Linking Activity of Common Pipistrelles, Pipistrellus Pipistrellus, in an Urbanised Area With a Nearby Mass Swarming Site

The Erňa cave, a mass winter hibernaculum and important swarming site of the common pipistrelle, Pipistrellus pipistrellus, is located in the Slovak Karst, near the Košice urban agglomeration in eastern Slovakia. Over the past two decades, the unusual behaviour of late summer or autumn accidental but abundant occurrences of this species have been observed in buildings (so-called invasions) in Košice. It has been hypothesised that these events are related to bats swarming and hibernating in the Erňa cave; however, causality has not been confirmed. We measured the relative activity of bats from the end of the breeding season through the invasions and autumn swarming prior to the onset of hibernation by recording their echolocation calls on car-based transects in order to find any spatial and temporal linkage between activity in the urban area and the swarming site. Over two years we recorded 6,253 sequences with echolocation calls of P. pipistrellus and 5,239 records of other bats along four transects totalling 7,121 km in length. Spatial pattern analysis found that the city agglomeration presented a local hotspot of the species’ activity, especially during the invasion season. Multivariate generalised additive modelling confirmed an increased density of records of P. pipistrellus between the urban area and the hibernaculum in the pre-hibernation season, whereas this pattern was not found to be consistent on the control transects near the city. Contrary to that, other bat species showed little variation in their activity between transects and seasons. The obtained results suggest that the relatively short geographical distance between the urban agglomeration and the large swarming site is likely a clue to the frequent city invasions of the species, although the role of the city as a hibernation area cannot be completely omitted.

Bats (Plecotus auritus) use contact calls for communication among roost mates

Communication between group members is mediated by a diverse range of signals. Contact calls are produced by many species of birds and mammals to maintain group cohesion and associations among individuals. Contact calls in bats are typically relatively low-frequency social calls, produced only for communication. However, echolocation calls (higher in frequency and used primarily for orientation and prey detection) can also facilitate interaction among individuals and location of conspecifics in the roost. We studied calling behaviour of brown long-eared bats (Plecotus auritus) during return to maternity roosts in response to playbacks of social and echolocation calls. We hypothesised that calling by conspecifics would elicit responses in colony members. Bat responses (inspection flights and social calls production) were significantly highest during social call and echolocation call playbacks than during noise (control) playbacks. We suggest that social calling in maternity roosts of brown long-eared bat evolved to maintain associations among roostmates, rather than to find roosts or roostmates, because this species is strongly faithful to roosts and the social groups and roosts are stable over time and space. Living in a stable social group requires recognition of group members and affiliation of social bonds with group members, features that may be mediated by vocal signals.

Discriminating predation attempt outcomes during natural foraging using the post-buzz pause in Japanese large-footed bat Myotis macrodactylus

Bats emit a series of echolocation calls with an increasing repetition rate (the terminal buzz), when attempting to capture prey. This is often used as an acoustic indicator of prey-capture attempts. However, because it is directly linked to foraging efficiency, predation success is a more useful measure than predation attempts in ecological research. The characteristics of echolocation calls that consistently signify predation success across different situations have not been identified. Due to additional influencing factors, identification of these characteristics is particularly challenging for wild bats foraging in their natural environment compared to those in flight chambers. This study documented the natural foraging behavior of wild Japanese large-footed bat Myotis macrodactylus using synchronized acoustic and video recordings. From the video recordings, we could assign 137 attacks to three outcome categories: prey captured (51.8%), prey dropped (29.2%), and failed attempt (19%). Based on previous indications from laboratory studies that the length of the silent interval following the terminal buzz (post-buzz pause) might reflect the prey capture outcome, we compared post-buzz pause durations among categories of attack outcomes. The post-buzz pause was longest in the case of successful capture, suggesting that the length of the post-buzz pause is a useful acoustic indicator of predation success during natural foraging in M. macrodactylus. Our finding will advance the study of bat foraging behavior using acoustic data, including estimations of foraging efficiency and analyses of feeding habitat quality.Summary statementWe investigated the natural foraging behavior of wild Myotis macrodactylus and found that the length of the post-buzz pause is a useful acoustic indicator of predation success.

Hearing sensitivity: An underlying mechanism for niche differentiation in gleaning bats

Tropical ecosystems are known for high species diversity. Adaptations permitting niche differentiation enable species to coexist. Historically, research focused primarily on morphological and behavioral adaptations for foraging, roosting, and other basic ecological factors. Another important factor, however, is differences in sensory capabilities. So far, studies mainly have focused on the output of behavioral strategies of predators and their prey preference. Understanding the coexistence of different foraging strategies, however, requires understanding underlying cognitive and neural mechanisms. In this study, we investigate hearing in bats and how it shapes bat species coexistence. We present the hearing thresholds and echolocation calls of 12 different gleaning bats from the ecologically diverse Phyllostomid family. We measured their auditory brainstem responses to assess their hearing sensitivity. The audiograms of these species had similar overall shapes but differed substantially for frequencies below 9 kHz and in the frequency range of their echolocation calls. Our results suggest that differences among bats in hearing abilities contribute to the diversity in foraging strategies of gleaning bats. We argue that differences in auditory sensitivity could be important mechanisms shaping diversity in sensory niches and coexistence of species.

Bat Calls of Britain and Europe

A comprehensive guide to the calls of the 44 species of bat currently known to occur in Europe. Following on from the popular British Bat Calls by Jon Russ, this new book draws on the expertise of more than forty specialist authors to substantially update all sections, further expanding the volume to include sound analysis and species identification of all European bats. Aimed at volunteers and professional alike, topics include the basics of sound, echolocation in bats, an introduction to acoustic communication, equipment used and call analysis. For each species, detailed information is given on distribution, emergence, flight and foraging behaviour, habitat, echolocation calls – including parameters of common measurements – and social calls. Calls are described for both heterodyne and time expansion/full spectrum systems. A simple but complete echolocation guide to all species is provided for beginners, allowing them to analyse call sequences and arrive at the most likely species or group. The book also includes access to a downloadable library of over 450 calls presented as sonograms in the species sections.

Record of bats and their echolocation calls from southern Dolakha, central Nepal

With 52 species, bats make up almost a quarter of all the mammal species in Nepal, and yet remains the least understudied group of mammals. Owing to its diverse geography and climate, more species of bats potentially occur in the country, and acoustic surveys could improve the knowledge of their ecology. So, a study was conducted in Sailung and Melung Rural Municipalities of Dolakha district of Nepal, with the objectives of assessing bat species richness and preparing digital records of their echolocation calls. Using mist-netting and roost survey during three periods in 2018 (late March, late May to early June and mid-November), 10 species of bats were recorded: Cynopterus sphinx, Lyroderma lyra, Rhinolophus ferrumequinum, Rhinolophus luctus, Rhinolophus pearsonii, Rhinolophus sinicus, Hipposideros armiger, Myotis formosus, Myotis sp. and Nyctalus noctula. Ten roosts (mostly caves) were located. Echolocation calls of six of these species were documented, including signals of three species described for the first time in Nepal. This study also reports the fifth record of Myotis formosus in Nepal and new locality record of the species after two decades. The echolocation calls documented hereafter may serve as a reference for species identification for non-invasive studies of bats.

Effects of Environmental Clutter on Synthesized Chiropteran Echolocation Signals in an Anechoic Chamber

Ultrasonic bat detectors are useful for research and monitoring purposes to assess occupancy and relative activity of bat communities. Environmental “clutter” such as tree boles and foliage can affect the recording quality and identification of bat echolocation calls collected using ultrasonic detectors. It can also affect the transmission of calls and recognition by bats when using acoustic lure devices to attract bats to mist-nets. Bat detectors are often placed in forests, yet automatic identification programs are trained on call libraries using echolocation passes recorded largely from open spaces. Research indicates that using clutter-recorded calls can increase classification accuracy for some bat species and decrease accuracy for others, but a detailed understanding of how clutter impacts the recording and identification of echolocation calls remains elusive. To clarify this, we experimentally investigated how two measures of clutter (i.e., total basal area and number of stems of simulated woody growth, as well as recording angle) affected the recording and classification of a synthesized echolocation signal under controlled conditions in an anechoic chamber. Recording angle (i.e., receiver position relative to emitter) significantly influenced the probability of correct classification and differed significantly for many of the call parameters measured. The probability of recording echo pulses was also a function of clutter but only for the detector angle at 0° from the emitter that could receive deflected pulses. Overall, the two clutter metrics were overshadowed by proximity and angle of the receiver to the sound source but some deviations from the synthesized call in terms of maximum, minimum, and mean frequency parameters were observed. Results from our work may aid efforts to better understand underlying environmental conditions that produce false-positive and -negative identifications for bat species of interest and how this could be used to adjust survey accuracy estimates. Our results also help pave the way for future research into the development of acoustic lure technology by exploring the effects of environmental clutter on ultrasound transmission.

Phylogenetic Patterns in Mouth Posture and Echolocation Emission Behavior of Phyllostomid Bats

While phyllostomid bats show an impressive range of feeding habits, most of them emit highly similar echolocation calls. Due to the presence of an often prominent noseleaf, it has long been assumed that all phyllostomids emit echolocation calls exclusively through the nostrils rather than through the mouth. However, photo evidence documents also phyllostomid bats flying with an opened mouth. We hypothesized that all phyllostomid species emit echolocation calls only through the nostrils and therefore fly consistently with a closed mouth, and that observations of an open mouth should be a rare and random behavior among individuals and species. Using a high-speed camera and standardized conditions in a flight cage, we screened 40 phyllostomid species. Behavior varied distinctly among the species and mouth posture shows a significant phylogenetic signal. Bats of the frugivorous subfamilies Rhinophyllinae and Carolliinae, the nectarivorous subfamilies Glossophaginae and Lonchophyllinae, and the sanguivorous subfamily Desmodontinae all flew consistently with open mouths. So did the animalivorous subfamilies Glyphonycterinae, Micronycterinae and Phyllostominae, with the notable exception of species in the omnivorous genus Phyllostomus, which consistently flew with mouths closed. Bats from the frugivorous subfamily Stenodermatinae also flew exclusively with closed mouths with the single exception of the genus Sturnira, which is the sister clade to all other stenodermatine species. Further, head position angles differed significantly between bats echolocating with their mouth closed and those echolocating with their mouths opened, with closed-mouth phyllostomids pointing only the nostrils in the direction of flight and open-mouth phyllostomids pointing both the nostrils and mouth gape in the direction of flight. Ancestral trait reconstruction showed that the open mouth mode is the ancestral state within the Phyllostomidae. Based on the observed behavioral differences, we suggest that phyllostomid bats are not all nasal emitters as previously thought and discuss possible reasons. Further experiments, such as selectively obstructing sound emission through nostrils or mouth, respectively, will be necessary to clarify the actual source, plasticity and ecological relevance of sound emission of phyllostomid bats flying with their mouths open.

Bat echolocation plasticity in allopatry: a call for caution in acoustic identification of Pipistrellus sp.

Animals modify their behaviours and interactions in response to changing environments. In bats, environmental adaptations are reflected in echolocation signalling that is used for navigation, foraging and communication. However, the extent and drivers of echolocation plasticity are not fully understood, hindering our identification of bat species with ultrasonic detectors, particularly for cryptic species with similar echolocation calls. We used a combination of DNA barcoding, intensive trapping, roost and emergence surveys and acoustic recording to study a widespread European cryptic species complex (Pipistrellus pipistrellus and Pipistrellus pygmaeus) to investigate whether sibling bat species could exhibit extreme echolocation plasticity in response to certain environmental conditions or behaviours. We found that P. pygmaeus occupied the acoustic niche of their absent congeneric species, producing calls with P. pipistrellus’ characteristic structure and peak frequencies and resulting in false positive acoustic records of that species. Echolocation frequency was significantly affected by the density of bats and by maternity rearing stage, with lower frequency calls emitted when there was a high density of flying bats, and by mothers while juveniles were non-volant. During roost emergence, 29% of calls had peak frequencies typical of P. pipistrellus, with calls as low as 44 kHz, lower than ever documented. We show that automatic and manual call classifiers fail to account for echolocation plasticity, misidentifying P. pygmaeus as P. pipistrellus. Our study raises a vital limitation of using only acoustic sampling in areas with high densities of a single species of a cryptic species pair, with important implications for bat monitoring. Significance statement Ultrasonic acoustic detectors are widely used in bat research to establish species inventories and monitor species activity through identification of echolocation calls, enabling new methods to study and understand this elusive understudied group of nocturnal mammals. However, echolocation call signalling in bats is intrinsically different to that of other taxa, serving a main function of navigation and foraging. This study demonstrates an extreme level of plasticity, showing large variation in call frequency and structure in different situations. We showcase the difficulty and limitation in using acoustic sampling alone for bat monitoring and the complications of setting parameters for species identification for manual and automatic call classifiers. Our observations of call frequency variation correlated with density and absence of congenerics provide novel insights of behavioural echolocation plasticity in bats.

Winter activity of boreal bats

Natural hibernation sites used by bats in areas that lack cave features have long remained unresolved. To investigate hibernation site selection and winter activity of boreal bats, we recorded bat calls using passive acoustic monitoring at 16 sites in South-Western Finland. These sites included four rock outcrops with crevices and cave features, three glacial erratics or boulder fields, three ancient shores, three root cellars and three control sites where we did not expect bats to be overwintering. Our results revealed echolocation calls of Eptesicus nilssonii, Plecotus auritus and Myotis sp. We recorded significantly more activity near rock outcrops compared to other habitats, excluding root cellars. We also found that ambient temperature had a positive effect on bat activity and found evidence that P. auritus may be using low barometric pressure as a proxy for suitable foraging conditions during the winter. Our results suggest that rock outcrops may be more important to bats than previously acknowledged, highlighting the need to take these sites in account in planning of conservation measures. Furthermore, our findings underline the suitability of using acoustic monitoring in homing on hibernation sites that are not otherwise accessible.