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Concept: Pipistrellus

166

Echolocating bats construct an auditory world sequentially by analyzing successive pulse-echo pairs. Many other mammals rely upon a visual world, acquired by sequential foveal fixations connected by visual gaze saccades. We investigated the scanning behavior of bats and compared it to visual scanning. We assumed that each pulse-echo pair evaluation corresponds to a foveal fixation and that sonar beam movements between pulses can be seen as acoustic gaze saccades. We used a two-dimensional 16 microphone array to determine the sonar beam direction of succeeding pulses and to characterize the three dimensional scanning behavior in the common pipistrelle bat (Pipistrellus pipistrellus) flying in the field. We also used variations of signal amplitude of single microphone recordings as indicator for scanning behavior in open space. We analyzed 33 flight sequences containing more than 700 echolocation calls to determine bat positions, source levels, and beam aiming. When searching for prey and orienting in space, bats moved their sonar beam in all directions, often alternately back and forth. They also produced sequences with irregular or no scanning movements. When approaching the array, the scanning movements were much smaller and the beam was moved over the array in small steps. Differences in the scanning pattern at various recording sites indicated that the scanning behavior depended on the echolocation task that was being performed. The scanning angles varied over a wide range and were often larger than the maximum angle measurable by our array. We found that echolocating bats use a “saccade and fixate” strategy similar to vision. Through the use of scanning movements, bats are capable of finding and exploring targets in a wide search cone centered along flight direction.

Concepts: Dimension, Sequence, Animal echolocation, Bat, Microbat, Common Pipistrelle, Pipistrellus, Soprano Pipistrelle

16

Wind turbines represent a source of hazard for bats, especially through collision with rotor blades. With increasing technical development, tall turbines (rotor-swept zone 50-150 m above ground level) are becoming widespread, yet we lack quantitative information about species active at these heights, which impedes proposing targeted mitigation recommendations for bat-friendly turbine operation. We investigated vertical activity profiles of a bat assemblage, and their relationships to wind speed, within a major valley of the European Alps where tall wind turbines are being deployed. To monitor bat activity we installed automatic recorders at sequentially increasing heights from ground level up to 65 m, with the goal to determine species-specific vertical activity profiles and to link them to wind speed. Bat call sequences were analysed with an automatic algorithm, paying particular attention to mouse-eared bats (Myotis myotis and Myotis blythii) and the European free-tailed bat (Tadarida teniotis), three locally rare species. The most often recorded bats were the Common pipistrelle (Pipistrellus pipistrellus) and Savi’s pipistrelle (Hypsugo savii). Mouse-eared bats were rarely recorded, and mostly just above ground, appearing out of risk of collision. T. teniotis had a more evenly distributed vertical activity profile, often being active at rotor level, but its activity at that height ceased above 5 ms-1 wind speed. Overall bat activity in the rotor-swept zone declined with increasing wind speed, dropping below 5% above 5.4 ms-1. Collision risk could be drastically reduced if nocturnal operation of tall wind turbines would be restricted to wind speeds above 5 ms-1. Such measure should be implemented year-round because T. teniotis remains active in winter. This operational restriction is likely to cause only small energy production losses at these tall wind turbines, although further analyses are needed to assess these losses precisely.

Concepts: Little brown bat, Wind power, Bats, Wind turbine, Common Pipistrelle, Pipistrellus, Soprano Pipistrelle, Mouse-eared bat

13

Conflict can arise when bats roost in human dwellings and householders are affected adversely by their presence. In the United Kingdom, the exclusion of bats from roosts can be licensed under exceptional circumstances to alleviate conflict, but the fate of excluded bats and the impact on their survival and reproduction is not well understood. Using radio-tracking, we investigated the effects of exclusion on the soprano pipistrelle Pipistrellus pygmaeus, a species that commonly roosts in buildings in Europe. Exclusions were performed under licence at five roosts in England in spring, when females were in the early stages of pregnancy. Following exclusion, all bats found alternative roosts and colonies congregated in nearby known roosts that had been used by radio-tagged bats prior to exclusion. We found no difference in roosting behaviour before and after exclusion. Both the frequency of roost switching and the type of roosts used by bats remained unchanged. We also found no change in foraging behaviour. Bats foraged in the same areas, travelled similar distances to reach foraging areas and showed similar patterns of habitat selection before and after exclusion. Population modelling suggested that any reduction in survival following exclusion could have a negative impact on population growth, whereas a reduction in productivity would have less effect. While the number of soprano pipistrelle exclusions currently licensed each year is likely to have little effect on local populations, the cumulative impacts of licensing the destruction of large numbers of roosts may be of concern.

Concepts: Demography, Population, World population, License, Common Pipistrelle, Pipistrellus, Soprano Pipistrelle, Eastern Pipistrelle

10

Extinctions typically have ecological drivers, such as habitat loss. However, extinction events are also influenced by policy and management settings that may be antithetical to biodiversity conservation, inadequate to prevent extinction, insufficiently resourced, or poorly implemented. Three endemic Australian vertebrate species - the Christmas Island pipistrelle (Pipistrellus murrayi), Bramble Cay melomys (Melomys rubicola), and Christmas Island forest skink (Emoia nativitatis) - became extinct from 2009 to 2014. All 3 extinctions were predictable and probably preventable. We sought to identify the policy, management, research, and other shortcomings that contributed their extinctions or failed to prevent them. Factors that contributed to these extinctions included a lack within national environmental legislation and policy of explicit commitment to the prevention of avoidable extinctions, lack of explicit accountability, inadequate resources for conservation (particularly for species not considered charismatic or not of high taxonomic distinctiveness), inadequate biosecurity, a slow and inadequate process for listing species as threatened, recovery planning that failed to consider the need for emergency response, inability of researchers to identify major threatening factors, lack of public engagement and involvement in conservation decisions, and limited advocacy. From these 3 cases, we recommend environmental policy explicitly seek to prevent extinction of any species and provide a clear chain of accountability and an explicit requirement for public inquiry following any extinction; implementation of a timely and comprehensive process for listing species as threatened and for recovery planning; reservation alone not be assumed sufficient to maintain species; enhancement of biosecurity measures; allocation of sufficient resources to undertake actions necessary to prevent extinction; monitoring be considered a pivotal component of the conservation response; research provide timely identification of factors responsible for decline and of the risk of extinction; effective dissemination of research results; and advocacy by an informed public for the recovery of threatened species; and public involvement in governance of the recovery process. These recommendations should be applicable broadly to reduce the likelihood and incidence of extinctions. This article is protected by copyright. All rights reserved.

Concepts: Conservation biology, Evolution, Ecology, Endangered species, Extinction, Dinosaur, Pipistrellus, Christmas Island Pipistrelle

10

While the novel Middle East Respiratory Syndrome Coronavirus (MERS-CoV) is closely related to Tylonycteris bat CoV HKU4 (Ty-BatCoV HKU4) and Pipistrellus bat CoV HKU5 (Pi-BatCoV HKU5) in bats from Hong Kong, and other potential lineage C betacoronaviruses in bats from Africa, Europe and America, its animal origin remains obscure. To better understand the role of bats in its origin, we examined the molecular epidemiology and evolution of lineage C betacoronaviruses among bats. Ty-BatCoV HKU4 and Pi-BatCoV HKU5 were detected in 29% and 25% of alimentary samples from lesser bamboo bat (Tylonycteris pachypus) and Japanese pipistrelle (Pipistrellus abramus) respectively. Sequencing of their RdRp, S and N genes revealed that MERS-CoV is more closely related to Pi-BatCoV HKU5 in RdRp (92.1-92.3% aa identities) but to Ty-BatCoV HKU4 in S (66.8-67.4% aa identities) and N (71.9-72.3% aa identities). Although both viruses were under purifying selection, the S of Pi-BatCoV HKU5 displayed marked sequence polymorphisms and more positively selected sites than that of Ty-BatCoV HKU4, suggesting that Pi-BatCoV HKU5 may generate variants to occupy new ecological niches along with its host which faces diverse habitats. Molecular clock analysis showed that they diverged from a common ancestor with MERS-CoV at least several centuries ago. Although MERS-CoV may have diverged from potential lineage C betacoronaviruses in European bats more recently, these bat viruses were unlikely the direct ancestor of MERS-CoV. Intensive surveillance for lineage C betaCoVs in Pipistrellus and related bats with diverse habitats, and other animals from the Middle East may fill the evolutionary gap.

Concepts: Evolution, Severe acute respiratory syndrome, Middle East, Asia, Vesper bats, Pipistrellus, Lesser Bamboo Bat, Tylonycteris

5

Urbanisation and climate change are two global change processes that affect animal distributions, posing critical threats to biodiversity. Due to its versatile ecology and synurbic habits, Kuhl’s pipistrelle (Pipistrellus kuhlii) offers a unique opportunity to explore the relative effects of climate change and urbanisation on species distributions. In a climate change scenario, this typically Mediterranean species is expected to expand its range in response to increasing temperatures. We collected 25,132 high-resolution occurrence records from P. kuhlii European range between 1980 and 2013 and modelled the species' distribution with a multi-temporal approach, using three bioclimatic variables and one proxy of urbanisation. Temperature in the coldest quarter of the year was the most important factor predicting the presence of P. kuhlii and showed an increasing trend in the study period; mean annual precipitation and precipitation seasonality were also relevant, but to a lower extent. Although urbanisation increased in recently colonised areas, it had little effect on the species' presence predictability. P. kuhlii expanded its geographical range by about 394 % in the last four decades, a process that can be interpreted as a response to climate change.

Concepts: Biodiversity, Earth, Precipitation, Climate, Temperature, Monsoon, Pipistrellus, Kuhl's Pipistrelle

3

Bats are natural reservoirs for many mammalian coronaviruses, which have received renewed interest after the discovery of the severe acute respiratory syndrome (SARS) and the Middle East respiratory syndrome (MERS) CoV in humans. This study describes the identification and molecular characterization of alphacoronaviruses and betacoronaviruses in bats in Italy, from 2010 to 2012. Sixty-nine faecal samples and 126 carcasses were tested using pan-coronavirus RT-PCR. Coronavirus RNAs were detected in seven faecal samples and nine carcasses. A phylogenetic analysis of RNA-dependent RNA polymerase sequence fragments aided in identifying two alphacoronaviruses from Kuhl’s pipistrelle (Pipistrellus kuhlii), three clade 2b betacoronaviruses from lesser horseshoe bats (Rhinolophus hipposideros), and 10 clade 2c betacoronaviruses from Kuhl’s pipistrelle, common noctule (Nyctalus noctula), and Savi’s pipistrelle (Hypsugo savii). This study fills a substantive gap in the knowledge on bat-CoV ecology in Italy, and extends the current knowledge on clade 2c betacoronaviruses with new sequences obtained from bats that have not been previously described as hosts of these viruses.

Concepts: RNA, RNA interference, Severe acute respiratory syndrome, RNA-dependent RNA polymerase, Vesper bats, Pipistrellus, Nyctalus, Kuhl's Pipistrelle

1

Frequency shifts in signals of bats flying near conspecifics have been interpreted as a spectral jamming avoidance response (JAR). However, several prerequisites supporting a JAR hypothesis have not been controlled for in previous studies. We recorded flight and echolocation behavior of foraging Pipistrellus pipistrellus while flying alone and with a conspecific and tested whether frequency changes were due to a spectral JAR with an increased frequency difference, or whether changes could be explained by other reactions. P. pipistrellus reacted to conspecifics with a reduction of sound duration and often also pulse interval, accompanied by an increase in terminal frequency. This reaction is typical of behavioral situations where targets of interest have captured the bat’s attention and initiated a more detailed exploration. All observed frequency changes were predicted by the attention reaction alone, and do not support the JAR hypothesis of increased frequency separation. Reaction distances of 1-11 m suggest that the attention response may be elicited either by detection of the conspecific by short range active echolocation or by long range passive acoustic detection of echolocation calls.

Concepts: Scientific method, Psychology, Hypothesis, Ultrasound, Bat, Common Pipistrelle, Pipistrellus, Soprano Pipistrelle

1

Artificial lighting is a particular problem for animals active at night. Approximately 69% of mammal species are nocturnal, and one-third of these are bats. Due to their extensive movements-both on a nightly basis to exploit ephemeral food supplies, and during migration between roosts-bats have an unusually high probability of encountering artificial light in the landscape. This paper reviews the impacts of lighting on bats and their prey, exploring the direct and indirect consequences of lighting intensity and spectral composition. In addition, new data from large-scale surveys involving more than 265 000 bat calls at more than 600 locations in two countries are presented, showing that prevalent street-lighting types are not generally linked with increased activity of common and widespread bat species. Such bats, which are important to ecosystem function, are generally considered ‘light-attracted’ and likely to benefit from the insect congregations that form at lights. Leisler’s bat (Nyctalus leisleri) may be an exception, being more frequent in lit than dark transects. For common pipistrelle bats (Pipistrellus pipistrellus), lighting is negatively associated with their distribution on a landscape scale, but there may be local increases in habitats with good tree cover. Research is now needed on the impacts of sky glow and glare for bat navigation, and to explore the implications of lighting for habitat matrix permeability.

Concepts: Light, Mammal, Lighting, Bat, Light pollution, Vesper bats, Common Pipistrelle, Pipistrellus

1

In Germany, rabies in bats is a notifiable zoonotic disease, which is caused by European bat lyssaviruses type 1 and 2 (EBLV-1 and 2), and the recently discovered new lyssavirus species Bokeloh bat lyssavirus (BBLV). As the understanding of bat rabies in insectivorous bat species is limited, in addition to routine bat rabies diagnosis, an enhanced passive surveillance study, i.e. the retrospective investigation of dead bats that had not been tested for rabies, was initiated in 1998 to study the distribution, abundance and epidemiology of lyssavirus infections in bats from Germany. A total number of 5478 individuals representing 21 bat species within two families were included in this study. The Noctule bat (Nyctalus noctula) and the Common pipistrelle (Pipistrellus pipistrellus) represented the most specimens submitted. Of all investigated bats, 1.17% tested positive for lyssaviruses using the fluorescent antibody test (FAT). The vast majority of positive cases was identified as EBLV-1, predominately associated with the Serotine bat (Eptesicus serotinus). However, rabies cases in other species, i.e. Nathusius' pipistrelle bat (Pipistrellus nathusii), P. pipistrellus and Brown long-eared bat (Plecotus auritus) were also characterized as EBLV-1. In contrast, EBLV-2 was isolated from three Daubenton’s bats (Myotis daubentonii). These three cases contribute significantly to the understanding of EBLV-2 infections in Germany as only one case had been reported prior to this study. This enhanced passive surveillance indicated that besides known reservoir species, further bat species are affected by lyssavirus infections. Given the increasing diversity of lyssaviruses and bats as reservoir host species worldwide, lyssavirus positive specimens, i.e. both bat and virus need to be confirmed by molecular techniques.

Concepts: Bat, Rabies, Mammals of Africa, Mammals of Europe, Mammals of Asia, Vesper bats, Vesper bat, Pipistrellus