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Map indicating the location of Nový Hrádek composed of three ruin complexes — Old and New Castle and ruins of old stables. Numbers (1–10) denote recording sites, diamonds denote hibernacula in cellars
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In comparison with Pipistrellus pipistrellus, very little is known about the mating behaviour and hibernation of P. pygmaeus. The phenology of display and swarming behaviour were studied using bat detectors and mist nettings in the ruins of Nový Hrádek castle (southwestern Moravia, Czech Republic), where P. pipistrellus and P. pygmaeus hibernate in...
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... the ruins of Nový Hrádek, 10 stationary sites were selected. All sites were on open passages separated from each other by high stone walls (Fig. 1). The night was divided into thirds (c.f., McAney and Fairley, 1988). Monitoring lasted five minutes at each stationary site every third of the ...
Citations
... To outline the general distribution of the species outside Eastern Europe and the Caucasus, we used georeferenced datasets of genetically identified samples provided by Mayer and von Helversen (2001), Racey et al. (2007), Kaňuch et al. (2007Kaňuch et al. ( , 2010, Bryja et al. (2009), Hulva et al. (2010, Kruskop et al. (2012), Sztencel-Jabłonka and Bogdanowicz (2012), Çoraman et al. (2013), Boston et al. (2014), Bogdanowicz et al. (2015), Bartoničková et al. (2016), Nusová et al. (2017), Montauban et al. (2021). In addition, we performed a selective mapping of published mentions of the species using the same approach as described above. ...
... Hulva et al. (2010) also pointed out the absence of winter observations of Pipistrelus pygmaeus in Central Europe. According to the available data, there is only one reliable winter record of the species, from the Czech Republic (Kaňuch et al. 2010, Bartoničková et al. 2016). The aforementioned records may testify to the expansion of Pipistrelus pygmaeus winter range and attempts of the species to hibernate in northern regions. ...
Knowledge of species distribution and, for migratory species, seasonal occurrence is particularly important for vulnerable and protected animals such as bats. The former European bat species Pipistrellus pipistrellus was split into two, Pipistrelus pipistrellus s.s. and Pipistrelus pygmaeus , over 20 years ago. However, their distribution, breeding and winter ranges as well as migratory status in Eastern Europe and the Caucasus remained obscure.
In our study, we scrutinised records of both species and assessed the sex ratio during their breeding season in this region.
We show that Pipistrellus pipistrellus s.s. has a significantly smaller range than previously assumed, being restricted to the southern part of the study area. On the contrary, Pipistrelus pygmaeus has a broader distribution, covering the Caucasus and Eastern Europe, up to ca. 59° N. Hence, all historical records and long‐distance movements of Pipistrellus pipistrellus sensu lato from Eastern Europe within the inferred allopatric occurrence of Pipistrelus pygmaeus should be attributed to Pipistrelus pygmaeus .
The high portion of adult males in breeding localities, mass wintering within the breeding range and lacking evidence of long‐distance seasonal movements imply that Pipistrellus pipistrellus s.s. is a sedentary species. In Pipistrelus pygmaeus , the sex ratio among adults during the breeding season varies latitudinally. The female‐biased sex ratio, absence of mass wintering and the historical long‐distance recoveries of ringed individuals suggest that this species is migratory in Eastern Europe. The male‐biased sex ratio during the breeding season, along with late autumn records, suggests that the Caucasus represents its winter range.
... Songflight calls are produced throughout the mating season, and their timing shifts to become progressively earlier as the season progresses (Sachteleben and von Helversen 2006;Bartoničková et al. 2016). The mean FMAXE for P. pipistrellus songflight social calls is 18 kHz (Sachteleben and von Helversen 2006;Barlow and Jones 1997a), compared with 21-22 kHz for P. pygmaeus (Pfalzer and Kusch 2003;Barlow and Jones 1997a). ...
... The mating period occurs from late summer into autumn with peaks in courtship behavior at the end of August and mid-November (Bartoničková et al. 2016;Park et al. 1998). It is possible that there are some geographical differences in the timing of mating, with mating groups forming and behavior being displayed in late summer in the UK (Park et al. 1996), compared with mating during or after seasonal movements in Central Europe (Sachteleben and von Helversen 2006). ...
... While patrolling courtship territories, males have a distinctive style of flight consisting of regular flapping interspersed with short periods of gliding, and are occasionally observed chasing competing males out of their territories (Sachteleben and von Helversen 2006). Courtship activity is at its highest in the earlier stages of the night, with activity peaking in August, and again in November, when more individuals begin to arrive at the hibernacula (Bartoničková et al. 2016). ...
This comprehensive species-specific chapter covers all aspects of the mammalian biology, including paleontology, physiology, genetics, reproduction and development, ecology, habitat, diet, mortality, and behavior. The economic significance and management of mammals and future challenges for research and conservation are addressed as well. The chapter includes a distribution map, a photograph of the animal, and a list of key literature.
... Songflight activity was highest between mid-August and mid-September close to a hibernaculum in the Czech Republic, though the bats probably mated away from the hibernaculum. At the start of the mating period, songflight activity is highest in the second third of the night, though peak activity is in the first third of the night after late September (Bartoničková et al. 2016). P. pygmaeus also emits single component Type C social calls with a median duration of 19 ms and median FMAXE of 31 kHz (Pfalzer and Kusch 2003). ...
... The main swarming activity in common pipistrelles at hibernacula is observed in early August (van Schaik et al. 2015;Bartoničková et al. 2016), and this behaviour is associated with frequent emission of social calls (e.g., Barlow and Jones 1997;Budenz et al. 2009;Middleton et al. 2014;Chaverri et al. 2018;Fig. 4 Predicted probability of the number of records of all bat species but P. pipistrellus at four transects (a, urban; b, link; c, control A; d, control B; locations see in Fig. 1) as a response to date, holding the outdoor temperature constant, derived from a generalised additive model. ...
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 so-called invasions of this species have been observed in buildings in Košice. This unusual behaviour occurs in late summer or autumn and it is characterized by numerous accidental or unforeseen occurrences of bats in various spaces of these houses. 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.
... Evidence that females are attracted was not presented. Bartonickova et al., (2016) reported that soprano pipistrelles behaved differently in a comparable situation. Lundberg (1990) reported that non-territorial males were never observed to perform a song flight. ...
... The main swarming activity in common pipistrelles at hibernacula is observed in early August (van Schaik et al. 2015, Bartoničková et al. 2016), and this behaviour is associated with frequent emission of social calls (e.g., Barlow In conclusion, invasions into the city could perhaps be considered as a twin swarming site of the same common pipistrelle population(s) related to a particular mass hibernaculum. In addition, such cities can serve as hibernation sites, too, where on warmer days bats can even nd a suitable amount of prey. ...
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.
... Although males of several species court females at known roost sites and may copulate during the daytime, the data are scant and we do not know the mating habits of many other species (Ortega and Arita 1999, Behr et al. 2004, Chaverri and Kunz 2006, Tan et al. 2009, Toth and Parsons 2013. Despite their secretive nature, several studies describe mating calls by males to court females or defend territories (Behr et al. 2004, Lin et al. 2015, Smotherman et al. 2016, Bartoničková et al. 2016. To collect acoustic data that can unequivocally assign specific behaviours (i.e., courtship or territoriality) to certain calls, it may be necessary to use synchronized audio and video in concert with directional microphones. ...
... Although males of several species court females at roosts and may copulate during the daytime, the data are scant, and we do not know the mating habits of most species (Ortega and Arita 1999;Behr et al. 2004;Chaverri and Kunz 2006;Tan et al. 2009;Toth and Parsons 2013). Several studies describe mating calls by males to court females or defend territories (Behr et al. 2004;Knörnschild et al. 2014;Lin et al. 2015;Smotherman et al. 2016;Bartoničková et al. 2016). To collect acoustic data that can be used to unequivocally assign specific behaviors (i.e., courtship or territoriality) to certain calls, it may be necessary to use synchronized audio and video in concert with directional microphones. ...
The Handbook summarizes all the key steps in conducting an acoustic survey of a bat community, including project planning, strategies for data collection, approaches to analysis and interpretation, a guide to purchasing a bat detector, and a series of case studies. Chapter 1 (“Introduction to bat echolocation”) provides a broad introduction to the theme, including a discussion of why and how bats echolocate, and a brief description of acoustic data, as well as what can be discerned about a bat community using acoustic techniques. Chapter 2 (“Acoustic survey design”) focuses on acoustic survey design, stressing the importance of identifying a clear research question and approach, and summarizing some of the most common questions that researchers investigate using acoustic techniques. Chapter 3 (“Bat detector choice and deployment”) discusses the difficult task of choosing the appropriate detector and summarizing the different technological approaches, as well as the trade-offs involved with selecting one style of detector over another. Chapter 4 (”Echolocation call identification”) focuses on strategies for identifying recordings of echolocation calls, starting with a discussion of the challenges associated with this task, an overview of both manual and automated approaches, and a section on using and creating call libraries, which is crucial for researchers working in areas where bat communities have received little or no study. Chapter 5 “Data, analysis, and inference”) deals with data management, analysis, and inference. It includes a discussion about strategies for data management that contains a section on the nature and use of databases. Furthermore, it describes different approaches to statistical analyses, many of which are intuitively linked to the suggestions for study design in Chapter 2. Chapters 2 through 5 each conclude with a “Some additional suggestions” section, which were sent to us when we asked a group of bat acoustic experts what they considered to be some common pitfalls associated with the technique. The final chapter of the Handbook (“Case studies”) includes five case studies, each of which summarizes a previously published study or studies that used acoustic survey techniques. The goal of this section is to demonstrate how many of the principles discussed throughout the Handbook have been applied in real-life scenarios. We selected the case studies to provide examples from a range of geographic locations, using various detecting technologies, and asking diverse questions about bat communities. Throughout the Handbook, when photos or recordings of individual species are provided and labeled, we have identified the species of interest by scientific name and by the common name provided by the online resource, Bats of the World: A Taxonomic and Geographic Database (Simmons and Cirranello 2020), unless stated otherwise.
On behalf of all contributors to the Handbook, we hope that this guide will help demystify the process of eavesdropping on bats and promote high standards in future acoustic studies of bat activity.
Erin Fraser, Alex Silvis, Mark Brigham, and Zenon Czenze
... If one considers an equal sex ratio at the population level, the following explanation may account for the difference in sex ratio at the swarming site. First, we found that males are present or visit the site more often or for a much longer period than females, a result consistent with a previous study in the Czech Republic (Bartoničková et al., 2016). Until midsum mer, adult females are reproducing in nurseries and joined the swarming site only after having weaned their pup. ...
The annual life cycle of temperate bats is typically accompanied by seasonal movements. In autumn, individuals of several species display very active flying behaviour around preferred locations, often hibernacula, a behaviour called swarming. This behaviour is usually characterized by a strong male-biased sex ratio and is often considered to be related to mating. Alternatively, these groupings may be a social behaviour related to the location and quality assessment of hibernacula and may enable this information to be transferred to juveniles, the maternal guidance hypothesis. Our study reports the results of a weekly survey of a bat community in an abandoned mine in Switzerland from April to October 2009, a period longer than the swarming period. The site is characterized by numerous visiting species as well as a high number of common pipistrelle bats displaying autumnal swarming like behaviour at the entrance of the mines. Thirteen species were present at the site with a total of 1,589 individuals of which 1,404 (88%) were common pipistrelle bats allowing us to focus on intersexual and age variations in their arrival phenology. Although the recapture rate was too low to obtain a reliable population size estimator, our data suggest that the site attracts individuals from a large area, possibly from several different nursing colonies and therefore requires a national conservation status. Adult males were present at the site throughout the study period with a peak from mid-July to the end of August. Females arrived later and both sexes displayed a swarming like behaviour in July and August. During this period, the sex ratio was slightly female biased, an unexpected pattern when compared with other swarming species. The absence of a male-biased sex ratio suggests that swarming behaviour in pipistrelle bats may have functions other than mating. Juveniles of both sexes joined the site about two weeks later suggesting that the maternal guidance hypothesis also did not explain this phenology pattern. The number of days between recaptures was greater for males than females indicating that males stay longer or return more often to the sites than females. Further studies may bring insights to the understanding of the function of swarming in pipistrelle bats.
... Individual bats may visit several swarming sites during a given season (Glover and Altringham 2008). At underground sites with large swarming activity, conspecifics from different breeding colonies may meet and search for unrelated mating partners or they may inform subadults about suitable hibernation sites (e.g., Thomas et al. 1979;Kerth et al. 2003;Rivers et al. 2006;Furmankiewicz 2008; Bartoničková et al. 2016). Thus, the main purpose of large swarming sites seems to be to facilitate communication across familiar and unfamiliar Handling editor: Danilo Russo. ...
Mass swarming of tens of thousands of common pipistrelles in front of hibernacula of the Carpathian Mountains suggests that bats may originate from a large catchment area. However, until now neither banding nor molecular data have resolved the geographic origin of common pipistrelles at these sites. Here, we measured the acoustic activity of bats and the stable hydrogen isotope ratios (δ2H) in metabolically inert fur keratin of bats to infer the relative swarming activity and the putative summer origin of bats, respectively, observed in autumn at Erňa cave, one of the largest bat hibernacula in Europe. Swarming activity declined with decreasing ambient temperature during the early season, while it increased during colder days towards the onset of hibernation. Based on δ2H values, we deduced that about 50% of the animals did not have a local origin. Provenance of all but one of these migrants was identified as the Pannonian Basin, while a single long-distance migrant may have originated from the northern margin of the species’ European distribution range. Modelling the variation in δ2H values of bats in response to sex, body condition and season suggested that, towards the onset of hibernation, males of low body condition were likely to be of distant geographic origin. Throughout the swarming season, females were mostly of local origin, yet towards the onset of hibernation, their body condition was not as variable as in males. We conclude that common pipistrelles observed at this mass hibernacula site are facultative migrants, which may undertake long-distance seasonal movements occasionally. At our study site, common pipistrelles are more likely to include long-distance migrants because of the mass occurrence of this species at this large hibernaculum.
