Activity Patterns of Two Syntopic and Closely Related Aerial-Hawking Bat Species During Breeding Season in Białowieża Primaeval Forest
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Mamm Res DOI 10.1007/s13364-016-0298-5 ORIGINAL PAPER Activity patterns of two syntopic and closely related aerial-hawking bat species during breeding season in Białowieża Primaeval Forest I. Ruczyński1 & P. Zahorowicz1 & T. Borowik1 & Z. Hałat1 Received: 7 July 2016 /Accepted: 14 October 2016 # The Author(s) 2016. This article is published with open access at Springerlink.com Abstract Temporal and spatial activity of bats is species specif- Keywords Nyctalus noctula . Nyctalus leisleri . ic and shaped by many factors such as energy requirements, Reproduction . Temperature . Niche segregation . Species climate conditions and food distribution. Pregnancy and lactation competition are the most energy-demanding periods throughout the female life cycle. During these periods, females have to optimize their activity patterns to maximize foraging success; however, they simultaneously need to take care of their young. In addition, daily Introduction and seasonal fluctuations of insect availability strongly affect bat foraging activity. If females, which are under strong energy con- Bats are the only mammals capable of active flight. The ability to straints, belong to closely related species, they may potentially fly allows bats, especially aerial insectivorous species, to cover suffer from competition. One of the mechanisms that allows large distances during the night and decrease risk of predation them to avoid competition is temporal and spatial niche (Rydell et al. 1996). Although flight ability is beneficial from an partitioning. Noctule and Leisler’s bats are closely related evolutionary perspective, it is also a time- and energy-consuming forest-dwelling species whose diet is similar and consists mainly process (Norberg and Rayner 1987; Muijres et al. 2012). Thus, of ephemeral insects. The aim of our study was to test if they the pattern of daily flight activity should maximize energy intake exhibit similar patterns in relation to the time and duration of their and minimize the costs of flight and predation risk (Rydell et al. nocturnal activity. In Białowieża Forest, we demonstrated that 1996; Speakman and Thomas 2003; Voigt et al. 2010). Temporal female nocturnal activity of both noctule and Leisler’sbatswas and spatial patterns of activity of bats are shaped by many factors, shaped mostly by reproductive period and ambient temperature. such as temporal and spatial changes in food distribution and the We did not observe significant differences in the activity patterns energetic demands of individuals (Neuweiler 2000; Dietz and of the two noctule species, which suggests that physiological Kalko 2007; Barclay and Jacobs 2011). Because the diet of bats constraints connected with reproduction and environmental con- occurring in the temperate zone contains mainly small flying ditions affect these species in a similar way and outweigh the invertebrates, the activity of which is known to be reliant on competition between species. ambient temperature (Abdullah 1961;Taylor1963;Baleetal. 2002), bat activity is also highly dependent on climate and weather conditions (Clare et al. 2011; Frick et al. 2012;Clare Communicated by: Karol Zub et al. 2014). Night activity of females during the breeding season Electronic supplementary material The online version of this article is mainly connected with the necessity for foraging, parental care (doi:10.1007/s13364-016-0298-5) contains supplementary material, and roost finding (Neuweiler 2000). The most energy- which is available to authorized users. demanding periods for females are pregnancy and lactation (Kunz 1974; Anthony and Kunz 1977; Racey et al. 1987; * Z. Hałat [email protected] Barclay 1989). Females need to balance their time out of the roost, which is required to get enough food, with the necessity 1 Mammal Research Institute, Polish Academy of Sciences, for regularly returning to feed and warm their young (Rydell Waszkiewicza 1, 17-230 Białowieża, Poland 1993; Shiel and Fairley 1999; Vonhof and Betts 2010). Mamm Res Females of most bat species deal with the same constraints (Ruczyński and Bogdanowicz 2005;Ruczyński et al. 2007; relating to parental care; however, nocturnal activity patterns Ruczyński and Bogdanowicz 2008; Ruczyński and Bartoń in bats are highly species specific. Even in the same climate 2012) and foraging activity during pregnancy and lactation conditions and reproductive period, females belonging to dif- (Kronwitter 1988;Jones1995; Shiel and Fairley 1999;Shiel ferent species may exhibit different temporal activity patterns. et al. 1999;Kaňuch et al. 2005;Kaňuch 2007; Nad’o and The differences are mainly connected with the various food Kaňuch 2015) for both species, there was no comparative preferences and hunting strategies (Neuweiler 2000; Dietz study of these two species inhabiting the same forest area. et al. 2009). Bats that feed on ephemeral insects that occur Our study aims to compare the patterns of nocturnal activity in high abundance and unpredictably in space and time usu- of the two noctule bats in primaeval forest conditions. ally make only one short bout per night (Racey and Swift We hypothesized that in both species, the temporal forag- 1985; Brigham and Fenton 1991; Shiel et al. 1999; ing patterns, i.e. the duration and time of nocturnal activity Dechmann et al. 2009). Contrarily, bats whose food is contin- and number of flights, are shaped by the reproductive status of uously available throughout the whole night feed for a longer females. To test our hypotheses, we searched the bat roosts timeandmakemoreflights(Audet1990; Chruszcz and and monitored their emergences and nocturnal activity with Barclay 2003). radiotelemetry. We expected females of two Nyctalus species The theory of niche differentiation indicates that closely to (i) increase number of flights per night and their duration related sympatric species use the environment in a way that after parturitions and hasten the onset of emergence when allows them to reduce competition for resources. The separa- lactating, as well as (ii) lengthen their nocturnal activity in tion of resource utilization can be based on temporal or spatial higher ambient temperatures and (iii) exhibit similar activity partitioning (Kronfeld-Schor and Dayan 2003; Bergeson et al. patterns. 2013; Symes et al. 2013). To test if closely related insectivo- rous bat species exhibit similar patterns, we conducted re- search on two syntopic, noctule bat species: noctule bats (Nyctalus noctula) and Leisler’sbats(Nyctalus leisleri). The Study area, methods and material diets of both species consist mainly of ephemeral insects and their night activity lasts only a few hours. Majority of noctule Study area bat prey consists of swarm insects, i.e. Trichoptera, Diptera (mostly Chironomidae) and Lepidoptera. In spring and au- Our study was conducted in the Białowieża Primaeval Forest tumn, they also forage on mass emergences of large insects (BPF). This large temperate mixed lowland forest covers an from Hemiptera and Coleoptera (Beck 1995; Gloor et al. area of 1450 km2, partially in Poland (600 km2) and partially 1995; Mackenzie and Oxford 1995). Leisler’sbatshavesim- in Belarus. BPF in Poland (52° 45′ N, 23° 50′ E) is divided ilar food preferences and they forage predominantly on small- into BiałowieżaNationalPark(105km2) and managed forest. and medium-sized insects forming swarms. Their diet consists The study area (100 km2) covered well-preserved stands of primarily of Lepidoptera, Diptera (mainly Tipulidae), Białowieża National Park (BNP), as well as parts of the man- Trichoptera (Beck 1995; Shiel et al. 1998) and Hymenoptera aged forest with old-growth remnants (over 80–100 years). (Kaňuch et al. 2005). These two bat species are dependent on The climate is continental (Olszewski 1986), with the highest forest as they roost mainly in trees (Bihari 2004; Bogdanowicz average temperature in July (17.8 °C) and lowest in January and Ruprecht 2004; Gebhard and Bogdanowicz 2004). The (−4.7 °C). The daily temperature amplitude is high in spring noctule bat is widespread across its range and known to be one and summer. During the study period, the night’s duration of the most common forest-dwelling bats in Europe, unlike varied from 7:05–9:25 h. The five main forest types in this Leisler’s bat, which is rather rare apart from in Spain, southern area are deciduous forest (Quercus robur, Tilia cordata and France, Germany and the Balkans (Horàcek et al. 2000;Dietz Carpinus betulus), mixed deciduous forest (Picea abies, et al. 2009). Both species migrate seasonally from the south or Q. robur, T. cordata and C. betulus), black alder bog forest southwest to Białowieża Primaeval Forest, which is one of and streamside alder-ash forest (Alnus glutinosa and Fraxinus their important breeding sites (Bogdanowicz and Ruprecht excelsior), mixed coniferous forest (Pinus sylvestris, P. abies 2004; Gebhard and Bogdanowicz 2004;Kaňuch et al. and Q. robur) and coniferous forest (P. sylvestris and P, abies) 2004). Aggregations last from May until August (Ruczyńi sk (Faliński 1986; Bernadzki et al. 1998). The superabundance of and Bogdanowicz 2005). Breeding colonies usually comprise holes in this forest provides shelters for a wide variety of 20–60 individuals in noctule bats and 20–50 in Leisler’sbats animals; there are 16 secondary cavity nesting bird species (Bogdanowicz and Ruprecht 2004). In the study area, parturi- inhabiting BPF (Wesołowski 2007). The area is inhabited by tions in both species take place in mid-June and females wean 15