Suckling behaviour in the brown long-eared bat (Plecotus auritus)

Thirty-two adult female brown long-eard bats were taken into captivity. Eight individuals gave birth to single young in captivity (known mother-young pairs), 10 were lactating when captured (putative mother-young pairs), and the remaining 14 bats were non-reproductive. Bats were maintained in five groups consisting of females from sigle(n=3) or mixed (n=2) wild roosts. All bats were housed in outdoor, free-flight enclosures and fed mainly on free-flying noctuid moths. Bats were individually were determined daily (n=152) for a single gruup of bats containing four known mother-young pairs and five non-reproductive bats. The probability of a being attached to the nipple declined from 100% of records at 1-5 days of age to 5% of records at 36-40 days of age. Females were always found suckling their own young. Suckling associations were determined using infra-red sensitive video-recordings of bat behaviour within the roost box. For both known (n=8) and putative mother-young pairs (n=10), there were no records of young attached to lactating females other than their own mothers (from the same or different wild roosts).     

Food habits of northern (Eptesicus nilssoni) and brown long-eared (Plecotus auritus) bats in Sweden

Food habits of sympatric northern and brown long-eared bats were examined by analysis of droppings collected in six maternity roosts of each species. Relatively large, nocturnal flying insects such as moths (Lepidoptera), June-beetles (Coleoptera: Scarabaeidae), crane-flies (Diptera: Tipulidae) and caddis-flies (Trichoptera), were eaten by both bat species, representing 47% (by volume) of the northern and 57% of the long-eared bat's diet. Small dipterans were important food items for the northern bat (47% of the diet) but not for the long-eared bat, which instead consumed large amounts of diurnal or predominantly non-flying taxa such as blow-flies (Diptera: Calliphoridae), earwigs (Dermaptera), spiders (Araneae) and harvestmen (Opiliones) (40% of the diet). The difference in food selection by the two species is probably due to the different foraging strategies used.     

The role of swarming sites for maintaining gene flow in the brown long-eared bat (Plecotus auritus)

Bat-swarming sites where thousands of individuals meet in late summer were recently proposed as 'hot spots' for gene flow among populations. If, due to female philopatry, nursery colonies are genetically differentiated, and if males and females of different colonies meet at swarming sites, then we would expect lower differentiation of maternally inherited genetic markers among swarming sites and higher genetic diversity within. To test these predictions, we compared genetic variance from three swarming sites to 14 nursery colonies. We analysed biparentally (five nuclear and one sex-linked microsatellite loci) and maternally (mitochondrial D-loop, 550 bp) inherited molecular markers. Three mtDNA D-loop haplolineages that were strictly separated at nursery colonies were mixed at swarming sites. As predicted by the 'extra colony-mating hypothesis', genetic variance among swarming sites (V(ST)) for the D-loop drastically decreased compared to the nursery population genetic variance (V(PT)) (31 and 60%, respectively), and genetic diversity increased at swarming sites. Relatedness was significant at nursery colonies but not at swarming sites, and colony relatedness of juveniles to females was positive but not so to males. This suggests a breakdown of colony borders at swarming sites. Although there is behavioural and physiological evidence for sexual interaction at swarming sites, this does not explain why mating continues throughout the winter. We therefore propose that autumn roaming bats meet at swarming sites across colonies to start mating and, in addition, to renew information about suitable hibernacula.     

The foraging ecology of the mountain long-eared bat Plecotus macrobullaris revealed with DNA mini-barcodes

Molecular analysis of diet overcomes the considerable limitations of traditional techniques for identifying prey remains in bat faeces. We collected faeces from individual Mountain Long-eared Bats Plecotus macrobullaris trapped using mist nets during the summers of 2009 and 2010 in the Pyrenees. We analysed their diet using DNA mini-barcodes to identify prey species. In addition, we inferred some basic features of the bat's foraging ecology that had not yet been addressed. P. macrobullaris fed almost exclusively on moths (97.8%). As prey we detected one dipteran genus (Tipulidae) and 29 moth taxa: 28 were identified at species level (23 Noctuidae, 1 Crambidae, 1 Geometridae, 1 Pyralidae, 1 Sphingidae, 1 Tortricidae), and one at genus level (Rhyacia sp., Noctuidae). Known ecological information about the prey species allowed us to determine that bats had foraged at elevations between 1,500 and 2,500 m amsl (above mean sea level), mostly in subalpine meadows, followed by other open habitats such as orophilous grasslands and alpine meadows. No forest prey species were identified in the diet. As 96.4% of identified prey species were tympanate moths and no evidence of gleaning behaviour was revealed, we suggest P. macrobullaris probably forages by aerial hawking using faint echolocation pulses to avoid detection by hearing moths. As we could identify 87.8% of the analysed sequences (64.1% of the MOTUs, Molecular Operational Taxonomic Units) at species level, we conclude that DNA mini-barcodes are a very useful tool to analyse the diet of moth-specialist bats.     

Conflicting molecular phylogenies of European long-eared bats (Plecotus) can be explained by cryptic diversity

Conflicting phylogenetic signals of two data sets that analyse different portions of the same molecule are unexpected and require an explanation. In the present paper we test whether (i) differential evolution of two mitochondrial genes or (ii) cryptic diversity can better explain conflicting results of two recently published molecular phylogenies on the same set of species of long-eared bats (genus Plecotus). We sequenced 1714bp of three mitochondrial regions (16S, ND1, and D-loop) of 35 Plecotus populations from 10 European countries. A likelihood ratio test revealed congruent phylogenetic signals of the three data partitions. Our phylogenetic analyses demonstrated that the existence of a previously undetected Plecotus lineage caused the incongruities of previous studies. This lineage is differentiated on the species level and lives in sympatry with its sister lineage, Plecotus auritus, in Switzerland and Northern Italy. A molecular clock indicates that all European Plecotus species are of mid or late Pliocene origin. Plecotus indet. was previously described as an intergrade between P. auritus and Plecotus austriacus since it shares morphological characters with both. It is currently known from elevations above 800 m a.s.l. in the Alps, the Dinarian Alps and the Pindos mountains in Greece. Since we could demonstrate that incongruities of two molecular analyses simply arose from the mis-identification of one lineage, we conclude that molecular phylogenetic analyses do not free systematists from a thorough inclusion of morphological and ecological data.     

Analysis of the diet of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus in the West of Ireland

The diets of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus were investigated at a nursery colony of each species by analysis of droppings collected monthly from May to September. Respectively, 68% and 42% of the diets comprised items presumed to have been gleaned from foliage or other surfaces: diurnal insects, insects which rarely fly, and non-flying arthropods. Such surfaces included the ground: centipedes were eaten by both bats, and the muscid, Scatophaga stercoraria , usually associated with cattle dung, was a common prey. Indeed, the prevalence of cattle-farming around both of the bat roosts almost certainly contributed indirectly, in the form of S. stercoraria , to a significant part of the Diptera consumed. The chief food of M. nattereri was the larger Diptera, but Trichoptera, Hymenoptera and Arachnida were also important. Lepidoptera, Coleoptera and Hemiptera were minor prey, with Dermaptera and Chilopoda taken occasionally. Diptera, closely followed by Lepidoptera, together accounted for nearly two-thirds of the diet of P. auritus. Also taken, in descending order of importance, were Trichoptera, Arachnida, Chilopoda, Coleoptera, Dermaptera, Hymenoptera and Hemiptera. Plecotus auritus , but not M. nattereri , was evidently able to take a few small insects such as aphids and lesser nematoceran Diptera.     

Mating patterns, relatedness and the basis of natal philopatry in the brown long-eared bat, Plecotus auritus

The brown long-eared bat, Plecotus auritus, is unusual among temperate zone bats in that summer maternity colonies are composed of adult males and females, with both sexes displaying natal philopatry and long-term association with a colony. Here, we describe the use of microsatellite analysis to investigate colony relatedness and mating patterns, with the aim of identifying the evolutionary determinants of social organization in P. auritus. Mean colony relatedness was found to be low (R=0.033 +/- 0.002), with pairwise estimates of R within colonies ranging from -0.4 to 0.9. The proportion of young fathered by males in their own colony was investigated using a Bayesian approach, incorporating parameters detailing the number of untyped individuals. This analysis revealed that most offspring were fathered by males originating from a different colony to their own. In addition, we determined that the number of paternal half-sibs among cohorts of young was low, inferring little or no skew in male reproductive success. The results of this study suggest that kin selection cannot account for colony stability and natal philopatry in P. auritus, which may instead be explained by advantages accrued through the use of familiar and successful roost sites, and through long-term associations with conspecifics. Moreover, because the underlying causes of male natal dispersal in mammals, such as risk of inbreeding or competition for mates, appear to be avoided via extra-colony copulation and low male reproductive skew, both P. auritus males and females are able to benefit from long-term association with the natal colony.     

Mitochondrial phylogeography of the long-eared bats (Plecotus) in the Mediterranean Palaearctic and Atlantic Islands

Long-eared bats of the genus Plecotus are widespread and common over most of the western Palaearctic. Based on recent molecular evidence, they proved to represent a complex of several cryptic species, with three new species being described from Europe in 2002. Evolutionary relationships among the different lineages are still fragmentary because of the limited geographic coverage of previous studies. Here we analyze Plecotus mitochondrial DNA sequences from the entire Mediterranean region and Atlantic Islands. Phylogenetic reconstructions group these western Palaearctic Plecotus into two major clades which split at least 5 Myr ago and that are each subdivided into further subgroups. An 'auritus group' includes the traditional P. auritus species and its sister taxon P. macrobullaris (=P. alpinus) plus related specimens from the Middle East. P. auritus and P. macrobullaris have broadly overlapping distributions in Europe, although the latter is apparently more restricted to mountain ranges. The other major clade, the 'austriacus group,' includes the European species P. austriacus and at least two other related taxa from North Africa (including P. teneriffae from the Canary Islands), the Balkans and Anatolia (P. kolombatovici). The sister species of this 'austriacus group' is P. balensis, an Ethiopian endemic. Phylogenetic reconstructions further suggest that P. austriacus reached Madeira during its relatively recent westward expansion through Europe, while the Canary Islands were colonized by a North African ancestor. Although colonization of the two groups of Atlantic Islands by Plecotus bats followed very distinct routes, neither involved lineages from the 'auritus group.' Furthermore, the Strait of Gibraltar perfectly segregates the distinct lineages, which confirms its key role as a geographic barrier. This study also stresses the biogeographical importance of the Mediterranean region, and particularly of North Africa, in understanding the evolution of the western Palaearctic biotas.     

Energy metabolism and evaporative water loss in the European free-tailed bat and Hemprich's long-eared bat (Microchiroptera): species sympatric in the Negev Desert

We compared the thermoregulatory abilities of two insectivorous bat species, Tadarida teniotis (mean body mass 32 g) and Otonycteris hemprichii (mean body mass 25 g), that are of different phylogenetic origins and zoogeographic distributions but are sympatric in the Negev Desert. At night, both were normothermic. By day, both were torpid when exposed to ambient temperatures (T(a)) below 25 degrees Celsius, with concomitant adjustments in metabolic rate (MR). Otonycteris hemprichii entered torpor at higher T(a) than T. teniotis, and, when torpid, their body temperatures (T(b)) were 1 degrees -2 degrees Celsius and 5 degrees -8 degrees Celsius above T(a), respectively; MR was correspondingly reduced. At night, the lower critical temperature of T. teniotis was 31.5 degrees Celsius, and that of O. hemprichii was 33 degrees Celsius. Mean nocturnal thermoneutral MR of T. teniotis was 37% greater than that of O. hemprichii. At high T(a), evaporative water loss (EWL) increased markedly in both species, but it was significantly higher in T. teniotis above 38 degrees Celsius. In both species, the dry heat transfer coefficient (thermal conductance) followed the expected pattern for small mammals, by day and by night. Total EWL was notably low in normothermic and torpid animals of both species, much lower than values reported for other bats, indicating efficient water conservation mechanisms in the study species. Comparing thermoregulatory abilities suggests that O. hemprichii is better adapted to hot, arid environments than T. teniotis, which may explain its wider desert distribution. By both standard and phylogenetically informed ANCOVA, we found no differences in basal metabolic rate (BMR) between desert and nondesert species of insectivorous bats, substantiating previous studies suggesting that low BMR is a characteristic common to insectivorous bats in general.     

The functional significance of ventilation frequency,and its relationship to oxygen demand in the resting brown long-eared bat,Plecotus auritus

Summary Mean oxygen consumption and simultaneous ventilation frequency of nine non-reproductive brown long-eared bats (body mass 8.53–13.33 g) were measured on 159 occasions. Ambient (chamber) temperature at which the measurements were made ranged from 10.8 to 41.1°C. Apneic ventilation occurred in 22 of the 59 measurements made when mean oxygen consumption was less than 0.5 ml·min^-1. No records of apneic ventilation were obtained when it was over 0.5 ml·min^-1. The relationship between ventilation frequency and mean oxygen consumption depended on whether ventilation was apneic or non-apneic. When ventilation was non-apneic the relationship was positive and log-linear. When ventilation was apneic the relationship was log-log. Within the thermoneutral zone ventilation frequency was not significantly different from that predicted from allometric equations for a terrestrial mammal of equivalent body mass, but was significantly greater than that predicted for a bird. A reduction in the amount of oxygen consumed per breath occurred at ambient temperatures above the upper critical temperature (39°C).Abbreviations RH relative humidity - T _a chamber temperature - vf ventilation frequency - VO₂ oxygen consumption     

Effects of body mass and reproduction on the basal metabolic rate of brown long-eared bats (Plecotus auritus)

We measured basal metabolic rate (BMR) of nonreproductive and of breeding (pregnant and lactating) female brown long-eared bats (Plecotus auritus) to investigate the effects of intra- and interindividual variation in body mass and of reproduction on metabolism. The BMR of six nonreproductive females was measured between five and seven times at approximately 2-wk intervals over a period of 2.5 mo. There was a highly significant effect (P<0.001) of body mass on BMR of these nonreproductive females. The pooled within-individual scaling exponent (1.88) significantly exceeded the established mammalian interspecific exponent (0.75). In addition, we made single observations on 14 nonreproductive females to establish the effects of differences in mass between individuals. The mean BMR across all 14 individuals was 82 mW (+/-24 SD). There was a significant positive relationship between BMR and body mass across these individuals (r2=0.39), with a between-individual scaling exponent of 0.75. Inter- and intraindividual effects of mass on BMR were combined in a regression analysis that included mean body mass and deviation from mean mass on any given day as predictors. This regression model explained 55% of the variation in BMR. We made longitudinal measurements of BMR throughout reproduction and compared these with the predicted BMR of nonreproductive bats of the same body mass. Reproductive females exhibited temporal flexibility in BMR. BMR during pregnancy increased on a whole-animal basis but was significantly lower (by, on average, 15%) than BMR predicted for nonreproductive females of the same mass. Over a period of 1-75 d following birth, whole-animal BMR was greater than that during pregnancy, even though body mass declined after parturition. Hence, postbirth BMR was greater than the level predicted for nonreproductive females of the same mass. This study indicates that the scaling of BMR with body mass differs significantly within and between individuals and that there is a reduction of BMR in pregnancy and an elevation of BMR during lactation.     

Frequency sensitivity and directional hearing in the gleaning bat,Plecotus auritus (Linnaeus 1758)

1. The neural audiogram of the common long-eared bat, Plecotus auritus was recorded from the inferior colliculus (IC). The most sensitive best frequency (BF) thresholds for single neurones are below 0 dB SPL between 7-20 kHz, reaching a best value of -20 dB SPL between 12-20 kHz. The lower and upper limits of hearing occur at 3 kHz and 63 kHz, respectively, based on BF thresholds at 80 dB SPL. BF threshold sensitivities are about 10 dB SPL between 25-50 kHz, corresponding to the energy band of the sonar pulse (26-78 kHz). The tonotopic organization of the central nucleus of the IC (ICC) reveals that neurones with BFs below 20 kHz are disproportionately represented, occupying about 30% of ICC volume, occurring in the more rostral and lateral regions of the nucleus. 2. The acoustical gain of the external ear reaches a peak of about 20 dB between 8-20 kHz. The gain of the pinna increases rapidly above 4 kHz, to a peak of about 15 dB at 7-12 kHz. The pinna gain curve is similar to that of a simple, finite length acoustic horn; expected horn gain is calculated from the average dimensions of the pinna. 3. The directional properties of the external ear are based on sound diffraction by the pinna mouth, which, to a first approximation, is equivalent to an elliptical opening due to the elongated shape of the pinna. The spatial receptive field properties for IC neurones are related to the directional properties of the pinna. The position of the acoustic axis of the pinna and the best position (BP) of spatial receptive fields are both about 25 degrees from the midline between 8-30 kHz but approach the midline to 8 degrees at 45 kHz. In elevation, the acoustic axis and the BP of receptive fields move upwards by 20 degrees between 9-25 kHz, remaining stationary for frequencies up to 60 kHz. 4. The extremely high auditory sensitivity shown by the audiogram and the directionality of hearing are discussed in terms of the adaptation of the auditory system to low frequencies and the role of a large pinna in P. auritus. The functional significance of low frequency hearing in P. auritus is discussed in relation to hunting for prey by listening and is compared to other gleaning species.     

The influence of body condition on sexual development of male Brown long-eared bats (Plecotus auritus) in the wild

The relationship between sexual maturity and body condition was examined in a sample of 57 male Brown long-eared bats ( Plecolus auritus ) captured in autumn 1984 from nursery roosts in north-east Scotland (57°N). A direct relationship was found between reproductive status, which was established by external examination of the genitalia, and body weight. No such relationship was established between reproductive status and forearm length. A strong direct relationship was found between reproductive status and an index of body condition (body weight/forearm length). Three of the males examined had distended epididymides, suggesting they were filled with spermatozoa, and that the bats concerned had recently undergone spermatogenesis.
Reproductive status also varied between maternity roosts. Bats from roosts in areas with a high proportion of tree cover were more sexually developed, and in better body condition, than bats from roosts in areas with less tree cover.     

Habitat selection of three cryptic Plecotus bat species in the European Alps reveals contrasting implications for conservation

Assessing the ecological requirements of species coexisting within a community is an essential requisite for developing sound conservation action. A particularly interesting question is what mechanisms govern the stable coexistence of cryptic species within a community, i.e. species that are almost impossible to distinguish. Resource partitioning theory predicts that cryptic species, like other sympatric taxa, will occupy distinct ecological niches. This prediction is widely inferred from eco-morphological studies. A new cryptic long-eared bat species, Plecotus macrobullaris, has been recently discovered in the complex of two other species present in the European Alps, with even evidence for a few mixed colonies. This discovery poses challenges to bat ecologists concerned with planning conservation measures beyond roost protection. We therefore tested whether foraging habitat segregation occurred among the three cryptic Plecotus bat species in Switzerland by radiotracking 24 breeding female bats (8 of each species). We compared habitat features at locations visited by a bat versus random locations within individual home ranges, applying mixed effects logistic regression. Distinct, species-specific habitat preferences were revealed. P. auritus foraged mostly within traditional orchards in roost vicinity, with a marked preference for habitat heterogeneity. P. austriacus foraged up to 4.7 km from the roost, selecting mostly fruit tree plantations, hedges and tree lines. P. macrobullaris preferred patchy deciduous and mixed forests with high vertical heterogeneity in a grassland dominated-matrix. These species-specific habitat preferences should inform future conservation programmes. They highlight the possible need of distinct conservation measures for species that look very much alike.     

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.