High latitude northern bats (Eptesicus nilssonii) reveal adaptations to both high and low ambient temperatures.

Insectivorous bats at northern latitudes need to cope with long periods of no food for large parts of the year. Hence, bats which are resident at northern latitudes throughout the year will need to undergo a long hibernation season and a short reproductive season where foraging time is limited by extended daylight periods. Eptesicus nilssonii is the northernmost occurring bat species worldwide and hibernates locally when ambient temperatures (Ta) limit prey availability. Therefore, we investigated the energy spent maintaining normothermy at different Ta, as well as how much bats limit energy expenditure while in torpor. We found that, despite being exposed to Ta as low as 1.1°C, bats did not increase torpid metabolic rate, thus indicating that E. nilssonii can survive and hibernate at low ambient temperatures. Furthermore, we found a lower critical temperature (Tlc) of 27.8°C, which is lower than in most other vespertilionid bats, and we found no indication of any metabolic response to Ta up to 37.1°C. Interestingly, carbon dioxide production increased with increasing Ta above the Tlc, presumably caused by a release of retained CO2 in bats that remained in torpor for longer and aroused at Ta above the Tlc. Our results indicate that E. nilssonii can thermoconform at near-freezing Ta, and hence maintain longer torpor bouts with limited energy expenditure, yet also cope with high Ta when sun exposed in roosts during long summer days. These physiological traits are likely to enable the species to cope with ongoing and predicted climate change.

Species and reproductive status influence element concentrations in bat fur.

To assess the impact of increasing pollutant levels on wildlife, we measured chemical contaminant loads in bats with different habitat and dietary preferences. Samples were taken from the fur of bats (Eptesicus nilssonii, Myotis brandtii, Myotis mystacinus and Plecotus auritus) to measure concentrations of 55 elements by inductively coupled plasma mass spectrometry (ICP-MS). Variations in element concentrations between different bat groups (species, sex, reproductive status) were analysed with a focus on arsenic (As), mercury (Hg) and lead (Pb) as these are known to cause specific health concerns in wildlife. For M. brandtii we found the highest As concentrations, especially in lactating bats, with a maximum exceeding those from other studies where bats had compromised health. Whereas for M. mystacinus there was a negative correlation between body condition index (BCI) and As concentration, indicating a potential danger for bats in the study area. In M. mystacinus and M. brandtii Hg concentrations were higher for sixteen individuals than in other studies where bats suffered genotoxic effects, although median levels were still below this threshold. Lactating bats from P. auritus and M. brandtii had higher Hg concentrations than bats of other reproductive status, which could endanger offspring as Hg can be transferred through lactation. In females from M. mystacinus Pb concentrations were more than three times higher compared to males. There was also a negative correlation between Pb concentration and BCI, which could mean that Pb has an adverse effect on health. Although many other biotic and abiotic factors should be considered, some of the variations in element concentrations could be due to different behaviours (foraging, roosting, etc.) in the studied species. The high levels of chemical contamination in some of the bats in our study, particularly reproductive individuals, is of conservation concern as bats are important agents for insect control.

A species-level trait dataset of bats in Europe and beyond.

Knowledge of species' functional traits is essential for understanding biodiversity patterns, predicting the impacts of global environmental changes, and assessing the efficiency of conservation measures. Bats are major components of mammalian diversity and occupy a variety of ecological niches and geographic distributions. However, an extensive compilation of their functional traits and ecological attributes is still missing. Here we present EuroBaTrait 1.0, the most comprehensive and up-to-date trait dataset covering 47 European bat species. The dataset includes data on 118 traits including genetic composition, physiology, morphology, acoustic signature, climatic associations, foraging habitat, roost type, diet, spatial behaviour, life history, pathogens, phenology, and distribution. We compiled the bat trait data obtained from three main sources: (i) a systematic literature and dataset search, (ii) unpublished data from European bat experts, and (iii) observations from large-scale monitoring programs. EuroBaTrait is designed to provide an important data source for comparative and trait-based analyses at the species or community level. The dataset also exposes knowledge gaps in species, geographic and trait coverage, highlighting priorities for future data collection.

Determining the different phases of torpor from skin- or body temperature data in heterotherms.

Technological innovations have made heat-sensitive data-loggers smaller, more efficient and less expensive, which has led to a growing body of literature that measures the skin- or body temperatures of small animals in their natural environments. Studies of this type on heterothermic endotherms have prompted much debate regarding how to best define 'torpor' expressions from skin- or body temperature data alone. We propose a new quantitative method for defining torpor 'entries', 'arousals' and 'stable torpor periods' whilst comparing the results to 'torpor bout' durations identified using only the torpor cut-off method. By decomposing a torpor bout into 'entries', 'stable torpor periods', and 'active arousals', we avoid biases introduced by using strict threshold temperature values for the onset of torpor, thereby allowing better insight into individual use of torpor. We present our method as an easy-to-use function written in R-code, offering an un-biased and consistent methodology to be applied on skin- or body temperature measurements across datasets and research groups. When testing the function on a large dataset of skin temperature data collected on three bat species in Norway (Plecotus auritus: Nind = 39; Eptesicus nilssonii: Nind = 11; Myotis brandtii: Nind = 10), we identified 461 complete torpor bouts across species. More than 40% of the torpor bouts (Nbouts = 192) did not contain stable torpor periods, because the bats aroused before they had reached a stable skin temperature level. Furthermore, only considering 'torpid' and 'euthermic' temperature values by applying strict cut-off thresholds led to potentially large underestimations of torpor bout durations compared to our quantitative determination of the onset and termination of each torpor bout. We highlight the importance of differentiating between torpor phases, especially for active arousals that can be very energetically expensive and may alter our evaluation of the actual energetic savings gained by an individual employing torpor.

State dependence of arousal from torpor in brown long-eared bats (Plecotus auritus).

To cope with periods of low food availability and unsuitable environmental conditions (e.g., short photoperiod or challenging weather), many heterothermic mammals can readily go into torpor to save energy. However, torpor also entails several potential costs, and quantitative energetics can, therefore, be influenced by the individual state, such as available energy reserves. We studied the thermal energetics of brown long-eared bats (Plecotus auritus) in the northern part of its distributional range, including torpor entry, thermoregulatory ability during torpor and how they responded metabolically to an increasing ambient temperature (Ta) during arousal from torpor. Torpor entry occurred later in bats with higher body mass (Mb). During torpor, only 10 out of 21 bats increased oxygen consumption (V̇O2) to a greater extent above the mean torpor metabolic rates (TMR) when exposed to low Ta. The slope of the torpid thermoregulatory curve was shallower than that of resting metabolic rate (RMR) during normothermic conditions, indicating a higher thermal insulation during torpor. During exposure to an increasing Ta, all bats increased metabolic rate exponentially, but the bats with higher Mb aroused at a lower Ta than those with lower Mb. In bats with low Mb, arousal was postponed to an Ta above the lower critical temperature of the thermoneutral zone. Our results demonstrate that physiological traits, which are often considered fixed, can be more flexible than previously assumed and vary with individual state. Thus, future studies of thermal physiology should to a greater extent take individual state-dependent effects into account.

Universality of Torpor Expression in Bats.

AbstractAlthough heterothermy is employed by species at a global level within the order of Chiroptera (bats), the possibility of torpor being expressed in bat species inhabiting warmer climate zones has been explored only in the past couple decades. Recent studies suggest that the benefit of expressing torpor is not limited to saving energy during cold exposure or food shortage but may be just as important for saving water during heat waves. Thus, even if the physiological challenges faced by bats may depend on the habitat they live in, species expressing torpor should be found in any climate zone where employing torpor may yield benefits and increase their survival probability. Here, we summarize available data on torpor metabolic rates and daily skin temperature patterns of bats across climate zones, emphasizing similarities found in the data. We also present data that we have collected from a southern subtropical species (Nyctophilus bifax) and a northern subarctic species (Plecotus auritus) to illustrate specific examples of torpor expressions in two bat species living in highly different environments. Our findings highlight that torpor metabolic rates and skin temperature patterns of bats outside of the hibernation season can be universal across vastly different habitats, although arid environments indicate potential divergence in mean minimum torpor metabolic rates compared with measurements of populations inhabiting other climate zones.

Energetics of whiskered bats in comparison to other bats of the family Vespertilionidae.

Bats inhabit a variety of climate types, ranging from tropical to temperate zones, and environmental differences may therefore affect the basal metabolic rate (BMR) of bats from different populations. In the present study, we provide novel data on the energetics of whiskered bats (Myotis mystacinus), which is the smallest species within Chiroptera measured to date. We investigated the thermoregulatory strategies of M. mystacinus close to the northern limits of this species' distribution range and compared these data to other vespertilionid bats living in different climates. As mammals living in colder areas experience elevated thermoregulatory costs, often leading to an increase in BMR, we hypothesised that BMR of this northern population of whiskered bats would be higher than that of bats from climates with warm environmental temperatures. From a systematic literature search we obtained BMR estimates (N=47) from 24 species within Vespertilionidae. Our metabolic measurements of M. mystacinus in Norway (body mass of 4.4 g; BMR of 1.48 ml O2 g-1 h-1) were not different from other vespertilionid bats, based on the allometric equation obtained from the systematic literature search. Further, there was no effect of environmental temperature on BMR within Vespertilionidae. How these tiny bats adapt metabolically to high latitude living is thus still an open question. Bats do have a suite of physiological strategies used to cope with the varying climates which they inhabit, and one possible factor could be that instead of adjusting BMR they could express more torpor. This article has an associated First Person interview with the first author of the paper.