Testing traditional assumptions about regional migration in bats

While some bats cover long distances during migration, moving thousands of kilometers, most migratory bats are considered regional migrants, thought to move relatively short distances (<?500 km) between hibernacula and maternity sites. However, behavior can vary considerably among species and our understanding of these movements has largely been limited to banding studies or detailed tracking of small numbers of bats by aircraft. Inferring population-wide behavior from small samples is difficult and can introduce bias. We tagged 108 Indiana bats (Myotis sodalis) in the Midwestern US and used a regional network of radiotelemetry receivers to study movement patterns. With this dataset, we tested the following traditional generalizations about regional migrants: (1) bats move away from hibernacula in spring in all directions with known maternity roosts, e.g., in a star-like pattern; (2) bats follow linear landscape features; (3) long-distance movements are uncommon; and (4) autumn migration comprises a single movement from summer maternity site to winter hibernaculum. In spring, bats left the hibernaculum immediately and primarily moved north despite available maternity roosts in all directions. We found no evidence that bats follow rivers, the predominant linear element in the landscape. Only six tagged bats traveled >?100 km, suggesting that longer-distance movements may be outliers. In autumn, only two bats visited multiple known hibernacula, and after swarming, some females moved >?100 km to areas without known hibernacula. Common generalizations about regional migrant movements may not be representative of population behavior and care should be taken with respect to management decisions based on those assumptions.     

Predation of wood mice (Apodemus sylvaticus) on hibernating bats

Bat hibernacula with high numbers of bats can become high-risk areas, as they attract flying and non-flying predators. In order to protect hibernating bats effectively, more knowledge about mortality factors is needed. During the winters of 2003–2015, we found 214 dead bats in 12 hibernacula in The Netherlands province of Zuid-Holland. Most bat remains were found in December and January, with a second peak in April. Their remains showed a typical pattern of lesions consistent with those caused by predation by the wood mouse (Apodemus sylvaticus). Trail camera surveys showed that wood mice actively searched for bats. Predation pressure seemed to vary between winters, with a peak in the winters of 2004, 2011 and 2015. The annual mortality (relative to the maximum winter population size) caused by wood mouse predation varied between 0.1 and 8.8 %, with a maximum local effect of 83.6 %. The years with high wood mouse predation pressure were characterized by a long frost period and a low mast production of common oak in the preceding autumn. The size of a hibernaculum and the population density of its bats had an effect on predation-dependent mortality. The highest predation risk occurred near the entrance of bunkers. From these results we tentatively conclude that predation is not incidental and that wood mice actively search for and kill hibernating bats or scavenge for weakened individuals.     

Hibernacula microclimate and declines in overwintering bats during an outbreak of white‐nose syndrome near the northern range limit of infection in North America

We document white‐nose syndrome (WNS), a lethal disease of bats caused by the fungus Pseudogymnoascus destructans (Pd), and hibernacula microclimate in New Brunswick, Canada. Our study area represents a more northern region than is common for hibernacula microclimate investigations, providing insight as to how WNS may impact bats at higher latitudes. To determine the impact of the March 2011 arrival of Pd in New Brunswick and the role of hibernacula microclimate on overwintering bat mortality, we surveyed bat numbers at hibernacula twice a year from 2009 to 2015. We also collected data from iButton temperature loggers deployed at all sites and data from HOBO temperature and humidity loggers at three sites. Bat species found in New Brunswick hibernacula include Myotis lucifugus (Little Brown Bat) and M. septentrionalis (Northern Long‐eared Bat), with small numbers of Perimyotis subflavus (Tricolored Bat). All known hibernacula in the province were Pd‐positive with WNS‐positive bats by winter 2013. A 99% decrease in the overwintering bat population in New Brunswick was observed between 2011 and 2015. We did not observe P. subflavus during surveys 2013–2015 and the species appears to be extirpated from these sites. Bats did not appear to choose hibernacula based on winter temperatures, but dark zone (zone where no light penetrates) winter temperatures did not differ among our study sites. Winter dark zone temperatures were warmer and less variable than entrance or above ground temperatures. We observed visible Pd growth on hibernating bats in New Brunswick during early winter surveys (November), even though hibernacula temperatures were colder than optimum for in vitro Pd growth. This suggests that cold hibernacula temperatures encountered near the apparent northern range limit for Pd do not sufficiently slow fungal growth to prevent the onset of WNS and associated bat mortality over the winter.     

Identifying unusual mortality events in bats: a baseline for bat hibernation monitoring and white‐nose syndrome research

Bat population trends are particularly affected by adult mortality, especially when large numbers of individuals die, as evidenced by white‐nose syndrome in North America. We obtained baseline mortality data from 318 European hibernacula. Mortality was low and negatively associated with elevation but not with fungal infestation. Mortality events involving more than seven bats at a hibernaculum should be considered unusual, and above this threshold, pathological or microbiological analysis should be carried out. To increase understanding of mortality in bats, there is an urgent need to develop and co‐ordinate national and international programs for monitoring and investigating mortality and diseases.     

Migratory behaviour of the Schreiber's bat: when, where and why do cave bats migrate in a Mediterranean region?

Regional migrations are important elements of the biology of bats, but remain poorly understood. We obtained a large dataset of recoveries of ringed Miniopterus schreibersii to study the patterns and drivers of migration of a Mediterranean cave-dwelling bat. In spite of the mildness of Mediterranean winters, in average years bats hibernated, and few movements were recorded during this period. After hibernation, females migrated to spring roosts, and again to maternity roosts just before parturition. This late arrival at nurseries could be a strategy to avoid a harmful build-up of parasites. Soon after the juveniles were weaned, the mothers migrated to the roosts where they spent autumn and sometimes also winter. Juveniles remained in the warm nurseries longer, presumably because high roost temperatures speed up growth. The pattern of migration of males was similar to that of females, but they left hibernacula later and remained more mobile during the maternity season. They also arrived at the hibernacula later, possibly because they needed time to build up fat stores after the energetically costly mating season. Maternity colonies spent the yearly cycle in well-defined home ranges (mean=19 030 km²), which overlapped greatly. Bats were furthest from the maternity sites during hibernation, but even then 80% remained within 90 km of them. Each hibernaculum attracted bats from multiple nurseries, from within a mean range of 10 770 km². We tested two potential drivers for migration – temperature in the roosts and at the foraging areas – but our results supported only the first one. Bats migrated to reach the roosts most thermally suited for each phase of their life cycle, indicating that roost temperature and associated metabolic advantages are key drivers for regional migrations of cave-dwelling bats.     

Going, going, gone: the impact of white-nose syndrome on the summer activity of the little brown bat (Myotis lucifugus)

Since its discovery in the winter of 2005-2006, white-nose syndrome (WNS) has killed over one million little brown bats (Myotis lucifugus) in the American northeast. Although many studies have reported die-offs of bats at winter hibernacula, it is important to understand how bat mortality linked to WNS at winter hibernacula affects bat activity levels in their summer ranges. In the summer (May-August) of 2007, 2008 and 2009, we recorded echolocation calls to determine bat activity at sites along the Hudson River, NY (within approx. 100 km of where WNS was first reported). We documented a 78 per cent decline in the summer activity of M. lucifugus, coinciding with the arrival and spread of WNS. We suggest that mortality of M. lucifugus in winter hibernacula is reflected by reduced levels of activity in the summer and that WNS affects the entire bat population of an area, and not only individual hibernacula.     

Roost selection by Formosan leaf-nosed bats (Hipposideros armiger terasensis)

Patterns of roost use by Formosan leaf-nosed bats (Hipposideros armiger terasensis) were studied from November 1998 to April 2000. Structural characteristics, microclimates, and disturbance levels of 17 roosts used by H. a. terasensis and 15 roosts either used by other bat species (2) or not occupied by any bat species were compared. Roosts used by these bats were significantly larger in size and had greater areas covered by water compared to unused roosts. Entrances of active roosts were more likely to be east-west oriented. Hibernacula had lower entrances and ceilings than did roosts used only in summer. Higher temperatures were recorded in non-breeding roosts than in breeding roosts, but temperature gradients in these two types of roosts did not differ. In winter, hibernacula were warmer, and the temperature fluctuated less than in non-hibernacula. The relative humidities in summer roosts and hibernacula were nearly 100%. Disturbance levels were significantly higher in non-breeding roosts than in breeding roosts, and in non-hibernacula than in hibernacula. These results suggest that the Formosan leaf-nosed bats are selective of their roosts, but the pattern of their roost selection differs from those reported for bats of temperate regions. The reasons for such differences may be related to differences in body size, behavior, and reproductive strategy of the Formosan leaf-nosed bats living in a subtropical climate in Taiwan.     

Characteristics of the northernmost population ofRhinolophus ferrumequinum in the Carpathian Basin

Roost utilisation byRhinolophus ferrumequinum (Schreber, 1774) was investigated between 1984 and 1998 in north-eastern Hungary. Exploration of summer and winter roosts, monitoring and bat-banding were implemented to find movements between the colonies. Data on roost utilisation by this species in south-eastern Slovakia, collected in a similar way, were included for comparison. Twenty-two marked bats were recaptured. The studied bats created nursery colonies in Hungarian churches and moved to Slovakian mines and caves to hibernate in winter. The population used two main hibernacula, two large nursery roosts and one temporary roost but several other roosts were also visited. The area occupied by the population was 5180 km².R. ferrumequinum living in SE Slovakia and NE Hungary formed probably a separate population on the northern edge of the species range. This population is a part of the metapopulation of the species.     

Winter behavior of bats and the progression of white‐nose syndrome in the southeastern United States

Understanding the winter behavior of bats in temperate North America can provide insight into how bats react to perturbations caused by natural disturbances such as weather, human‐induced disturbances, or the introduction of disease. This study measured the activity patterns of bats outside of their hibernaculum and asked how this winter activity varied by time, temperature, bat species, body condition, and WNS status. Over the course of three winters (2011–2013), we collected acoustic data and captured bats outside of five hibernacula in Tennessee, United States. During this time, Pseudogymnoascus destructans, the causative agent of white‐nose syndrome, became established in hibernacula throughout the region, allowing us to track disease‐related changes in the winter behavior of ten bat species. We determined that bats in the southeastern United States were active during winter regardless of disease. We recorded activity outside of hibernacula at temperatures as low as ?13°C. Although bat activity was best determined by a combination of variables, the strongest factor was mean daily temperature (R ² = .2879, F _1,1450 = 586.2, p  < .0001). Bats that left the hibernacula earlier in evening had lower body condition than those that left 2–4 hr after sunset (F _7,932 = 7.225, p  < .0001, Tukey HSD, p  < .05). The number of daytime emergences from hibernacula, as determined via acoustic detection, increased the longer a site was P. destructans positive (F _{3,17 808} = 124.48, p  < .0001, Tukey HSD, p  < .05). Through the use of passive acoustic monitoring and monthly captures, we determined that winter activity was driven by both ambient temperature and the presence of P. destructans .     

Density regulation in toad populations (Epidalea calamita,Bufotes viridis) by differential winter survival of juveniles

The size of amphibian populations varies considerably between years, so that systematic trends in dynamics are difficult to detect. Informed conservation management of presumably declining populations requires the identification of the most sensitive life stage. In temperate-zone anurans there is growing evidence that juveniles hibernating for the first time suffer from substantial winter losses. In two syntopic toads (Epidalea calamita, Bufotes viridis) we monitored survival of such juveniles during four consecutive winters in the natural habitat and in four temperature treatments (3°, 5 °C, 10°/15 °C or 20 °C, natural light-dark cycle) in temperature-controlled chambers during winter. Specifically, we tested the hypotheses that (1) winter mortality of juvenile toads which hibernate for the first time in their life is an important component of population dynamics, and that (2) mortality rates differed between the two species. Parameters quantified were size-dependent winter mortality and body condition of pre- and post-hibernating juveniles. Field data provided evidence for the important role of winter mortality of first-hibernators in population dynamics. Choice of hibernacula differed in E. calamita between small and medium-sized individuals and also between the two species suggesting distinct mortality risks. The inability of small E. calamita to reach frost-proof hibernacula by burrowing, and the exposure of small B. viridis to predators are the most probable causes of size-assortative winter mortality. In conclusion, E. calamita juveniles may benefit from rising average winter temperatures in the future by decreased risk of freezing to death, whereas predator-caused winter mortality of B. viridis juveniles will also depend on the effects of climate warming on predator phenology.     

OBSERVATIONS ON NOCTULE BATS (NYCTALUS NOCTULA) CAPTURED WHILE FEEDING

In the summer and early autumn of 1960, 1961 and 1962, noctule bats flying low, taking house crickets as these flew from a municipal rubbish tip, were captured in mist nets, ringed, released and in many cases recaptured a number of times. The flying bats showed no fear of human beings or predatory birds and did not learn to avoid the net. In June and July of each year the majority of bats caught were adult females, the flying young of the year first appearing in August though some did not fly until September and October. Young males did not reach sexual maturity in the year of their birth, though five out of fourteen females recaptured at a year old did. There was a considerable movement of adult males in the late summer, adult bats being captured in approximately equal numbers of both sexes during August and September. In October the females seemed to disappear, the majority of the bats caught during that month being males: by November the crickets had ceased to fly and no more bats could be captured though a few wero still flying on warm nights. There was a marked difference in feeding behaviour over these three years, the bats concentrating more on crickets in 1960 than in 1961 and 1962. Though the differences are not statistically significant there were indications of an increase in body weight between July and October in the years when less cricket feeding was occurring. About 50 per cent of the females captured in each of the years 1960 and 1961 were recaptured feeding on the same site in the following year: the recovery rate of males was about 30 per cent in 1961, 60 per cent in 1962.     

The trans-boundary importance of artificial bat hibernacula in managed European forests

Many European migratory bat species hibernate in large hollow trees, a decreasing resource in present day silviculture. Here, we report on the importance of man-made hibernacula to support trans-boundary populations of noctule bats (Nyctalus noctula), a species that performs seasonal long distance movements throughout Europe. In winter, we surveyed nine bat roosts (eight artificial and one natural) in Germany and collected small tufts of fur from a total of 608 individuals. We then measured the stable isotope ratios of the non-exchangeable hydrogen in fur keratin and estimated the origin of migrants using a refined isoscape origin model that included information on expected flight distances and migration directions. According to the stable isotope signature, 78 % of hibernating bats originated from local populations. The remaining 22 % of hibernacula occupants originated from distant populations, mostly from places in northern or eastern countries such as Sweden, Poland and Baltic countries. Our results confirm that many noctule bats cross one or several political borders during migration. Data on the breeding origin of hibernating noctule bats also suggest that artificial roosts may not only be important for local but also for distant populations. Protection of natural and artificial hibernacula in managed forests may support the trans-boundary populations of migratory bats when hollow trees are scarce in managed forests.     

Emergence activity at hibernacula differs among four bat species affected by white‐nose syndrome

Prior to the introduction of white‐nose syndrome (WNS) to North America, temperate bats were thought to remain within hibernacula throughout most of the winter. However, recent research has shown that bats in the southeastern United States emerge regularly from hibernation and are active on the landscape, regardless of their WNS status. The relationship between winter activity and susceptibility to WNS has yet to be explored but warrants attention, as it may enable managers to implement targeted management for WNS‐affected species. We investigated this relationship by implanting 1346 passive integrated transponder (PIT) tags in four species that vary in their susceptibility to WNS. Based on PIT‐tag detections, three species entered hibernation from late October to early November. Bats were active at hibernacula entrances on days when midpoint temperatures ranged from −1.94 to 22.78°C (mean midpoint temperature = 8.70 ± 0.33°C). Eastern small‐footed bats (Myotis leibii), a species with low susceptibility to WNS, were active throughout winter, with a significant decrease in activity in mid‐hibernation (December 16 to February 15). Tricolored bats (Perimyotis subflavus), a species that is highly susceptible to WNS, exhibited an increase in activity beginning in mid‐hibernation and extending through late hibernation (February 16 to March 31). Indiana bats (M. sodalis), a species determined to have a medium–high susceptibility to WNS, remained on the landscape into early hibernation (November 1 to December 15), after which we did not record any again until the latter portion of mid‐hibernation. Finally, gray bats (M. grisescens), another species with low susceptibility to WNS, maintained low but regular levels of activity throughout winter. Given these results, we determined that emergence activity from hibernacula during winter is highly variable among bat species and our data will assist wildlife managers to make informed decisions regarding the timing of implementation of species‐specific conservation actions.     

Digital Photography Improves Consistency and Accuracy of Bat Counts in Hibernacula

ABSTRACT The size and distribution of measurement errors associated with major techniques for estimating numbers of hibernating bats are unstudied, although this is the principle method of enumerating several endangered bat species. However, decisions concerning the listing status of a species under the Endangered Species Act require consistent and accurate estimation of population size and trends. Recent advances in digital photography have improved the ability to produce a quantitative record of the numbers of bats in hibernacula. We surveyed clusters of Indiana bats in a hibernaculum and compared results from counts of digital photographs of clusters to results from 4 variations of visual estimation. We counted bats in photographs using Geographic Information System digitization over the photograph. Total counts from 2 sets of photographs varied by <1.5%. Nonphotographic estimation techniques varied from 76% to 142% of counts from photographs for clusters for which estimation (rather than counting) was used. Where feasible, photography can improve status and trend information for species of concern, permitting more timely and specific management actions.     

Present status of commercial stocks of sturgeons in the Caspian Sea basin

Catches for the last 25 years are analyzed for beluga Huso huso, stellate sturgeon A. stellatus and Russian sturgeon Acipenser gueldenstaedtii, which are the three commercially important species of sturgeons found in the Caspian Sea Basin. Population sizes for generations born between 1961 and 1970 are estimated, and found to depend on natural reproduction and the number of young fish stocked annually from sturgeon hatcheries located in the Volga River Delta. A ban on sea fishing from 1962 to 1991 positively impacted the number and total biomass of commercial stocks. Sturgeon growth rates depend on water levels in the Caspian Sea. In order to preserve Caspian Sea sturgeon populations, it will be necessary to coordinate efforts of all countries surrounding the Caspian Sea to achieve rational harvests, preserve juveniles, and produce at least 100 million juveniles annually from hatcheries.     

河南济源人工引水渠隧道蝙蝠冬眠生态学特征

2017-2020年期间,每年1月份对河南省济源市邵原镇布袋沟水库人工引水渠隧道内蝙蝠进行冬眠生态学特征调查,共发现2科5属7种蝙蝠在此冬眠,包括马铁菊头蝠（Rhinolophus ferrumequinum）、小菊头蝠（R.pusillus）、华南水鼠耳蝠（Myotis laniger）、白腹管鼻蝠（Murina leucogaster）、金管鼻蝠（Mu.aurata）、奥氏长耳蝠（Plecotus ognevi）和亚洲宽耳蝠（Barbastella leucomelas）。马铁菊头蝠是优势种（约52%-73%的冬眠个体）,其次是小菊头蝠（约19%-37%）、华南水鼠耳蝠（约5%-8%）,其余蝙蝠物种数量不足3%。2017-2020年冬眠蝙蝠个体总数呈增长趋势,但仍少于早期报道的数量。有42个隧道每年均有蝙蝠冬眠,而且不同年度冬眠数量也不尽相同。通过多元线性回归分析发现,隧道长度可能是影响蝙蝠冬眠栖息场所选择的主要影响因子（Adjusted R²=0.208,P=0.001）。每个隧道内,蝙蝠具有不同的冬眠栖点位置,约4/5的蝙蝠选择温暖且环境相对稳定的隧道深处（> 30 m）作为冬眠栖点,超过95%的个体选择长度> 60 m的隧道冬眠。蝙蝠具有不同的冬眠方式,绝大多数个体采用独栖方式进行冬眠（> 90%）,少数采用聚集方式。不同的冬眠栖点和冬眠方式可能有利于冬眠成本优化。此外,栖点温度与蝙蝠体温之间呈显著正相关（R²=0.98,P < 0.001）,而且蝙蝠冬眠期间的栖点温度具有种内和种间差异。研究结果为我国蝙蝠种群保护和冬眠场所管理提供科学依据。     

Nesting behaviour and seasonal body mass changes in a rural Irish population of the Western hedgehog (<Emphasis Type="Italic">Erinaceus europaeus</Emphasis>)

There was previously no information on nesting, seasonal body mass or the hibernation behaviour of the hedgehog in Ireland. Between 2008 and 2009, hedgehogs were caught, weighed weekly and monitored by radio tracking at a rural Irish site. Day nests were recorded in the active period and hibernacula thereafter. Arable land has been reported to be poorly utilised by hedgehogs in the UK and Denmark. In Ireland, day nests and hibernacula were constructed in the hedgerows of arable land indicating that these areas will be exploited if hedgerow is maintained and highlights the importance of maintaining hedgerows in arable areas. Individual females returned to the same day nest significantly more than males. Over the 2 years, individuals were found to occupy a mean of 1.8 (±0.9; SD) hibernacula (maximum of 3) and they rotated between nests up to four times (mean of 2.5?±?1.6). When hedgehogs occupied multiple hibernacula, those occupied in mid winter (December and January) were occupied for significantly longer than those occupied at the start (October and November) and end (February and March) of hibernation. Studies in the UK and Denmark have reported on hedgehog hibernation. However, the winter climate in southern Ireland is milder than other areas of Europe and thus it was expected that hibernation characteristics would also differ, i.e. shorter hibernation periods, earlier emergence, lower body mass loss and the ability to survive hibernation at a lower body mass. This proved to be the case with a mean hibernation period (±SE) of 148.9 (±0.5) days, a mean body mass loss of 17.0 (±0.53) %, emergence in March and the ability of late juveniles to survive at a pre-hibernation body mass of 475 g.     

Mark–recapture may reveal more about ecology than about population trends: Demography of a threatened ghost bat (Macroderma gigas) population

Since European settlement in Australia, the geographical range of ghost bats (Macroderma gigas) has contracted northwards. Ghost bats are thought to occur in disjunct populations with little interpopulation migration, raising concerns over the current status and future viability of the southernmost colony, which has also been threatened by mining activity. To address these concerns, demographic parameters of the southernmost colony were estimated from a mark–recapture study conducted during 1975–1981. Female bats gave birth to a single young in late spring, but only 40% (22–70%, 95% CI) of females bred in their second year, increasing to 93% (87–97%, 95% CI) for females ≥ 2 years old. Sixty‐five percent of juveniles caught were female. Annual adult survival ranged between 0.57–0.77 for females and 0.43–0.66 for males, and was lowest over winter–spring and greatest in autumn–winter. Juvenile survival for the first year ranged between 0.35–0.46 for females and 0.29–0.42 for males. Adult survival varied among seasons, was negatively associated with rainfall, but was not associated with temperature beyond being lower in late winter. Poor survival may result from the inferior daytime roosts that bats must use if water seepage forces them to leave their normal roosts. Although these age‐specific rates of fecundity and survival suggested a declining population, mark–recapture estimates of the population trend indicated stability over the study period. Counts at daytime roosts also suggested a population decline, but were considered unreliable because of an increasing tendency of bats to avoid detection. It is therefore likely that some assumptions in estimating survival were violated. These results provide a caution against the uncritical use of population projections derived from mark–recapture estimates of demographic parameters, and the use of untested indices as the basis for conservation decisions.     

Subtropical mouse-tailed bats use geothermally heated caves for winter hibernation

We report that two species of mouse-tailed bats (Rhinopoma microphyllum and R. cystops) hibernate for five months during winter in geothermally heated caves with stable high temperature (20°C). While hibernating, these bats do not feed or drink, even on warm nights when other bat species are active. We used thermo-sensitive transmitters to measure the bats’ skin temperature in the natural hibernacula and open flow respirometry to measure torpid metabolic rate at different ambient temperatures (T_a, 16–35°C) and evaporative water loss (EWL) in the laboratory. Bats average skin temperature at the natural hibernacula was 21.7 ± 0.8°C, and no arousals were recorded. Both species reached the lowest metabolic rates around natural hibernacula temperatures (20°C, average of 0.14 ± 0.01 and 0.16 ± 0.04 ml O₂ g⁻¹ h⁻¹ for R. microphyllum and R. cystops, respectively) and aroused from torpor when T_a fell below 16°C. During torpor the bats performed long apnoeas (14 ± 1.6 and 16 ± 1.5 min, respectively) and had a very low EWL. We hypothesize that the particular diet of these bats is an adaptation to hibernation at high temperatures and that caves featuring high temperature and humidity during winter enable these species to survive this season on the northern edge of their world distribution.     

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.