Winter activity of bats is common, yet poorly understood. Other studies suggest a relationship between winter activity and ambient temperature, particularly temperature at sunset.We recorded echolocation calls to determine correlates of hourly bat-activity in Dinosaur Provincial Park, Alberta, Canada. We documented bat activity in temperatures as low as -10.4°C.We observed big brown bats (Eptesicus fuscus (Palisot de Beauvois, 1796)) flying at colder temperatures than species of Myotis bats (genus Myotis (Kaup 1829). We show that temperature and wind are important predictors of winter activity by E. fuscus and Myotis, and that Myotis may also use changes in barometric pressure to cue activity. In the absence of foraging opportunity, we suggest these environmental factors relate to heat loss and thus the energetic cost of flight. To understand the energetic consequences of bat flight in cold temperatures, we estimated energy expenditure during winter flights of E. fuscus and Myotis lucifugus using species-specific parameters. We estimated that winter flight uses considerable fat stores and that flight thermogenesis could mitigate energetic costs by 20% or more. We also show that temperature-dependent interspecific differences in winter activity likely stem from differences between species in heat loss and potential for activity-thermoregulatory heat substitution.
IntroductionDuring hibernation, metabolic rate (MR) and body temperature (T b ), are decreased for days to weeks (Ruf and Geiser 2015). Hibernating animals save considerable energy and can survive on limited resources (i.e., body fat or cached food) for extended periods, up to months at a time (Humphries et al. 2003). Benefits of hibernation are obvious (Geiser 2004;Geiser and Brigham 2012) but there are associated costs, such as suppressed molecular synthesis (Lillegraven et al. 1987), ceased or delayed reproduction (Racey 1969; Barnes et al. 1986), and immunosuppression (Bouma et al. 2010). For these reasons and likely others, nearly all hibernating mammals arouse periodically (Willis 1982; French 1985), ostensibly to excrete metabolic wastes (Baumber et al. 1971), mount immune responses (Burton and Reichman 1999), mate (Thomas et al. 1979), eat (Humphries et al. 2001), and possibly drink (Thomas and Cloutier 1992;Thomas and Geiser 1997; Ben-Hamo et al. 2013).Much of what we know about arousals and winter activity of temperate-zone bats comes from research on caverniculous species. In eastern North America, bats often roost in large groups in sizeable, humid, thermally-stable hibernacula (Webb et al. 1996;Perry 2012). Most activities typical of arousals occur within the hibernacula; caves and mines allow flight within hibernacula and may contain open water sources and hibernating insects that can be consumed (Swanson and Evans 1936;Rysgaard 1942). Opportunistic mating is also possible with mixedsex groups typically found in cave hibernacula (Thomas et al. 1979).Bats are also active outside of hibernacula for reasons poorly understood. Early ...