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1.
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.  相似文献   

2.
Group fission is an important dispersal mechanism for philopatric adults. In Cypress Hills Interprovincial Park, Saskatchewan, tree-roosting big brown bats (Eptesicus fuscus) exhibit fission-fusion roosting behaviour. During 2004-2007, the majority of females previously resident to roosting area 1 (RA1) moved to a new roosting area (RA4). We examined how genetic relationships, inferred from data for microsatellite loci and mitochondrial DNA, influenced new roost area (RA) selection during 2006 when colony members were split between the RAs. We found that females who moved to RA4 had higher average relatedness than those that remained in RA1. We found that nearly all females belonging to matrilines with high average relatedness moved to RA4 while females from matrilines with low average relatedness were split between the two RAs. These results suggest that closely related maternal kin preferentially move to new RAs. However, daily roosting preferences within a RA are not based on genetic relationships probably because daily roosting associations between kin and non-kin are used to ensure adequate roost group size. Studying the effects of kinship on the fission and movements of groups not only enhances our understanding of social behaviour and population genetics but also informs conservation decisions.  相似文献   

3.
We used radiotelemetry to quantify roost switching and assess associations between members of maternity colonies of forest-dwelling big brown bats. Bats remained loyal to small roosting areas of forest within and between years and switched trees often (). For radiotagged bats from the colony in one of these areas, roost-switching frequency was positively correlated with the number of different individuals with which tagged bats shared roosts. We quantified associations between pairs of bats using a pairwise sharing index and found that bats associated more often than predicted when roost and roostmate selection were random but that all tagged bats spent at least some days roosting in different trees, apart from preferred roostmates. Our results suggest that forest-dwelling big brown bats conform to a fission-fusion roosting pattern. Roost switching in forests may reflect the maintenance of long-term social relationships between individuals from a colony that is spread among a number of different trees on a given night. In this fission-fusion scenario, switching between trees, within a local area, could serve to increase the numbers of individuals with which bats maintain associations. We contend that roosting areas in forests are analogous to spatially large roosts in caves, mines and buildings.  相似文献   

4.
The prevalent view of group splitting during group decisions is that a beneficial consensus has not been reached because time constraints, different individual information, or inter-individual conflicts lead to fission instead of a compromise. However, societies with high fission-fusion dynamics may allow their members to avoid consensus decisions that are not in their favour without foregoing grouping benefits that arise from collective behaviour. Moreover, by forming temporary subgroups that represent individual preferences better than the group as a whole fission-fusion societies could avoid a permanent break up even in situations where conflicts among their members are to strong to reach a consensus.  相似文献   

5.
Logging is one of the greatest threats to global biodiversity, while forests are one of the most important habitats for bats. Bats that roost in tree cavities require a large number of potential roosts due to their frequent roost switching. However, the density of tree cavities and hollows sufficient to sustain large populations of bat species in forests is unknown. The fission-fusion dynamics of bat groups in forest environment is associated with ritualised dawn swarming behaviour at potential tree cavities that serves to exchange information in a non-centralised decision-making process. We used a computer model based on the swarm algorithm, SkyBat, that resembles this complex process and aimed to determine how population size changes over time when cavity trees are removed from roosting territory of the local population of Leisler's bats (Nyctalus leisleri), which inhabit a forest habitat in Central Europe. Simulations revealed that social bonds between bats, maintained by frequent switching among groups, play an important role in this highly dynamic system. When strong social contact was not considered, reducing the original number of trees with cavities (20 cavities × ha−1) to 50% was still acceptable to bats, but further interventions and/or increased demand for social contact would have led to local extinction of the species. Results suggest that potential bat roosts in mature forest stands should be preserved as much as possible and that non-intensive logging and management can be beneficial to tree-dwelling bats.  相似文献   

6.
The decision where to live has far-reaching fitness consequences for animals. In contrast to most other mammals or birds that use sheltered nest sites, female Bechstein's bats frequently switch day roosts during one breeding season, and therefore must often decide where to spend the day. Selecting the right roost is important, because roost quality, e.g. microclimatic condition, influences survival and reproduction in bats. Although thermal factors are very important for the quality of roosts occupied by bats, whether bats base their day roost selection directly on roost temperature has not been tested in the field. Over one summer, we examined and tested the roost choice of 21 individually marked female Myotis bechsteinii living in one maternity colony. In a field experiment, we allowed the bats to choose between relatively warm versus cold bat boxes, while controlling for site preferences. We expected females to exhibit a preference for warm roosts during pregnancy and lactation to accelerate gestation and shorten the period of growth of their young. Roost occupancy over 160 census days reflected significant temperature differences among 89 surveyed roosts (14 tree holes and 75 bat boxes), and preferences changed with the season. Females significantly preferred cold roosts before parturition, whereas post-partum, they significantly favoured warm roosts. Temperature preferences were independent of the roost site, and thus roost selection was based directly on temperature. Boxes with significantly different daytime temperatures did not differ significantly at night. Consequently, bats would have to spend at least 1 day in a new roost to test it. Information transfer among colony members might facilitate knowledge of roost availability. Access to many roosts providing different microclimates is likely to be important for successful reproduction in the endangered Bechstein's bat.  相似文献   

7.
Animals can use signals emitted by other animals as sources of information. Auditory signals are important in communication networks, as they can potentially convey information about the location and state of conspecifics and other species over long distances. Signalling is important in fission–fusion societies, in which animals from the same social group temporarily split into subgroups and frequently change roost sites. We used playbacks of social calls of the noctule Nyctalus noctula produced in roosts, to show how bats might maintain group cohesion and to test the hypothesis that noctules can locate conspecifics when returning from foraging trips by eavesdropping on or communicating with roosting individuals. Noctules responded strongly to broadcasted social calls. Their reactions included inspections and landing on a loudspeaker broadcasting social calls and occasional social vocalisation. Responses by other bat species to the noctule social calls were negligible. The high amplitude, low‐frequency vocalisations emitted by noctules in roosts can propagate over long distances and allow group members to announce their position. Bats can extract information about the location of roosts containing conspecifics by eavesdropping or by communication. Social calls may thus be sufficient to locate conspecifics in roosts and maintain spatial associations of groups in mammals.  相似文献   

8.
We evaluated the spatial and temporal patterns of roost switching behaviour by a tree-dwelling population of barbastelle bats Barbastella barbastellus in a beech forest of central Italy. Switching behaviour was common to both sexes and did not depend on group size. We observed both individual and group switching, the latter often involving the abandonment of a roost tree on a single night. We suggest that behaviours such as flight activity around roosts or cavity inspection by bats play a role in recruiting group mates and coordinating their occupation of another site. Bats almost never crossed mountain ridges to use roosts located beyond them, possibly because ridges are regarded as boundaries delimiting main roosting areas. The rate of switching was lowest during the middle of the lactation period, probably to minimise problems related to the transportation of non-volant young by their mothers. Although the maintenance of social relationship among bats spread over large forest areas may partly explain the occurrence of roost switching, the persistence of this behaviour in solitary bats and the movement of entire groups best fit the hypothesis that roost switching represents a way to maintain or increase knowledge of alternative roosts.  相似文献   

9.
Dusk to dawn observations, using a bat detector and occasionally an image intensifier, were made outside two nursery roosts of lesser horseshoe bats Rhinolophus hipposideros from late May to September. Emergence was correlated with sunset but delayed by extended twilight. Light intensity was important in triggering departure and cloud cover advanced it. Light-testing behaviour was invariably undertaken, in the form of brief flights out and back into the roosts. The exit from one roost was shaded by trees and exploratory flights were generally more extended there. Heavy rain inhibited emergence. There was almost always intermittent activity throughout the night, with many individuals returning and departing, and no indication of seasonal or overnight peaks. A bat detector inside a third roost confirmed overnight observations at the other two. Some bats often returned to the roost for the night before dawn. Dawn return was linked to sunrise, prolonged twilight in midsummer hastening it. Colony size varied appreciably over periods of a few days and even overnight. There is some limited evidence that increased colony size, perhaps through social interaction, may have influenced timing of departure at dusk and return at dawn.  相似文献   

10.
Social calls in bats have many functions, including mate attraction and maintaining contact during flight. Research suggests that social calls may also be used to transfer information about roosts, but no studies have yet demonstrated that calls are used to actively attract conspecifics to roosting locations. We document the social calls used by Spix''s disc-winged bat (Thyroptera tricolor) to actively recruit group members to roosts. In acoustic trials, we recorded two sets of calls; one from flying individuals termed ‘inquiry calls’, and another from roosting bats termed ‘response calls’. Inquiry calls were emitted by flying bats immediately upon release, and quickly (i.e. 178 ms) elicited production of response calls from roosting individuals. Most flying bats entered the roost when roosting individuals responded, while few bats entered the roost in the absence of a response. We argue that information transfer concerning roost location may facilitate sociality in T. tricolor, given the ephemeral nature of roosting structures used by this species.  相似文献   

11.
Reckardt K  Kerth G 《Oecologia》2007,154(3):581-588
Ectoparasites of vertebrates often spend part of their life cycle in their hosts’ home. Consequently, hosts should take into account the parasite infestation of a site when selecting where to live. In a field study, we investigated whether colonial female Bechstein’s bats (Myotis bechsteinii) adapt their roosting behaviour to the life cycle of the bat fly Basilia nana in order to decrease their contact with infective stages of this parasite. B. nana imagoes live permanently on the bat’s body but deposit puparia in the bat’s roosts. The flies metamorphose independently in the roosts, but after metamorphosis emerge only in the presence of a potential host. In a field experiment, the bats preferred non-contagious to contagious day-roosts and hence were able to detect either the parasite load of roosts or some correlate with infestation, such as bat droppings. In addition, 9 years of observational data on the natural roosting behaviour of female Bechstein’s bats indicate that the bats largely avoid re-occupying roosts when highly contagious puparia are likely to be present as a result of previous occupations of the roosts by the bat colony. Our results indicate that the females adapted their roosting behaviour to the age-dependent contagiousness (emergence probability) of the puparia. However, some infested roosts were re-occupied, which we assume was because these roosts provided advantages to the bats (e.g. a beneficial microclimate) that outweighed the negative effects associated with bat fly infestation. We suggest that roost selection in Bechstein’s bats is the outcome of a trade-off between the costs of parasite infestation and beneficial roost qualities.  相似文献   

12.
Hibernation is a physiological adaptation that allows animals to survive adverse environmental conditions. A commonly assumed cost of hibernation is impaired memory retention. So far, however, the effects of hibernation on memory retention have been assessed on only a few behavioral tasks, and exclusively under laboratory conditions. Taking advantage of the longevity and strict colony fidelity of female Bechstein's bats, we were able to evaluate memory retention in the same individuals over two consecutive summers in the field. We used a pairwise roost choice experiment with automatic monitoring of RFID-tagged bats. Roosts’ suitability as day roost was associated with a distinctive external echo-acoustic cue. Experiments were separated by a natural hibernation period of eight months. We determined associative learning and memory retention by comparing the bats’ proportion of visits to suitable roosts within and between breeding seasons, respectively. During the first breeding season, bats quickly learned to associate the suitable roosts with their external cue. After hibernating, we found no evidence that individuals remembered the association between the roosts’ suitability and their respective external cue, suggesting a lack of memory retention. Nevertheless, bats quickly re-learned the same association during the second breeding season, emphasizing the high behavioral flexibility of Bechstein's bats.  相似文献   

13.
Roost microclimate plays an important role in the survival, growth and reproduction in microbats. Entering torpor is one of the main energy saving mechanisms commonly used by microbats. The use of torpor is affected by roost microclimate and seasonally differs between the two sexes in relation to their reproductive condition. Consequently, thermal properties of male and female roosts should differ. To test this hypothesis, we compared temperature parameters of two anthropogenic day roosts of Daubenton’s bats with a different structure of the population inhabiting them. In accordance with our predictions, the roost occupied by a male-dominated colony was colder and more fluctuant than the maternity roost with a female-dominated population. However, using of the two roosts changed during the season in response to changing energetic demands of the two sexes. While males were almost absent in the warmer maternity roost during pregnancy and lactation, they appeared in this roost during the post-lactation when mating starts. In contrast, females did not use the colder (male) roost until the time of weaning of juveniles, i.e., the time when their thermoregulatory needs change and they may benefit from using colder roost. Our study provides the evidence that the same roost may be used by individuals of different sex and reproductive state in different periods of the year. Generalizations about roost selection without knowledge of temporal variation in roost use and microclimatic conditions should be taken with caution. Anthropogenic roosts may be advantageous to Daubenton’s bats as these can provide a variety of suitable microclimates and/or more space for roosting than tree cavities.  相似文献   

14.
Theodore H.  Fleming 《Ibis》1981,123(4):463-476
This study presents data on the roosting and feeding behaviour of Pied Wagtails around Oxford, England. During the winter of 1977–78, from two to 1200 wagtails roosted in a Phragmites reed-bed. Use of this roost was greatest during mild, windless weather and the birds apparently used alternate roosts during harsh weather. Movement between roosts sometimes occurred between sunset and sunrise. Morning ‘departure group’ size, number leaving per unit time and diversity of departure directions increased with roost size. Wagtails quickly left the vicinity in the morning. In the afternoon, they joined one or more pre-roost gatherings before entering the roost for the night. Behaviour upon arrival at the roost was variable: birds might enter the reeds quickly or circle in large groups before landing. Aerial revolutions and generally ‘restless’ behaviour often accompanied increases in roost size. Wagtail feeding rates varied significantly between and within habitats. Number of wagtails feeding on the flooded Port Meadow, located 2–3 km south of the main roost, varied from about five to over 60 on different days; these numbers were not correlated with feeding rates. In contrast, the number of birds feeding at a sewage farm was nearly constant all winter. Some wagtails show high fidelity to feeding areas but others do not. Five short-term food supplementation experiments indicated that wagtails knowledgeable about a dense food source are not followed in the morning by naive birds. Results of this study are discussed in relation to the predation, physiology and information centre hypotheses that have been suggested to explain communal roosting in birds. I conclude that the communal roosting system of Pied Wagtails has physiological and anti-predator functions. Wagtails appear to choose certain roosts because of the protection that they provide from adverse climate and predation.  相似文献   

15.
For habitat specialists, fragmentation has major consequences as it means less suitable habitat for the species to live in. In a fragmented landscape, we would expect larger, but spatially more clustered, foraging ranges. We studied the impact of landscape fragmentation on the foraging range and habitat exploitation of a specialised forest bat by radiotracking 16 female lesser horseshoe bats Rhinolophus hipposideros in a landscape with connected woodland structures and in a highly fragmented landscape in Carinthia, Austria.Contrary to our expectations, spatial foraging behaviour was not influenced by fragmentation. No differences in the behaviour of the bats between the sites were evident for the foraging ranges (minimum convex polygon, MCP), the core foraging areas (50% kernel), nor the mean or the maximum distances from the roost. However, in the highly fragmented landscape, the foraging activity of individuals was spatially more clustered and the overall MCP of all bats of a colony was greater compared to the less fragmented landscape.Woodland was the most important foraging habitat for the lesser horseshoe bats at both study sites. Habitat selection at the individual MCPs was evident only at the site with low fragmentation. However, in the core foraging areas, woodland was significantly selected over all other habitat types at both study sites.We conclude that (1) conservation measures for colonies of lesser horseshoe bats should be undertaken within 2.5 km of the nursery roost, (2) woodland is the key foraging habitat particularly in the vicinity of the roost, and (3) any loss of woodland near the colonial roosts are likely to negatively influence the colony, since these bats do not seem to be able to adapt their spatial foraging behaviour in a degraded landscape. The inflexible spatial behaviour of this specialised bat highlights the need to compensate for any habitat loss within the foraging range of a bat colony.  相似文献   

16.
We studied the roosting ecology of the long-tailed bat (Chalinolobus tuberculatus) during the springautumn months from 1998–2002 at Hanging Rock in the highly fragmented landscape of South Canterbury, South Island, New Zealand. We compared the structural characteristics and microclimates of roost sites used by communally and solitary roosting bats with those of randomly available sites, and roosts of C. tuberculatus occupying unmodified Nothofagus forest in the Eglinton Valley, Fiordland. Roosting group sizes and roost residency times were also compared. We followed forty radio-tagged bats to 94 roosts (20% in limestone crevices, 80% in trees) at Hanging Rock. Roosts were occupied for an average of 1 day and 86% were only used once during the study period. Colony size averaged 9.8 ± 1.1 bats (range 2–38) and colonies were dominated by breeding females and young. Indigenous forest, shrubland remnants and riparian zones were preferred roosting habitats. Communally roosting bats selected roosts in split trunks of some of the largest trees available. Selection of the largest available trees as roost sites is similar to behaviour of bat species occupying unmodified forested habitats. Temperatures inside 12 maternity roosts measured during the lactation period were variable. Five roosts were well insulated from ambient conditions and internal temperatures were stable, whereas the temperatures inside seven roosts fluctuated in parallel with ambient temperature. Tree cavities used by bats at Hanging Rock were significantly nearer ground level, had larger entrance dimensions, were less well insulated, and were occupied by fewer bats than roosts in the Eglinton Valley. These characteristics appear to expose their occupants to unstable microclimates and to a higher risk of threats such as predation. We suggest that roosts at Hanging Rock are of a lower quality than those in the Eglinton Valley, and that roost quality may be one of the contributory factors in the differential reproductive fitness observed in the two bat populations. The value of introduced willows (especially Salix fragilis) as bat roosts should be acknowledged. We recommend six conservation measures to mitigate negative effects of deterioration of roosting habitat: protection and enhancement of the quality of existing roosts, replanting within roosting habitat, provision of high quality artificial roosts, predator control, and education of landowners and statutory bodies.  相似文献   

17.
Conflict can arise when bats roost in human dwellings and householders are affected adversely by their presence. In the United Kingdom, the exclusion of bats from roosts can be licensed under exceptional circumstances to alleviate conflict, but the fate of excluded bats and the impact on their survival and reproduction is not well understood. Using radio-tracking, we investigated the effects of exclusion on the soprano pipistrelle Pipistrellus pygmaeus, a species that commonly roosts in buildings in Europe. Exclusions were performed under licence at five roosts in England in spring, when females were in the early stages of pregnancy. Following exclusion, all bats found alternative roosts and colonies congregated in nearby known roosts that had been used by radio-tagged bats prior to exclusion. We found no difference in roosting behaviour before and after exclusion. Both the frequency of roost switching and the type of roosts used by bats remained unchanged. We also found no change in foraging behaviour. Bats foraged in the same areas, travelled similar distances to reach foraging areas and showed similar patterns of habitat selection before and after exclusion. Population modelling suggested that any reduction in survival following exclusion could have a negative impact on population growth, whereas a reduction in productivity would have less effect. While the number of soprano pipistrelle exclusions currently licensed each year is likely to have little effect on local populations, the cumulative impacts of licensing the destruction of large numbers of roosts may be of concern.  相似文献   

18.
Social dynamics are an important but poorly understood aspect of bat ecology. Herein we use a combination of graph theoretic and spatial approaches to describe the roost and social network characteristics and foraging associations of an Indiana bat (Myotis sodalis) maternity colony in an agricultural landscape in Ohio, USA. We tracked 46 bats to 50 roosts (423 total relocations) and collected 2,306 foraging locations for 40 bats during the summers of 2009 and 2010. We found the colony roosting network was highly centralized in both years and that roost and social networks differed significantly from random networks. Roost and social network structure also differed substantially between years. Social network structure appeared to be unrelated to segregation of roosts between age classes. For bats whose individual foraging ranges were calculated, many shared foraging space with at least one other bat. Compared across all possible bat dyads, 47% and 43% of the dyads showed more than expected overlap of foraging areas in 2009 and 2010 respectively. Colony roosting area differed between years, but the roosting area centroid shifted only 332 m. In contrast, whole colony foraging area use was similar between years. Random roost removal simulations suggest that Indiana bat colonies may be robust to loss of a limited number of roosts but may respond differently from year to year. Our study emphasizes the utility of graphic theoretic and spatial approaches for examining the sociality and roosting behavior of bats. Detailed knowledge of the relationships between social and spatial aspects of bat ecology could greatly increase conservation effectiveness by allowing more structured approaches to roost and habitat retention for tree-roosting, socially-aggregating bat species.  相似文献   

19.
Many aspects of animal behaviour are affected by real‐time changes in the risk of predation. This conclusion holds for virtually all taxa and ecological systems studied, but does it hold for bats? Bats are poorly represented in the literature on anti‐predator behaviour, which may reflect a lack of nocturnal predators specialized on bats. If bats actually experience a world with minimal anti‐predator concerns, then they will provide a unique contrast within the realm of vertebrate ecology. Alternatively, such predator‐driven behaviour in bats may not yet be fully understood, given the difficulties in working with these highly mobile and nocturnal animals. We provide a wide‐ranging exploration of these issues in bat behaviour. We first cover the basic predator‐prey information available on bats, both on potential predators and the ways in which bats might perceive predators and respond to attacks. We then cover work relevant to key aspects of bat behaviour, such as choice of daytime roosts, the nature of sleep and torpor, evening roost departures, moonlight avoidance, landscape‐related movement patterns, and habitat selection. Overall, the evidence in favour of a strong influence of predators on bat behaviour is equivocal, with the picture clouded by contradictory results and a lack of information on potential predators and the perception of risk by bats. It seems clear that day‐active bats run a considerable risk of being killed by diurnal raptors, which are able to capture bats with relative ease. Thus, bats taking advantage of a pulse of insects just prior to sunset are likely taking risks to gain much‐needed energy. Further, the choice of daytime roosts by bats is probably strongly influenced by roost safety. Few studies, however, have directly addressed either of these topics. As a group, insectivorous temperate‐zone bats show no clear tendency to avoid apparently risky situations, such as activity on moonlit nights. However, some observations are consistent with the idea that predation risk affects choice of movement paths and feeding areas by temperate‐zone bats, as well as the timing of roost departures. The behaviour of tropical bats, on the other hand, seems more generally influenced by predators; this is especially true for tropical nectarivores and frugivores, but also for insectivorous bats. Presumably there are more serious predators on bats in the tropics (e.g. specialized raptors or carnivorous bats), but the identity of these predators is unclear. More information is needed to assess fully the influence of predators on bat behaviour. There is much need for work on the ways in which bats perceive predators via auditory, visual, and olfactory cues, and whether bats have some knowledge of the risks posed by different predators. Also needed is information on how predators attack bats and how bats react to attacking predators. Difficult to obtain, but of critical value, will be information on the nature of the predation risk experienced by bats while away from roosts and during the full darkness of night.  相似文献   

20.
Closely related, ecologically similar species often roost in distinctly different habitats, and roosting patterns also vary within species in relation to sex, age and season. The causes of such variation are not well understood at either a proximate or ultimate level. We studied communal roosting in two congeneric species of Prionostemma harvestmen at a rainforest site in Nicaragua. Previous research showed that Prionostemma sp. 1 forms male‐biased communal roosts in tree‐root cavities, while Prionostemma sp. 2 forms communal roosts of variable but temporally stable sex ratios on spiny palms. Here, we investigate potential mechanisms underlying variation in roosting site choice between and within these syntopic species. First, we present the results of a field experiment designed to probe the mechanism underlying skewed roost sex ratios in Prionostemma sp. 2. Previous studies have suggested that these harvestmen use conspecific scent to locate communal roosts and that new roosts can be established via group translocation. Therefore, to test the hypothesis that skewed roost sex ratios in this species arise from sex differences in scent marks, we translocated single‐sex groups of ca. 30 individuals to each of 20 previously unoccupied spiny palms. Female release sites attracted new recruits of both sexes, while male release sites attracted almost exclusively males. We infer that Prionostemma sp. 2 females preferentially roost in sites scent‐marked by females and that this mechanism is sufficient to explain the skewed roost ratios. Further adding to knowledge of Prionostemma roosting behavior, we show that Prionostemma sp. 1 forms female‐biased communal roosts on spiny palms, that some roosts contain both species, and that the species composition is stable on a time scale of at least 2 weeks. To the best of our knowledge, this study is the first experimental test of mechanisms underlying sexual segregation at communal roosts in any taxon.  相似文献   

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