, one species was native to India (Neopsittaconirmus lybartota), where alien populations of ring-necked parakeet may have been originated, and one species from South America (Paragoniocotes fulvofasciatus), which is typically found of the monk parakeet in its native range. The other three species of arthropod parasites were native to Italy and commonly found on native species, suggesting the possibility of spillback processes.

Host specificity of the badger's flea (Paraceras melis) and first detection on a bat host.

Defining the whole spectrum of potential hosts of a parasite has large epidemiological and evolutionary implications in biology. Specialized parasites might be able to occasionally exploit a range of different host species, increasing the individual survival and the chances of successful dispersal. For long time Paraceras melis has been considered a specific flea of European badger Meles meles. Anyway, it has occasionally been reported on different hosts. In this work, we summarize the host spectrum of P. melis from literature and we report its first detection on a bat host. Ten species were identified as occasional hosts, man included, and the plasticity of this flea in host exploitation is noteworthy because of possible increase of pathogens transmission to humans and domestic species.

Alien ants spreading through Europe: Brachyponerachinensis and Nylanderiavividula in Italy.

The number of known alien ant species throughout Europe has been steadily increasing during the last few decades and Italy has been no exception, with four new taxa reported in the last five years. Here, we document new data on the Asian needle ant Brachyponerachinensis (Emery, 1895), an invasive alien species whose first establishment in Europe was detected in the southern Italian city of Naples in 2022 and which has now been found near Lake Como in northern Italy, representing the second European record, about 730 km distant from the first. Furthermore, we report for the first time the presence of Nylanderiavividula (Nylander, 1846) in the country, based on specimens collected both in Rome and near Lake Como. This is at least the second Nylanderia species established in the country after N.jaegerskioeldi, first reported in 2018. Unlike B.chinensis, N.vividula is not considered an ecological and health threat in the invaded range and is already known to occur in several other European countries. While only a few introduced ants in Europe are considered serious ecological, economic or health threats, the increasing circulation of several alien species and the poor ability to swiftly track their movements and detect their establishment can render management very difficult.

Response to Genovesi et al.: Ant biosurveillance should come before invasion.

Menchetti et al. respond to the letter of Genovesi et al. and contribute new records of the red imported fire ant in Sicily.

The genetic legacy of the Quaternary ice ages for West Palearctic butterflies.

The interplay between geographic barriers and climatic oscillations over the past 2.6 million years structured genetic variation at the continental scale. The genetic legacy of the Quaternary ice ages (GLQ) hypothesis outlines this phenomenon for Europe, but a comprehensive data-driven assessment is lacking. Using innovative genetic landscape methods, we model the GLQ in the West Palearctic based on 31,653 Cytochrome c oxidase subunit 1 (COI) sequences from 494 butterfly species and three functional traits. Seven distinct bioregions with varying levels of genetic endemicity emerge, revealing a latitudinal gradient in variation that confirms the "southern richness, northern purity" hypothesis. Through shift from case studies to a comparative approach, we objectively identify the main glacial refugia, colonization routes, and barriers to dispersal. Our findings offer a quantitative model of the GLQ across Europe, North Africa, and neighboring Asia, with broader applicability to other taxa and potentially scalable to encompass life on Earth.

A genetic atlas for the butterflies of continental Canada and United States.

Multi-locus genetic data for phylogeographic studies is generally limited in geographic and taxonomic scope as most studies only examine a few related species. The strong adoption of DNA barcoding has generated large datasets of mtDNA COI sequences. This work examines the butterfly fauna of Canada and United States based on 13,236 COI barcode records derived from 619 species. It compiles i) geographic maps depicting the spatial distribution of haplotypes, ii) haplotype networks (minimum spanning trees), and iii) standard indices of genetic diversity such as nucleotide diversity (π), haplotype richness (H), and a measure of spatial genetic structure (GST). High intraspecific genetic diversity and marked spatial structure were observed in the northwestern and southern North America, as well as in proximity to mountain chains. While species generally displayed concordance between genetic diversity and spatial structure, some revealed incongruence between these two metrics. Interestingly, most species falling in this category shared their barcode sequences with one at least other species. Aside from revealing large-scale phylogeographic patterns and shedding light on the processes underlying these patterns, this work also exposed cases of potential synonymy and hybridization.

Pollen metabarcoding reveals the origin and multigenerational migratory pathway of an intercontinental-scale butterfly outbreak.

Migratory insects may move in large numbers, even surpassing migratory vertebrates in biomass. Long-distance migratory insects complete annual cycles through multiple generations, with each generation's reproductive success linked to the resources available at different breeding grounds. Climatic anomalies in these grounds are presumed to trigger rapid population outbreaks. Here, we infer the origin and track the multigenerational path of a remarkable outbreak of painted lady (Vanessa cardui) butterflies that took place at an intercontinental scale in Europe, the Middle East, and Africa from March 2019 to November 2019. Using metabarcoding, we identified pollen transported by 264 butterflies captured in 10 countries over 7 months and modeled the distribution of the 398 plants detected. The analysis showed that swarms collected in Eastern Europe in early spring originated in Arabia and the Middle East, coinciding with a positive anomaly in vegetation growth in the region from November 2018 to April 2019. From there, the swarms advanced to Northern Europe during late spring, followed by an early reversal toward southwestern Europe in summer. The pollen-based evidence matched spatiotemporal abundance peaks revealed by citizen science, which also suggested an echo effect of the outbreak in West Africa during September-November. Our results show that population outbreaks in a part of species' migratory ranges may disseminate demographic effects across multiple generations in a wide geographic area. This study represents an unprecedented effort to track a continuous multigenerational insect migration on an intercontinental scale.

The invasive ant Solenopsis invicta is established in Europe.

The red imported fire ant (Solenopsis invicta) is classified as one of the worst invasive alien species1 and as the fifth costliest worldwide2, impacting ecosystems, agriculture and human health3. We report the establishment of S. invicta in Europe for the first time, documenting a mature population in Sicily. We use genetic analyses to assess its putative origin, as well as wind tracking and species distribution modelling to predict its potential range on the continent. We show that half of the urban areas in Europe are already suitable and that climate warming expected under current trends will favor the expansion of this invasive ant.

Unrecognized for centuries: distribution and sexual caste descriptions of the West European Aphaenogaster species of the subterranea group (Hymenoptera, Formicidae).

There are only two Aphaenogaster species from the subterranea group in the western Mediterranean: A.ichnusa Santschi, 1925, from south-western Europe, and A.subterranea (Latreille, 1798), also occurring in central and eastern Europe. Historically, the two species have been widely misunderstood: A.ichnusa was long considered a Sardinian endemic subspecies of A.subterranea, while its continental populations were misidentified as A.subterranea s. str. Recently, A.ichnusa was elevated to species rank and its worker caste was redescribed with that of A.subterranea, allowing for their correct identification. Yet their distribution was documented in detail only for France and Sardinia. Furthermore, no morphological characters were described to distinguish the males and queens of the two species. By investigating private and museum collections, 276 new records of A.ichnusa are provided here and 154 of A.subterranea from the western Mediterranean. Additionally, qualitative and quantitative morphological characters were combined to identify their males and queens. We present the new southernmost, easternmost, and westernmost distribution limits for A.ichnusa. Based on our results, this species is widely distributed in Italy and Catalonia (Spain), also occurring on several Mediterranean islands, avoiding areas with continental climate and high altitudes. Sicily is the only island to host the less thermophilous A.subterranea, which otherwise extends westward to Galicia (Spain). Sympatric occurrence is not rare along the contact zone. Additional natural history observations are reported regarding foraging habits, associated myrmecophiles, habitat preferences, and colony structure in the two species.

Observer-oriented approach improves species distribution models from citizen science data.

Citizen science platforms are increasingly growing, and, storing a huge amount of data on species locations, they provide researchers with essential information to develop sound strategies for species conservation. However, the lack of information on surveyed sites (i.e., where the observers did not record the target species) and sampling effort (e.g., the number of surveys at a given site, by how many observers, and for how much time) strongly limit the use of citizen science data. Thus, we examined the advantage of using an observer-oriented approach (i.e., considering occurrences of species other than the target species collected by the observers of the target species as pseudo-absences and additional predictors relative to the total number of observations, observers, and days in which locations were collected in a given sampling unit, as proxies of sampling effort) to develop species distribution models. Specifically, we considered 15 mammal species occurring in Italy and compared the predictive accuracy of the ensemble predictions of nine species distribution models carried out considering random pseudo-absences versus observer-oriented approach. Through cross-validations, we found that the observer-oriented approach improved species distribution models, providing a higher predictive accuracy than random pseudo-absences. Our results showed that species distribution modeling developed using pseudo-absences derived citizen science data outperform those carried out using random pseudo-absences and thus improve the capacity of species distribution models to accurately predict the geographic range of species when deriving robust surrogate of sampling effort.

First records of two alien land flatworms (Tricladida, Geoplanidae) from Northeastern Italy.

During a period of intense rainfall (May 2019), several specimens of land flatworms were collected from a private garden in Palazzolo dello Stella (Friuli Venezia Giulia, Udine, Italy: 45°47'40.5"N, 13°05'17.2"E). Planarians were found both in a cultivated part of the garden and in a part covered with gravel and with trees and shrubs (Pyracantha sp., Olea europaea, Pyrus communis). The animals were observed under branches, stones, tufa blocks, and pots close to a small artificial pond, but also in other parts of the garden, as well as inside buildings.

Updated distribution of the invasive Megachile sculpturalis (Hymenoptera: Megachilidae) in Italy and its first record on a Mediterranean island.

Megachile sculpturalis (Smith, 1853) (Hymenoptera: Megachilidae) is an invasive solitary bee that is rapidly spreading all over Europe. The present study aims to update the distribution of this species in Italy. The research led to the collection of 177 records, obtained through bibliographic research and data-mining from websites, blogs and social networks. We here present the first record of M. sculpturalis on a Mediterranean island and discuss its possible effect on the native ecosystem. Given the particular discovery of M. sculpturalis on Elba Island (Tuscany), we suggest possible monitoring, containment and possible eradication measures of the species.