Information transfer about roosts in female Bechstein's bats: an experimental field study

Information transfer among group members is believed to play an important part in the evolution of coloniality in both birds and bats. Although information transfer has received much scientific interest, field studies using experiments to test the underlying hypotheses are rare. We used a field experiment to test if communally breeding female Bechstein's bats (Myotis bechsteinii) exchange information regarding novel roosts. We supplied a wild colony, comprising 17 adult females of known relatedness, with pairs of suitable and unsuitable roosts and monitored the arrival of individuals marked with transponders (PIT-tags) over 2 years. As expected with information transfer, significantly more naive females were recruited towards suitable than towards unsuitable roosts. We conclude that information transfer about roosts has two functions: (i) it generates communal knowledge of a large set of roosts; and (ii) it aids avoidance of colony fission during roost switching. Both functions seem important in Bechstein's bats, in which colonies depend on many day roosts and where colony members live together for many years.     

Environmental conditions, rather than season, determine torpor use and temperature selection in large mouse-eared bats (Myotis myotis)

We tested whether food availability, thermal environment and time of year affect torpor use and temperature selection in the large mouse-eared bat (Myotis myotis) in summer and winter. Food-deprived bats were torpid longer than bats offered food ad libitum. Bats placed in a gradient of low (0 degrees C-25 degrees C) ambient temperatures (T(a)) spent more time in torpor than bats in a gradient of high (7 degrees C-43 degrees C) T(a)'s. However, we did not observe seasonal variations in the use of torpor. Moreover, even when food deprived in winter, bats never entered prolonged torpor at T(a)'s characteristic of their natural hibernation. Instead, bats preferred shallow torpor at relatively high T(a), but they always maintained a difference between body and ambient temperatures of less than 2 degrees C. Calculations based on respirometric measurements of metabolic rate showed that food deprived bats spent less energy per unit of time in torpor than fed individuals, even when they entered torpor at higher T(a)'s. We conclude that T(a) likely serves as a signal of food availability and daily torpor is apparently an adaptation to unpredictable changes in food availability, such as its decrease in summer or its increase in winter. Thus, we interpret hibernation to be a second step in the evolution of heterothermy in bats, which allows survival in seasonal environments.     

Thermoregulation and roost selection by reproductive female big brown bats (Eptesicus fuscus) roosting in rock crevices

A free-ranging maternity colony of big brown bats Eptesicus fuscus roosting in rock crevices along the South Saskatchewan River in south-eastern Alberta, Canada, was studied to understand better the discrepancy that exists in the literature regarding torpor use by reproductive female bats. Using radio-telemetry, thermoregulatory patterns and roost microclimate were recorded for pregnant, lactating and post-lactating females. Relative torpor use is described in several ways: the proportion of days on which torpor was used, depth, minimum body temperature, time spent in torpor, and a comprehensive torpor unit (degree-min). Pregnant and lactating female E. fuscus used torpor to the same extent overall (degree-min), but pregnant bats used torpor less frequently and with more time in deep torpor. Torpor was used to the greatest extent after weaning (post-lactation). Evidence is presented that the cost:benefit ratio for deep and prolonged periods of torpor may be highest during lactation. Microclimates of rock-crevice roosts mirrored the use of torpor throughout reproduction by bats. Lactation roosts (deeper, larger opening size) were more thermally stable and remained warmer at night compared to the shallow roosts used by pregnant and post-lactating females. It is shown that conclusions about relative use of torpor can differ depending on the units of comparison, necessitating measurement of all aspects of torpor (depth, duration and frequency). Comprehensive measurements, individual-based normothermic temperatures, and a definition of torpor that accounts for all energy savings, allow a more accurate depiction of patterns and facilitates inter-study comparisons.     

Social interactions among wild female Bechstein's bats (<Emphasis Type="Italic">Myotis bechsteinii</Emphasis>) living in a maternity colony

Although sociality is common in bats, few studies have investigated individual social behaviour in free-ranging colonies. This study quantifies social interactions among wild female Bechstein's bats (Myotis bechsteinii) belonging to one maternity colony. Our main goal was to analyse allogrooming and nose rubbing, which are both regularly displayed by adult females. Based on data of individually marked bats with known degrees of pairwise relatedness, we suggest that allogrooming has both a social and a hygienic function. Females groomed colony mates mainly on parts of the body that are difficult to reach by a bat itself. Thus, allogrooming may function to remove ectoparasites from inaccessible body parts. Allogrooming was rare compared to self-grooming (on average 0.7% vs 37.7% of a female's total observation time), and there was no significant correlation between the rate at which a bat groomed itself and the frequency with which it was groomed by conspecifics. Therefore, we assume that allogrooming also has a social purpose in addition to its assumed hygienic function. We suggest that allogrooming could strengthen social bonds among colony members that live together for many years. Mothers and adult daughters groomed each other preferentially. Thus, allogrooming may reflect special mother–daughter bonds. Nose rubbing occurred mainly within minutes (median: 80 s) after the arrival of a female in a night roost, and there was no correlation with relatedness. Therefore, it probably allows recognition of colony mates and may also be a greeting behaviour. Communicated by M.E. dos Santos     

Plasma progesterone concentrations in female little brown bats (Myotis lucifugus) during hibernation

Monthly collections of hibernating little brown bats contained (1) nulliparous females with small uteri and no antral follicles, (2) nulliparous females with swollen uteri and mature follicles, and (3) parous females, which, despite obvious differences in reproductive status, had equivalent plasma progesterone values. During the principal study season, mean monthly progesterone concentrations (measured by radioimmunoassay) showed recurrent increases with an apparent periodicity of about 60 days, but limited data obtained in the subsequent season did not. However, comparison of activity patterns in the two seasons with monthly progesterone concentrations suggests that ovarian activity during hibernation is affected by variations in metabolic level. We saw no evidence that nulliparous bats with small uteri developed antral follicles during hibernation. Despite their apparent immaturity, however, they had cornified vaginae and most were demonstrably inseminated. These indications of oestrus and the lack of differences between their plasma progesterone concentrations and those of patently mature females suggest that they were physiologically post-pubertal but failed to complete folliculogenesis before entering hibernation.     

Resource partitioning in two species of vespertilionid bats (Chiroptera) occupying the same roost

Colonies of Plecotus auritus (L.) and Myotis daubentoni (Kuhl) occupying the same roost showed no interspecific interaction. They utilized separate areas of the roost, departed from and returned to it at different times and foraged in different habitats on different groups of insects. P. auritus emerged from the roost before dusk, foraged singly in woodland areas with slow, hovering flight and fed on insects commonly found flying in woodland, on insects and spiders gleaned from foliage and on insects caught inside the roost. M. daubentoni emerged from the roost after dusk, flew fast and low over water and around riparian vegetation and foraged on long, straight beats. They frequently foraged in groups, and fed opportunistically on insects which fly in swarms in riparian habitats. The diets of the two species occupying the same roost did not differ from those established in areas where each species was found alone.     

Sperm-epithelium relationships in relation to the time of insemination in little brown bats (Myotis lucifugus)

Copulation lasted for up to 46 min in little brown bats. Spermatozoa were stored in both the uterus and the utero-tubal junction, although intimate relationships between spermatozoa and the epithelium were particularly evident in the utero-tubal junction, and were established at the beginning of the period of sperm storage. Polymorphonuclear leucocytes were present in all uteri irrespective of whether or not they had been inseminated but were not generally present in the utero-tubal junction or oviduct. Engulfment of spermatozoa by the epithelial cells of the utero-tubal junction and by polymorphonuclear leucocytes in the uterine glands was evident soon after copulation. It is suggested that this may effect the removal of defective spermatozoa and allow luminal spermatozoa access to the spatially restricted storage sites. Uninseminated female bats attempted to elicit copulation from torpid males, and were also observed adjacent to copulating pairs. Female bats also uttered copulation calls.     

Energy, water balance and the roost microenvironment in three Australian cave-dwelling bats (Microchiroptera)

The ghost bat, Macroderma gigas, and the orange leaf-nosed bat, Rhinonycteris aurantius, occupy similar ranges across northern Australia and are often found in the same roost caves. Both species are considered rare and vulnerable to further population decline. A third small species, the large bent-wing bat, Miniopterus schreibersii, has a similar body mass to R. aurantius, but has one of the largest ranges of any Australian mammal. In the present study we examine the effect and sensitivity of the animals' roosting microclimates on their energy and water balance. M. schreibersii exhibits a basal metabolic rate about 40% greater than other bats of similar body mass, whereas the other two species are close to predicted levels. R. aurantius shows a decrease in body temperatures below thermoneutrality. R. aurantius has levels of pulmocutaneous water loss among the highest seen for a mammal, and calculations based on nasal tip temperatures suggest that most of this loss is across the skin. Calculated ambient temperatures at which metabolic water production is equal to pulmocutaneous water loss in dry air are −14.7 °C for R. aurantius, 9.8 °C for M. schreibersii and −0.3 °C for M. gigas. Exposing the animals to relative humidities of between 80% and 90% shifted these calculated temperatures to 5.6 °C, 25.2 °C, and 2.9 °C, respectively. For each species the ratio of metabolic water production to evaporative water loss has been treated as a joint function of humidity and ambient temperature. The resulting surface plot shows that under known roosting conditions in caves R. aurantius and M. schreibersii remain in positive water balance, whereas M. gigas does not. Accepted: 20 May 2000     

Fecundity of Thominx neopulchra (Nematoda:Capillariidae) from bats of the genus Myotis (Chiroptera:Vespertilionidae)

Variability of absolute fecundity of nematode Thominx neopulchra from three species of genus Myotis was studies on the territory Zhiguli State Reserve in 2007, 2008. Significant differences in the fecundity of the nematode females depending on host species and sex, size of the parasite, number of the helminthes in a given host, and season of year were revealed.     

Eating local: influences of habitat on the diet of little brown bats (Myotis lucifugus)

We employ molecular methods to profile the diet of the little brown bat, Myotis lucifugus, and describe spatial and temporal changes in diet over their maternity season. We identified 61 prey species of insects and 5 species of arachnid. The largest proportion of prey (～32%) were identified as species of the mass-emerging Ephemeroptera (mayfly) genus Caenis. Bats roosting in agricultural settings had lower dietary richness than those occupying a roost located on a forest fragment in a conservation area. We detected temporal fluctuations in diet over the maternity season. Dipteran (fly) species dominated the diet early in the season, replaced later by species of mayfly. Because our methodology provides species-level identification of prey, we were able to isolate environmental indicator species in the diet and draw conclusions about the location and type of their foraging habitat and the health of these aquatic systems. The species detected suggested that the bats use variable habitats; members of one agricultural roost foraged on insects originating in rivers or streams while those in another agricultural roost and the forest roost fed on insects from pond or lake environments. All source water for prey was of fair to good quality, though no species detected are intolerant of pollution thus the habitat cannot be classified as pristine. Our study outlines a model system to investigate the abiotic and biotic interactions between habitat factors through this simple food chain to the top predator.     

Hemoglobins from bats (Myotis myotis and Rousettus aegyptiacus): a possible example of molecular adaptation to different physiological requirements

The functional properties of the hemoglobin systems from two different species of bat i.e. Rousettus aegyptiacus and Myotis myotis have been studied as a function of chloride, polyphosphates, pH and temperature. Apart from overall similarities shared with most mammalian hemoglobins, the two systems show significant differences with respect to the effect of chloride and temperature sensitivity. These findings have been related to the different physiological needs of the two species.     

Social behavior of water bats (Myotis daubentonii, Chiroptera) in an open-air cage

The social behavior of Myotis daubentonii bats was studied in experimental groups kept in an openair cage. The repertoire of identification, aggressive, and friendly (integration) behaviors was described. Analysis of their frequencies showed that interactions between the animals were for the most part friendly (61.6%), while aggressive displays were rare (4.6%). Differences in behavior depending on sex and age were revealed: avoidance of contact and aggressive vocalization were observed significantly more frequently in young and adult males than in females. The bats in the experimental groups personally distinguished each other; in group 4, a two-level hierarchy was established (a dominant male vs. all other bats), which could be explained by specific conditions in the open-air cage.     

Molecular systematics of bats of the genus Myotis (Vespertilionidae) suggests deterministic ecomorphological convergences.

Based on extensive phenetic analyses, bats of the genus Myotis have been classically subdivided into four major subgenera each of which comprise many species with similar morphological and ecological adaptations. Each subgenus thus corresponds to a distinct "ecomorph" encompassing bat species exploiting their environment in a similar fashion. As three of these subgenera are cosmopolitan, regional species assemblages of Myotis usually include sympatric representatives of each ecomorph. If species within these ecomorphs are monophyletic, such assemblages would suggest extensive secondary dispersal across geographic areas. Conversely, these ecomorphological adaptations may have evolved independently through deterministic processes, such as adaptive radiation. In this case, phylogenetic reconstructions are not expected to sort species of the same ecomorph into monophyletic clades. To test these predictions, we reconstructed the phylogenetic history of 13 American, 11 Palaearctic, and 6 other Myotis species, using sequence data obtained from nearly 2 kb of mitochondrial genes (cytochrome b and nd1). Separate or combined analyses of these sequences clearly demonstrate the existence of several pairs of morphologically very similar species (i.e., sibling species) which are phylogenetically not closely related. None of the three tested subgenera constitute monophyletic units. For instance, Nearctic and Neotropical species currently classified into the three subgenera were clustered in a single, well-supported monophyletic clade. These species thus evolved independently of their ecological equivalents from the Palaearctic region. Independent adaptive radiations among species of the genus Myotis therefore produced strikingly similar evolutionary solutions in different parts of the world. Furthermore, all phylogenetic reconstructions based on mtDNA strongly supported the existence of an unsuspected monophyletic clade which included all assayed New World species plus M. brandtii (from the Palaearctic Region). This "American" clade thus radiated into a morphologically diverse species assemblage which evolved after the first Myotis species colonized the Americas. Molecular reconstructions support paleontological evidence that species of the genus Myotis had a burst of diversification during the late Miocene-early Pliocene epoch.     

Roosting and foraging behaviour of Natterer's bats (Myotis nattereri) close to the northern border of their distribution

Seven nursery roosts and four roosts of male Myotis nattereri , Kuhl 1818 were found in central Scotland at latitude 56–57 N. Most were in crevices in the stonework of man-made structures other than occupied houses. Emergence occurred late in the evening, at an average light intensity of 3.5 lux and emerging bats circled in dark, sheltered areas outside roosts before departing along flyways towards foraging areas. Individuals departed from, and returned to, roosts in groups of 2-6, and circling behaviour was repeated on returning to the roost. During pregnancy, bats from anursery roost made one flight each per night. This increased to an average maximum of 1.84 early in lactation and then decreased again to one around weaning. Night roosts were situated in foraging areas and were used by M. nattereri for resting and grooming, for suckling volant but incompletely weaned young and also, possibly, for information transfer. Important foraging habitats were woodland edges, parkland, roadside vegetation and sheltered areas of water. Arthropod prey was captured both on the wing and by gleaning from foliage, and the bats were able to vary their diet according to arthropod availability. Overall, important prey included Diptera (both Nematocera and higher flies), Trichoptera, Coleoptera and non-flying groups such as Hemiptera, Dermaptera, Arachnida and Opiliones.     

Post-prandial urine loss and its relation to ecology in brown long-eared (Plecotus auritus) and Daubenton's (Myotis daubentoni) bats (Chiroptera: Vespertilionidae)

Urine loss, over the first 12 hours after feeding, was positively and linearly dependent on food consumption in water-denied, brown long-eared bats ( Plecotus auritus ) and also in water-denied and water-provided, Daubenton's bats ( Myotis daubentoni ). The slope of the relationship (food-dependent urine loss) (363 μ 1 g dry mass food⁻¹, S.D.=±70, n=19 ) was not significantly different between the two species but predicted urine loss at zero food consumption (food-independent urine loss) was significantly lower in P. auritus (0.048 μl.min⁻¹, S.D. =±0.015, n= 12) than in M. duubentoni (0.217 μl min⁻¹ S.D. =±0.040, n = 7 ). The same results were apparent if the data for M. daubentoni were restricted to water-deprived animals only. Of total urine loss, 46% occurred in the first hour after feeding in M. daubentoni compared with only 20% in P. auritus . We suggest that the differences between the two species in the pattern of postprandial urine loss reflect their relative association with open water when foraging and roosting in the wild. In the course of the water-denied experiment, M. duubentoni lost 15% of pre-fed body mass and showed signs of severe dehydration, while P. auritus only lost 6% and did not. However, urine loss only accounted for 8–10% of body mass loss. A water budget model for wild P. auritus in the summer was developed and suggested that if bats did not drink, approximately 19% of water loss would be attributable to faecal water loss, 18–20% to urine loss, and 59–62% of intake would be available to support evaporation and reproductive losses.     

Causes and consequences of living in closed societies: lessons from a long-term socio-genetic study on Bechstein's bats

Understanding the ecological, behavioural and genetic factors influencing animal social systems is crucial to investigating the evolution of sociality. Despite the recent advances in population genetic methods and the analysis of social interactions, long-term studies exploring the causes and consequences of social systems in wild mammals are rare. Here, we provide a synthesis of 15?years of data on the Bechstein's bat (Myotis bechsteinii), a species that raises its young in closed societies of 10-45 females living together for their entire lives and where immigration is virtually absent. We discuss the potential causes and consequences of living in closed societies, based on the available data on Bechstein's bat and other species with similar social systems. Using a combination of observational and genetic data on the bats together with genetic data on an ecto-parasite, we suggest that closed societies in Bechstein's bats are likely caused by a combination of benefits from cooperation with familiar colony members and parasite pressure. Consequences of this peculiar social system include increased sensitivity to demographic fluctuations and limits to dispersal during colony foundation, which have broad implications for conservation. We also hope to illustrate by synthesizing the results of this long-term study the diversity of tools that can be applied to hypothesize about the factors influencing a species' social system. We are convinced that with the expansion of the number of social mammals for which comparably detailed socio-genetic long-term data are available, future comparative studies will provide deeper insights into the evolution of closed societies.