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The fast rate of replacement of natural areas by expanding cities is a key threat to wildlife worldwide. Many wild species occur in cities, yet little is known on the dynamics of urban wildlife assemblages due to species' extinction and colonization that may occur in response to the rapidly evolving conditions within urban areas. Namely, species' ability to spread within urban areas, besides habitat preferences, is likely to shape the fate of species once they occur in a city. Here we use a long-term dataset on mammals occurring in one of the largest and most ancient cities in Europe to assess whether and how spatial spread and association with specific habitats drive the probability of local extinction within cities. Our analysis included mammalian records dating between years 1832 and 2023, and revealed that local extinctions in urban areas are biased towards species associated with wetlands and that were naturally rare within the city. Besides highlighting the role of wetlands within urban areas for conserving wildlife, our work also highlights the importance of long-term biodiversity monitoring in highly dynamic habitats such as cities, as a key asset to better understand wildlife trends and thus foster more sustainable and biodiversity-friendly cities.
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Ecosistema , Humedales , Animales , Ciudades , Mamíferos , Biodiversidad , Animales SalvajesRESUMEN
Studying hybrid zones that form between morphologically cryptic taxa offers valuable insights into the mechanisms of cryptic speciation and the evolution of reproductive barriers. Although hybrid zones have long been the focus of evolutionary studies, the awareness of cryptic hybrid zones increased recently due to rapidly growing evidence of biological diversity lacking obvious phenotypic differentiation. The characterization of cryptic hybrid zones with genome-wide analysis is in its early stages and offers new perspectives for studying population admixture and thus the impact of gene flow. In this study, we investigate the population genomics of the Myotis nattereri complex in one of its secondary contact zones, where a putative hybrid zone is formed between two of its cryptic lineages. By utilizing a whole-genome shotgun sequencing approach, we aim to characterize this cryptic hybrid zone in detail. Demographic analysis suggests that the cryptic lineages diverged during the Pliocene, c. 3.6 million years ago. Despite this ancient separation, the populations in the contact zone exhibit mitochondrial introgression and a considerable amount of mixing in nuclear genomes. The genomic structure of the populations corresponds to geographic locations and the genomic admixture changes along a geographic gradient. These findings suggest that there is no effective hybridization barrier between both lineages, nevertheless, their population structure is shaped by dispersal barriers. Our findings highlight how such deeply diverged cryptic lineages can still readily hybridize in secondary contact.
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Quirópteros , Flujo Génico , Especiación Genética , Genética de Población , Hibridación Genética , Animales , Quirópteros/genética , Quirópteros/clasificación , ADN Mitocondrial/genética , Introgresión GenéticaRESUMEN
The widespread decline of biodiversity due to increasing urban development raises the need to timely identify areas most relevant to the conservation of native species, particularly within cities where natural areas are extremely limited. Here, we assess the multiple role of local geomorphological features in shaping patterns and dynamics of plant diversity, with the aim of identifying conservation values and priorities in an urbanised area of Southern Italy. Based on recent and historical lists of vascular plants, we compared the floristic composition of different portions of the area by considering species' conservation value, ecological and biogeographical traits. We found that landscape remnants, accounting for 5% of the study area, harbour over 85% of the whole plant diversity and a considerable set of exclusive species. Results of Generalised Linear Mixed Models show an outstanding role of landscape remnants for the conservation of native, rare and specialised species. Based on the compositional similarities among sampled sites resulting from hierarchical clustering, these linear landscape elements also play a key role in maintaining the floristic continuity and potential connectivity throughout the urban landscape. By comparing current biodiversity patterns with data from the early XX century, we also show that the considered landscape elements are significantly more likely to host populations of declining native species, underlining their role as refugia against past and future extinctions. Taken together, our findings represent an effective framework to tackle the challenging conservation of nature in cities, namely providing a valuable approach for the identification of priority areas for the conservation of diversity within anthropogenic landscapes.
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Conservación de los Recursos Naturales , Ecosistema , Conservación de los Recursos Naturales/métodos , Biodiversidad , Plantas , CiudadesRESUMEN
Human settlements, including cities, may provide wildlife with new ecological niches, in terms of habitat types and food availability, thus requiring plasticity for adaptation. The crested porcupine Hystrix cristata is a habitat-generalist, large-sized rodent, also recorded in some suburban areas, but no information is available on its habitat use in metropolitan landscapes. Here, we assessed the land-use factors influencing the presence of crested porcupines in a metropolitan area of Central Italy. We collected data on the occurrence of crested porcupines from the metropolitan area of Rome, following an observer-oriented approach to record occurrences and retreive pseudo-absences. We then related the presence/absence of H. cristata to landscape composition. Occupancy models showed that cultivations and scrubland were positively related to porcupine presence, most likely as they provide food resources and shelter sites, respectively. Although the crested porcupine has been confirmed as a "generalist" species in terms of habitat selection, a strong preference for areas limiting the risk of being killed and providing enough food and shelter was observed. We therefore suggest that the crested porcupine may adapt to deeply modified landscapes such as large cities by selecting specific favourable land-use types.
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Urbanisation exposes wildlife to new challenging conditions and environmental pressures. Some mammalian species have adapted to these novel environments, but it remains unclear which characteristics allow them to persist. To address this question, we identified 190 mammals regularly recorded in urban settlements worldwide, and used phylogenetic path analysis to test hypotheses regarding which behavioural, ecological and life history traits favour adaptation to urban environments for different mammalian groups. Our results show that all urban mammals produce larger litters; whereas other traits such as body size, behavioural plasticity and diet diversity were important for some but not all taxonomic groups. This variation highlights the idiosyncrasies of the urban adaptation process and likely reflects the diversity of ecological niches and roles mammals can play. Our study contributes towards a better understanding of mammal association to humans, which will ultimately allow the design of wildlife-friendly urban environments and contribute to mitigate human-wildlife conflicts.
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Ecología , Mamíferos , Urbanización , Adaptación Fisiológica , Animales , Ecosistema , Humanos , FilogeniaRESUMEN
Managing biological invasions requires rapid, cost-effective assessments of introduced species' occurrence, and a good understanding of the species' vegetation associations. This is particularly true for species that are elusive or may spread rapidly. Finlayson's squirrel (Callosciurus finlaysonii) is native to Thailand and southeastern Asia, and two introduced populations occur in peninsular Italy. One of the two introduced populations is rapidly expanding, but neither effective monitoring protocols nor reliable information on vegetation associations are available. To fill this gap, we conducted visual surveys and hair tube sampling in a periurban landscape of southern Italy to compare the effectiveness of these two methods in assessing presence of Finlayson's squirrel. We also determined the species' association with vegetation types at detection locations and nesting sites. Both visual and hair tube sampling effectively assessed the species' presence, but hair tubes resulted in fewer false absences. Moreover, when we controlled for the costs of labor and equipment, hair tubes were 33.1% less expensive than visual sampling. Presence of squirrels and their nests was positively correlated with shrub species richness, indicating that the occurrence of forests with well-developed understory may inhibit the spread of the species.
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Monitoreo del Ambiente/métodos , Bosques , Especies Introducidas/tendencias , Sciuridae/crecimiento & desarrollo , Animales , Italia , Dinámica Poblacional , TailandiaRESUMEN
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.
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Enfermedades de las Aves/parasitología , Especies Introducidas , Periquitos , Enfermedades Parasitarias en Animales/parasitología , Animales , Enfermedades de las Aves/epidemiología , Italia/epidemiología , Enfermedades Parasitarias en Animales/epidemiologíaRESUMEN
In this article we review the current knowledge of the effects of urban expansion on bats and assess the potential of these mammals as bioindicators of urbanization. The response of bats to this process is highly species-specific: some species tolerate urban habitat or are even favoured by its roosting or foraging opportunities, others are affected by the loss or fragmentation of key natural habitat, or by the physical and chemical pollution associated with urbanization. Species responses generally translate into altered community structures, with few markedly dominating species. We propose different hypothetical models of bat fitness along an urbanization gradient and discuss why bat population density may not be an effective fitness proxy to assess the reactions of these mammals to urban expansion. We also suggest that urban habitat may act as an ecological trap even for apparently synurbic species. Overall, bat sensitivity to urbanization makes these mammals promising candidates to track the effects of this process of land use change on the biota, but more studies, specifically tailored to explore this role, are needed.
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Bats are highly social mammals that often form large groups and represent good models to test the role played by individual status in shaping social relationships. Social cohesion relies on the ability of group and individual recognition, which is mediated by a range of sensorial cues. In this study, we selected the European free-tailed bat Tadarida teniotis as a model species to test the effects of familiarity, sex and age on aggressiveness and mutual tolerance. We hypothesize that T. teniotis is able to recognize group members and exhibit selective aggressiveness, and thus we predict fewer aggressive events and more amicable encounters between colony mates than between strangers. As female bats are generally more sociable and perform prolonged parental care to juveniles even after weaning, we hypothesize that sex and age of bats have significant influences on aggressive behaviours and thus predict that females will perform more amicable behaviours than males and that adults of both sexes will be less aggressive towards juveniles. Our results confirm that T. teniotis is able to discriminate between familiar and stranger individuals, showing higher rates of aggressive behaviours towards the latter. Females are more prone to exhibit amicable behaviours, particularly during same-sex interactions, while males show higher level of aggressiveness. Juveniles are subjected to fewer aggressive behaviours by adults of both sexes. Familiarity appears crucial for T. teniotis in determining the degree of aggressiveness during social interactions but the rate of aggressive events is also influenced by intrinsic individual factors such as sex and age.
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Agresión/fisiología , Quirópteros/fisiología , Reconocimiento en Psicología/fisiología , Factores de Edad , Animales , Femenino , Masculino , Factores SexualesRESUMEN
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.
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Quirópteros/parasitología , Siphonaptera , Animales , Femenino , Especificidad del Huésped , Mustelidae/parasitología , Siphonaptera/fisiologíaRESUMEN
European semi-natural dry grasslands are among the most endangered terrestrial ecosystems, being recognised as habitats of community interest by the EU Habitats Directive. The occurrence and preservation of these habitats depend on a combination of anthropogenic and natural factors, although little is known regarding the role of past land-use changes. Here, we investigated the role of time since cultivation abandonment as a major driver of grassland successional dynamics in the Mediterranean agro-pastoral system of Alta Murgia, southern Italy. By integrating cartographic information on the past agricultural land-use with the main abiotic constraints (patch area, slope and aspect), we used generalised additive mixed models to test for the probability of occurrence of current grassland habitat types along time since cultivation abandonment (10 to 200 years). Our results disclosed the successional sequence of grassland plant communities since crop abandonment in the study area, highlighting that the distribution of semi-natural grassland communities largely depends on land use history besides current environmental patterns. Among the habitat types protected under the EU Habitats Directive, we highlighted that xero-thermic communities may represent an intermediate step of grassland succession after cultivation abandonment, while more mesic perennial communities indicate a late successional stage. These successional dynamics are further modulated by mesoclimatic conditions associated with slope and aspect, especially in case of long-standing pastures that were not historically affected by agricultural transformations. Our findings can contribute to a deeper understanding of dynamics relevant to spontaneous vegetation recovery in open environments, which is a prerequisite for setting up effective grassland conservation and restoration actions. Furthermore, our results underline the value of integrating historical maps and current information for the assessment of habitat conservation status.
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Ecosistema , Pradera , Biodiversidad , Agricultura/métodos , ItaliaRESUMEN
Understanding how species respond to climate change is key to informing vulnerability assessments and designing effective conservation strategies, yet research efforts on wildlife responses to climate change fail to deliver a representative overview due to inherent biases. Bats are a species-rich, globally distributed group of organisms that are thought to be particularly sensitive to the effects of climate change because of their high surface-to-volume ratios and low reproductive rates. We systematically reviewed the literature on bat responses to climate change to provide an overview of the current state of knowledge, identify research gaps and biases and highlight future research needs. We found that studies are geographically biased towards Europe, North America and Australia, and temperate and Mediterranean biomes, thus missing a substantial proportion of bat diversity and thermal responses. Less than half of the published studies provide concrete evidence for bat responses to climate change. For over a third of studied bat species, response evidence is only based on predictive species distribution models. Consequently, the most frequently reported responses involve range shifts (57% of species) and changes in patterns of species diversity (26%). Bats showed a variety of responses, including both positive (e.g. range expansion and population increase) and negative responses (range contraction and population decrease), although responses to extreme events were always negative or neutral. Spatial responses varied in their outcome and across families, with almost all taxonomic groups featuring both range expansions and contractions, while demographic responses were strongly biased towards negative outcomes, particularly among Pteropodidae and Molossidae. The commonly used correlative modelling approaches can be applied to many species, but do not provide mechanistic insight into behavioural, physiological, phenological or genetic responses. There was a paucity of experimental studies (26%), and only a small proportion of the 396 bat species covered in the examined studies were studied using long-term and/or experimental approaches (11%), even though they are more informative about the effects of climate change. We emphasise the need for more empirical studies to unravel the multifaceted nature of bats' responses to climate change and the need for standardised study designs that will enable synthesis and meta-analysis of the literature. Finally, we stress the importance of overcoming geographic and taxonomic disparities through strengthening research capacity in the Global South to provide a more comprehensive view of terrestrial biodiversity responses to climate change.
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Quirópteros , Animales , Biodiversidad , Cambio Climático , Ecosistema , Dinámica PoblacionalRESUMEN
Knowledge of species' functional traits is essential for understanding biodiversity patterns, predicting the impacts of global environmental changes, and assessing the efficiency of conservation measures. Bats are major components of mammalian diversity and occupy a variety of ecological niches and geographic distributions. However, an extensive compilation of their functional traits and ecological attributes is still missing. Here we present EuroBaTrait 1.0, the most comprehensive and up-to-date trait dataset covering 47 European bat species. The dataset includes data on 118 traits including genetic composition, physiology, morphology, acoustic signature, climatic associations, foraging habitat, roost type, diet, spatial behaviour, life history, pathogens, phenology, and distribution. We compiled the bat trait data obtained from three main sources: (i) a systematic literature and dataset search, (ii) unpublished data from European bat experts, and (iii) observations from large-scale monitoring programs. EuroBaTrait is designed to provide an important data source for comparative and trait-based analyses at the species or community level. The dataset also exposes knowledge gaps in species, geographic and trait coverage, highlighting priorities for future data collection.
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Quirópteros , Animales , Biodiversidad , Quirópteros/fisiología , Ecosistema , Europa (Continente) , MamíferosRESUMEN
SARS-CoV-2, the virus that caused the COVID-19 pandemic, is genomically similar to a SARS-like beta-coronavirus found in Asian rhinolophid bats. This evolutionary relationship impressed the global media, which then emphasised bats as key actors in the spillover that resulted in the pandemic. In this study, we highlight changes in the traditional and new media coverage of bats and in Internet search volumes that occurred since the beginning of the COVID-19 pandemic in 2020.We analysed Google and Wikipedia searches for bats and coronaviruses in 21 countries and eight languages, as well as television broadcasts in the USA, some of which have global coverage, between January 2016 and December 2020. In January 2020, the amount of television news about bats boomed, and news associated with the term 'bat' shifted to COVID-19-related topics. A nearly identical pattern was observed in Google searches during 2020 at the global scale. The daily time series of television coverage and Internet search volumes on bats and coronavirus in the USA covaried in the first quarter of 2020, in line with the existence of a media bubble. Time-series analysis revealed that both the Google Trends index and visits to Wikipedia pages about bats boomed in early 2020, despite the fact that this time of year is usually characterised by low search volumes.Media coverage emphasised, correctly or not, the role of bats in the COVID-19 pandemic and amplified public interest in bats worldwide. The public image of these mammals, in many cases threatened and important ecosystem service providers, was seriously compromised. We therefore recommend that policymakers and journalists prioritise scientifically accurate communication campaigns about bats, which would help counteract the surge in bat persecution, and leverage interest towards positive human-bat interactions.
SARSCoV2, el virus causante de la pandemia COVID19, es genómicamente símil al SARStipo betacoronavirus encontrado en los murciélagos rinolófidos asiáticos. Esta relación evolutiva impresionó a los medios mundialmente, hasta tal punto que se ha enfatizado el papel de los murciélagos como uno de los principales vectores en la expansión de la pandemia. En este estudio, se resaltan los cambios producidos en la percepción de los murciélagos por los medios de comunicación (nuevos y tradicionales), y en las búsquedas en Internet que ocurrieron desde el inicio de la pandemia COVID19 en 2020.Para ello se han analizado las búsquedas del término 'murciélago' en Google y Wikipedia en 21 países y ocho lenguas, así como las retransmisiones televisivas en EE.UU. (algunas de las cuales con cobertura global) entre enero 2016 y diciembre 2020. En enero 2020, la cantidad de noticias televisadas sobre murciélagos se disparó, y el número de noticias previamente asociadas con el término 'murciélago' cambiaron por temáticas relacionadas con el COVID19. Un patrón idéntico puede ser observado a escala global en las búsquedas de Google durante el 2020. La serie temporal de cobertura televisiva y búsquedas en internet sobre murciélagos y coronavirus en EE.UU. covaría durante el primer cuarto del 2020, en líneo con la existencia de la burbuja mediática. El análisis de la cobertura diaria televisiva revela que tanto el índice Google Trend como las visitas a páginas de Wikipedia sobre murciélagos incrementaron al comienzo del 2020, a pesar de que esta época del año suele caracterizarse por bajos volúmenes de búsqueda.La cobertura mediática enfatizó, correctamente o no, el papel de los murciélagos en la propagación de la pandemia COVID19, ampliando el interés público a lo largo del globo. La imagen pública de estos mamíferos, en muchos casos especies amenazadas o proveedoras de importantes funciones ecosistémicas, ha sido seriamente comprometida. Por lo tanto, se recomienda a periodistas y responsables políticos la priorización de campañas de comunicación científicamente fundamentadas que ayuden a contrarrestar este aumento en la persecución de los murciélagos aprovechando el interés actual para concienciar al público de sus interacciones positivas con el hombre.
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Mimicry is one of the most fascinating phenomena in nature1. Mimicry traits often reflect complex, finely tuned, and sometimes extravagant relationships among species and have evolved to deceive predators or prey. Indeed, mimicry has most often evolved to discourage predation: the 'mimic' exhibits phenotypic convergence towards a non-related 'model' organism which is inedible or harmful, so that a given predator, or 'receiver', will refrain from attacking or ingesting the mimic. Traditionally, mimicry is mainly evident and has been mainly studied in the visual domain. Here, we report experiments that document the first case of interspecific acoustic mimicry in a mammal and demonstrate that the distress calls the greater mouse-eared bat (Myotis myotis) broadcasts when handled imitate sounds of stinging bees or wasps to discourage the bat's avian predators.
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Quirópteros , Acústica , Animales , Insectos , Conducta Predatoria , SonidoRESUMEN
Bats show responses to anthropogenic stressors linked to changes in other ecosystem components such as insects, and as K-selected mammals, exhibit fast population declines. This speciose, widespread mammal group shows an impressive trophic diversity and provides key ecosystem services. For these and other reasons, bats might act as suitable bioindicators in many environmental contexts. However, few studies have explicitly tested this potential, and in some cases, stating that bats are useful bioindicators more closely resembles a slogan to support conservation than a well-grounded piece of scientific evidence. Here, we review the available information and highlight the limitations that arise in using bats as bioindicators. Based on the limited number of studies available, the use of bats as bioindicators is highly promising and warrants further investigation in specific contexts such as river quality, urbanisation, farming practices, forestry, bioaccumulation, and climate change. Whether bats may also serve as surrogate taxa remains a controversial yet highly interesting matter. Some limitations to using bats as bioindicators include taxonomical issues, sampling problems, difficulties in associating responses with specific stressors, and geographically biased or delayed responses. Overall, we urge the scientific community to test bat responses to specific stressors in selected ecosystem types and develop research networks to explore the geographic consistency of such responses. The high cost of sampling equipment (ultrasound detectors) is being greatly reduced by technological advances, and the legal obligation to monitor bat populations already existing in many countries such as those in the EU offers an important opportunity to accomplish two objectives (conservation and bioindication) with one action.
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Bats are often unfairly depicted as the direct culprit in the current COVID-19 pandemic, yet the real causes of this and other zoonotic spillover events should be sought in the human impact on the environment, including the spread of domestic animals. Here, we discuss bat predation by cats as a phenomenon bringing about zoonotic risks and illustrate cases of observed, suspected or hypothesized pathogen transmission from bats to cats, certainly or likely following predation episodes. In addition to well-known cases of bat rabies, we review other diseases that affect humans and might eventually reach them through cats that prey on bats. We also examine the potential transmission of SARS-CoV-2, the causal agent of COVID-19, from domestic cats to bats, which, although unlikely, might generate a novel wildlife reservoir in these mammals, and identify research and management directions to achieve more effective risk assessment, mitigation or prevention. Overall, not only does bat killing by cats represent a potentially serious threat to biodiversity conservation, but it also bears zoonotic implications that can no longer be neglected.
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Artificial light at night (ALAN) is a pervasive form of pollution largely affecting wildlife, from individual behaviour to community structure and dynamics. As nocturnal mammals, bats are often adversely affected by ALAN, yet some "light-opportunistic" species exploit it by hunting insects swarming near lights. Here we used two potentially competing pipistrelle species as models, Kuhl's (Pipistrellus kuhlii) and common (Pipistrellus pipistrellus) pipistrelles, both known to forage in artificially illuminated areas. We set our study in a mountainous area of central Italy, where only recently did the two species become syntopic. We applied spatial modelling and radiotracking to contrast potential vs. actual environmental preferences by the two pipistrelles. Species distribution models and niche analysis showed a large interspecific niche overlap, including a preference for illuminated areas, presenting a potential competition scenario. Pipistrellus pipistrellus association with ALAN, however, was weakened by adding P. kuhlii as a biotic variable to the model. Radiotracking showed that the two species segregated habitats at a small spatial scale and that P. kuhlii used artificially illuminated sites much more frequently than P. pipistrellus, despite both species potentially being streetlamp foragers. We demonstrate that ALAN influences niche segregation between two potentially competing species, confirming its pervasive effects on species and community dynamics, and provide an example of how light pollution and species' habitat preferences may weave a tapestry of complex ecological interactions.
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Quirópteros , Animales , Ecosistema , Italia , Iluminación , MamíferosRESUMEN
Because of the high risk of going unnoticed, cryptic species represent a major challenge to biodiversity assessments, and this is particularly true for taxa that include many such species, for example, bats. Long-eared bats from the genus Plecotus comprise numerous cryptic species occurring in the Mediterranean Region and present complex phylogenetic relationships and often unclear distributions, particularly at the edge of their known ranges and on islands. Here, we combine Species Distribution Models (SDMs), field surveys and molecular analyses to shed light on the presence of a cryptic long-eared bat species from North Africa, Plecotus gaisleri, on the islands of the Sicily Channel, providing strong evidence that this species also occurs in Europe, at least on the islands of the Western Mediterranean Sea that act as a crossroad between the Old Continent and Africa. Species Distribution Models built using African records of P. gaisleri and projected to the Sicily Channel Islands showed that all these islands are potentially suitable for the species. Molecular identification of Plecotus captured on Pantelleria, and recent data from Malta and Gozo, confirmed the species' presence on two of the islands in question. Besides confirming that P. gaisleri occurs on Pantelleria, haplotype network reconstructions highlighted moderate structuring between insular and continental populations of this species. Our results remark the role of Italy as a bat diversity hotspot in the Mediterranean and also highlight the need to include P. gaisleri in European faunal checklists and conservation directives, confirming the usefulness of combining different approaches to explore the presence of cryptic species outside their known ranges-a fundamental step to informing conservation.