Dangerous neighbours: Birds and bird-eating bats sharing tree cavities.

Mounting evidence indicates the non-consumptive effects of predators significantly impact prey physiology, ecology and behaviour. Passerine birds experience adverse effects on nesting and reproductive success when in proximity to predators. Fear of predators is context-dependent and influenced by hunting habitats and foraging strategies. While some bat species prey on birds, the greater noctule (Nyctalus lasiopterus) stands out by specialising in avian prey, especially during peak bird migration. N. lasiopterus is thought to seize avian prey in flight, but direct evidence is lacking. If birds were taken from nests, they would likely avoid nesting near these bats. However, no observations support this view. This study documents the successful reproduction of Eurasian blue tits (Cyanistes caeruleus) nesting alongside a colony of approximately 25 greater noctules. This bird species is a prey species for greater noctules in Italy. Over about 1 month (April-May 2023), we observed parent birds provisioning food to chicks, with at least two chicks alive and fed outside the tree cavity by the end of the period. While acknowledging the limitations of a single observation, we propose that this previously unknown behaviour indirectly supports the idea that greater noctules only capture avian prey in flight, not within confined spaces. This observation challenges the perception that these bats pose a threat when sharing roosting spaces in trees, as evidenced in our observed case. We hope this novel observation inspires future research on variations in bird nesting behaviour and reproductive success in the presence of bird-eating bats, as well as an assessment of the long-term impact on population dynamics and reproductive success of nesting birds sharing such roosting spaces.

Climate is changing, are European bats too? A multispecies analysis of trends in body size.

Animal size, a trait sensitive to spatial and temporal variables, is a key element in ecological and evolutionary dynamics. In the context of climate change, there is evidence that some bat species are increasing their body size via phenotypic responses to higher temperatures at maternity roosts. To test the generality of this response, we conducted a >20-year study examining body size changes in 15 bat species in Italy, analysing data from 4393 individual bats captured since 1995. In addition to examining the temporal effect, we considered the potential influence of sexual dimorphism and, where relevant, included latitude and altitude as potential drivers of body size change. Contrary to initial predictions of a widespread increase in size, our findings challenge this assumption, revealing a nuanced interplay of factors contributing to the complexity of bat body size dynamics. Specifically, only three species (Myotis daubentonii, Nyctalus leisleri, and Pipistrellus pygmaeus) out of the 15 exhibited a discernible increase in body size over the studied period, prompting a reassessment of bats as reliable indicators of climate change based on alterations in body size. Our investigation into influencing factors highlighted the significance of temperature-related variables, with latitude and altitude emerging as crucial drivers. In some cases, this mirrored patterns consistent with Bergmann's rule, revealing larger bats recorded at progressively higher latitudes (Plecotus auritus, Myotis mystacinus, and Miniopterus schreibersii) or altitudes (Pipistrellus kuhlii). We also observed a clear sexual dimorphism effect in most species, with females consistently larger than males. The observed increase in size over time in three species suggests the occurrence of phenotypic plasticity, raising questions about potential long-term selective pressures on larger individuals. The unresolved question of whether temperature-related changes in body size reflect microevolutionary processes or phenotypic plastic responses adds further complexity to our understanding of body size patterns in bats over time and space.

Higher and bigger: How riparian bats react to climate change.

The altitudinal distribution of animals and changes in their body size are effective indicators of climate change. Bats are sensitive to climate change due to their dependence on temperature during critical life stages. However, long-term studies documenting responses over extended periods are rare. We present a 24-year investigation of Myotis daubentonii, a riparian bat known for altitudinal sexual segregation, along a river course in Central Italy. While males occupy the entire river course, females are confined to downstream warmer areas supporting successful reproduction due to improved foraging site productivity. In 2000, females were absent above 900 m a.s.l in our study area. We hypothesise that a) this altitude threshold is now higher, due to thermal gradient changes along the river course; and b) thermoregulatory costs for reproductive females have declined, leading to increased energy investment in offspring and subsequent generational growth in bat body size. Confirming our hypotheses, females exhibited a 175-m upward shift in altitude limit. Furthermore, we found a concurrent increase in body size (but not condition). Temperatures increased in the 24 years, likely allowing females to extend their range to higher elevations and favouring an increase in newborn body mass. Riparian vegetation remained unchanged, excluding habitat quality changes as the cause for the observed responses. The rapid female elevation rise might imply future disruption of established social structures, altering intra- and intersexual competition for roosts and food. Given the global decline in insect populations, larger bats might face future difficulties in finding food to sustain their body size, increasing mortality. However, the full impact of such changes on bat fitness remains unexplored and warrants further investigation, including other bat populations. This knowledge is crucial for informing conservation in the face of ongoing climate change and preserving the ecosystem services bats deliver in riparian ecosystems.

A First Assessment of SARS-CoV-2 Circulation in Bats of Central-Southern Italy.

One serious concern associated with the SARS-CoV-2 pandemic is that the virus might spill back from humans to wildlife, which would render some animal species reservoirs of the human virus. We assessed the potential circulation of SARS-CoV-2 caused by reverse infection from humans to bats, by performing bat surveillance from different sites in Central-Southern Italy. We restricted our survey to sampling techniques that are minimally invasive and can therefore be broadly applied by non-medical operators such as bat workers. We collected 240 droppings or saliva from 129 bats and tested them using specific and general primers for SARS-CoV-2 and coronaviruses, respectively. All samples (127 nasal swabs and 113 faecal droppings) were negative for SARS-CoV-2, and these results were confirmed by testing the samples with the Droplet Digital PCR. Additionally, pancoronavirus end-point RT-PCR was performed, and no sample showed specific bands. This outcome is a first step towards a better understanding of the reverse transmission of this virus to bats. Although the occurrence of a reverse zoonotic pattern can only be fully established by serological testing, the latter might represent an in-depth follow-up to a broad-scale preliminary assessment performed with our approach. We encourage the systematic surveillance of bats to help prevent reverse zoonotic episodes that would jeopardize human health, as well as biodiversity conservation and management.

Do We Need to Use Bats as Bioindicators?

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.

Artificial illumination influences niche segregation in bats.

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.

Detection and phylogenetic characterization of astroviruses in insectivorous bats from Central-Southern Italy.

In recent years, bats have been found to harbour many viruses, raising several questions about their role as reservoirs and potential disseminators of zoonotic viruses. We investigated the presence of six virus families in bats in three regions of Central-Southern Italy. Astroviruses were identified in seven of 13 bat species. Sequence analysis revealed marked genetic heterogeneity among the astroviruses identified, with nucleotide identity ranging between 60.26% and 87.62%. Astrovirus diversity was not associated with the bat species, the geographic areas or the bat colony, suggesting the circulation of several astrovirus strains in Italian ecosystems. Genetic diversification and interspecies transmission appear common in bat astroviruses and could provide, potentially, the bases for transmission to humans and other mammals. Yet overemphasizing this risk might have detrimental consequences for bat conservation and preservation of the important ecosystem services bats provide.

Sociality influences thermoregulation and roost switching in a forest bat using ephemeral roosts.

In summer, many temperate bat species use daytime torpor, but breeding females do so less to avoid interferences with reproduction. In forest-roosting bats, deep tree cavities buffer roost microclimate from abrupt temperature oscillations and facilitate thermoregulation. Forest bats also switch roosts frequently, so thermally suitable cavities may be limiting. We tested how barbastelle bats (Barbastella barbastellus), often roosting beneath flaking bark in snags, may thermoregulate successfully despite the unstable microclimate of their preferred cavities. We assessed thermoregulation patterns of bats roosting in trees in a beech forest of central Italy. Although all bats used torpor, females were more often normothermic. Cavities were poorly insulated, but social thermoregulation probably overcomes this problem. A model incorporating the presence of roost mates and group size explained thermoregulation patterns better than others based, respectively, on the location and structural characteristics of tree roosts and cavities, weather, or sex, reproductive or body condition. Homeothermy was recorded for all subjects, including nonreproductive females: This probably ensures availability of a warm roosting environment for nonvolant juveniles. Homeothermy may also represent a lifesaver for bats roosting beneath loose bark, very exposed to predators, because homeothermic bats may react quickly in case of emergency. We also found that barbastelle bats maintain group cohesion when switching roosts: This may accelerate roost occupation at the end of a night, quickly securing a stable microclimate in the newly occupied cavity. Overall, both thermoregulation and roost-switching patterns were satisfactorily explained as adaptations to a structurally and thermally labile roosting environment.

How to Be a Male at Different Elevations: Ecology of Intra-Sexual Segregation in the Trawling Bat Myotis daubentonii.

Intra-sexual segregation is a form of social segregation widespread among vertebrates. In the bat Myotis daubentonii, males are disproportionately abundant at higher elevations, while females are restricted to lower altitude. Intra-male segregation is also known to occur yet its ecological and behavioural determinants are unclear. We studied male segregation along a river in Central Italy where we tested the following predictions: 1. Upstream ( > 1000 m a.s.l.) males will rely on scarcer prey; 2. To deal with this limitation and exploit a cooler roosting environment, they will employ more prolonged and deeper torpor than downstream (< 900 m a.s.l.) males; 3. Body condition will be better in downstream males as they forage in more productive areas; 4. To cope with less predictable foraging opportunities, upstream males will use more habitat types. Consistent with our predictions, we found that prey were less common at higher altitudes, where bats exhibited prolonged and deeper torpor. Body condition was better in downstream males than in upstream males but not in all summer months. This result reflected a decrease in downstream males' body condition over the season, perhaps due to the energy costs of reduced opportunities to use torpor and/or intraspecific competition. Downstream males mainly foraged over selected riparian vegetation whereas upstream males used a greater variety of habitats. One controversial issue is whether upstream males are excluded from lower elevations by resident bats. We tested this by translocating 10 upstream males to a downstream roost: eight returned to the high elevation site in 1-2 nights, two persisted at low altitude but did not roost with resident bats. These results are consistent with the idea of segregation due to competition. Living at high altitude allows for more effective heterothermy and may thus be not detrimental for survival, but by staying at lower altitude males increase proximity to females and potentially benefit from summer mating opportunities.

What story does geographic separation of insular bats tell? A case study on Sardinian rhinolophids.

Competition may lead to changes in a species' environmental niche in areas of sympatry and shifts in the niche of weaker competitors to occupy areas where stronger ones are rarer. Although mainland Mediterranean (Rhinolophus euryale) and Mehely's (R. mehelyi) horseshoe bats mitigate competition by habitat partitioning, this may not be true on resource-limited systems such as islands. We hypothesize that Sardinian R. euryale (SAR) have a distinct ecological niche suited to persist in the south of Sardinia where R. mehelyi is rarer. Assuming that SAR originated from other Italian populations (PES)--mostly allopatric with R. mehelyi--once on Sardinia the former may have undergone niche displacement driven by R. mehelyi. Alternatively, its niche could have been inherited from a Maghrebian source population. We: a) generated Maxent Species Distribution Models (SDM) for Sardinian populations; b) calibrated a model with PES occurrences and projected it to Sardinia to see whether PES niche would increase R. euryale's sympatry with R. mehelyi; and c) tested for niche similarity between R. mehelyi and PES, PES and SAR, and R. mehelyi and SAR. Finally we predicted R. euryale's range in Northern Africa both in the present and during the Last Glacial Maximum (LGM) by calibrating SDMs respectively with SAR and PES occurrences and projecting them to the Maghreb. R. mehelyi and PES showed niche similarity potentially leading to competition. According to PES' niche, R. euryale would show a larger sympatry with R. mehelyi on Sardinia than according to SAR niche. Such niches have null similarity. The current and LGM Maghrebian ranges of R. euryale were predicted to be wide according to SAR's niche, negligible according to PES' niche. SAR's niche allows R. euryale to persist where R. mehelyi is rarer and competition probably mild. Possible explanations may be competition-driven niche displacement or Maghrebian origin.

Sensory ecology of water detection by bats: a field experiment.

Bats face a great risk of dehydration, so sensory mechanisms for water recognition are crucial for their survival. In the laboratory, bats recognized any smooth horizontal surface as water because these provide analogous reflections of echolocation calls. We tested whether bats also approach smooth horizontal surfaces other than water to drink in nature by partly covering watering troughs used by hundreds of bats with a Perspex layer mimicking water. We aimed 1) to confirm that under natural conditions too bats mistake any horizontal smooth surface for water by testing this on large numbers of individuals from a range of species and 2) to assess the occurrence of learning effects. Eleven bat species mistook Perspex for water relying chiefly on echoacoustic information. Using black instead of transparent Perspex did not deter bats from attempting to drink. In Barbastella barbastellus no echolocation differences occurred between bats approaching the water and the Perspex surfaces respectively, confirming that bats perceive water and Perspex to be acoustically similar. The drinking attempt rates at the fake surface were often lower than those recorded in the laboratory: bats then either left the site or moved to the control water surface. This suggests that bats modified their behaviour as soon as the lack of drinking reward had overridden the influence of echoacoustic information. Regardless of which of two adjoining surfaces was covered, bats preferentially approached and attempted to drink from the first surface encountered, probably because they followed a common route, involving spatial memory and perhaps social coordination. Overall, although acoustic recognition itself is stereotyped and its importance in the drinking process overwhelming, our findings point at the role of experience in increasing behavioural flexibility under natural conditions.

Postglacial colonization of Europe by the barbastelle bat: agreement between molecular data and past predictive modelling.

The barbastelle (Barbastella barbastellus) is a rare forest bat with a wide distribution in Europe. Here, we combine results from the analysis of two mtDNA fragments with species distribution modelling to determine glacial refugia and postglacial colonization routes. We also investigated whether niche conservatism occurs in this species. Glacial refugia were identified in the three southern European peninsulas: Iberia, Italy and the Balkans. These latter two refugia played a major role in the postglacial colonization process, with their populations expanding to England and central Europe, respectively. Palaeo-distribution models predicted that suitable climatic conditions existed in the inferred refugia during the last glacial maximum (LGM). Nevertheless, the overlap between the current and the LGM distributions was almost inexistent in Italy and in the Balkans, meaning that B. barbastellus populations were forced to shift range between glacial and interglacial periods, a process that probably caused some local extinctions. In contrast, Iberian populations showed a 'refugia within refugium' pattern, with two unconnected areas containing stable populations (populations that subsisted during both glacial and interglacial phases). Moreover, the match between LGM models and the refugial areas determined by molecular analysis supported the hypothesis of niche conservatism in B. barbastellus. We argue that geographic patterns of genetic structuring, altogether with the modelling results, indicate the existence of four management units for conservation: Morocco, Iberia, Italy and UK, and Balkans and central Europe. In addition, all countries sampled possessed unique gene pools, thus stressing the need for the conservation of local populations.