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Most previous studies of the use bats make of their foraging areas have been concerned with general habitat preferences rather than with microhabitats. The present study focuses on microhabitat preference within three landscape features: linear landscape elements, ponds and rivers. The importance of linear landscape elements to bats was investigated by placing recording stations next to treelines, and others in adjacent open spaces approximately 35 m away. Most pipistrelle (Pipistrellus pipistrellus and P. pygmaeus) bat activity was recorded next to treelines and very little over open spaces. Bats used treelines for both commuting and foraging, but flew closer to treelines when commuting than when foraging. More insects were caught, and more pipistrelle feeding attempts were recorded close to treelines than further away. The relationship between the number and type of landscape elements leading to and surrounding ponds, and the use pipistrelle and Daubenton's (Myotis daubentonii) bats make of such ponds, was similarly investigated. Bats preferred to commute to ponds along woodland edges and streams, and not along hedgerows. More bat activity was recorded over ponds that had little overhanging and surrounding vegetation in comparison to ponds that had more, and over large wide ponds in comparison to small narrow ones. The extent to which pipistrelle and Daubenton's bats' use of river corridors extends beyond the water body was also investigated. Bat activity decreased with increasing distance from rivers, up to a distance of 70 m. River sites which were wooded on both sides attracted more bat activity for a longer duration than sites which had no trees on either side. Pipistrelles made use of the wider river corridor whereas Daubenton's bats restricted their activity almost exclusively to the water body.
Maternity colonies of serotine bats inhabiting large roof spaces were studied at three localities in southwestern Germany and Luxembourg. Bats of all three colonies returned to the maternity roosts during the second or third week of April, and were strongly philopatric to their main roost. Roost switching occurred only rarely and for short periods. Measurements of temperature inside two roosts showed gradients according to height and aspect of the roof space with a mean roost temperature of 22°C during gestation and lactation. The availability of adequate microclimates within the roost during gestation and lactation was thought to favour roost philopatry. Females of all ages and reproductive classes inhabit the roosts throughout summer. Mean (± SD) inferred gestation length was 52 ± 6 days, and variations were not related to roost temperatures during gestation. The mean date of birth was 16th June with most births occurring within a period of 14 ± 6 days and of young first emerged from the roost 36 ± 7 days after birth. Prolonged adverse weather conditions, leading to low roost temperatures, resulted in mortality of 11–27% of preweaning young in four of the five years studied. Since immigration occurs, the number of young emerging from the roost cannot be deduced from the number of females present during gestation or lactation. Adult females and young start to disperse after the first young are weaned at the beginning of August and are last observed in October. The implications of these results for the epidemiology of European bat lyssavirus 1 are discussed.
Natterer's bat Myotis nattereri is one of the least known European bats. Understanding its patterns of movement between roosts is an important aspect of assessing the relative value of different types of roost for conservation of the species. We determined patterns of movement of Natterer's bat between roosts by radio-tracking successive animals from the same colonies during summer (May to September). For one maternity colony comprising 65 adult females, the attic of a large mediaeval church was the main roost site, accounting for 88% of radio-located bat days. The two other maternity colonies tracked comprised about 35 adult females each and used from 15 to 25 roost sites, some containing multiple roosts. For these two colonies, up to six roost sites per colony accounted for about 80% of bat occupancy days in any one summer and for each colony, roost home ranges for roosts used by subgroups of at least two bats together covered 0.4 km2. Although bats made frequent movements between roosts there was no apparent interchange between adjacent colonies and no overlap in the range used by adjacent colonies. Bats changed roosts every 3.0 days on average, moving 510 m (median) to an alternative roost. They departed late and returned early to roosts. Colonies assembled or disintegrated into larger or smaller groups occupying diverse roosts during summer, but there was high social cohesion between colony members. Bats that separated into subgroups from mid-summer (mid-July) onwards later roosted together again. Natterer's bat exhibits high behavioural flexibility in the type of roosts used: of two adjacent maternity colonies, one used mostly tree roosts and the other mostly buildings. However, both roosted mainly in roofs during early summer (late May to mid-July). Tree roosts were significantly preferred to those in buildings when ambient external maximum temperatures were ≥ 30°C or mean temperatures fell below 14°C. Natterer's bat appears to depend on the availability of a number of roosts of different types. Parturition roosts, roosts in heavily timbered barns and roosts within core roosting areas, potentially up to 1.2 km distant from the parturition roost, should have highest conservation priority but conservation management should guard against any roost loss.
The number of orientation, feeding and social calls emitted by pipistrelle batsPipistrellus pipistrellus Schreber, 1774 andP. pygmaeus Leach, 1825 was recorded throughout the night at eight different sites. Social calls were unaffected by weather variables, whereas orientation calls and feeding buzzes were both significantly affected by cloud and temperature conditions. The number of emissions of each call type was significantly different between sites, indicating that the bats used different sites for different activities. Significant positive correlations between all three combinations of call types occurred only during the middle of the night, corresponding with the nadir of flying insects. This suggests that bats were engaged in activities other than feeding at this time, such as territory protection or mate attraction.
The behaviour and habitat associations of aerial insectivorous bats are poorly understood despite constituting up to 65% of bat species in the Neotropics. In 2003, 2004 and 2005 we quantified the activity of insectivorous bats and their insect prey at pastureland and forest sites with and without cenotes (water-filled sinkholes) in the Yucatan Peninsula, Mexico. We used a time-expansion bat detector to survey each habitat for 24 nights and analysed 2,880 one-minute recorded sequences to determine bat activity. We identified 14 species and five phonic types belonging to four families. Bat activity and the average number of bat species acoustically sampled each night were significantly greater in habitats with cenotes than in those without. Pteronotus personatus and an unidentified molossid were recorded exclusively at cenotes. Peropteryx macrotis showed the highest activity of all bat species. In all habitats insects were more abundant during the rainy season but only in pastureland was bat activity significantly greater during the rainy season. Insect abundance was correlated with bat activity only at cenotes in pastureland. Cenotes are important foraging habitats for insectivorous bats as 16 species, 84% of those revealed by this study, were recorded feeding in these habitats and the number of feeding buzzes was higher in comparison to habitats without cenotes. Protection of cenotes and their surrounding vegetation should be a management priority in order to conserve the high diversity of insectivorous bats associated with these distinctive habitats.
Time-expanded echolocation calls were recorded from 29 species of Neotropical bats in lowland moist tropical forest in Trinidad, West Indies with three aims: (1) to describe the echolocation calls of the members of a diverse Neotropical bat community, especially members of the family Phyllostomidae, whose calls are not well documented (2) to investigate whether multivariate analysis of calls allows species and foraging guilds to be identified and (3) to evaluate the use of bat detectors in surveying the phyllostomids of Neotropical forests. The calls of 12 species of the family Phyllostomidae are described here for the first time and a total of 29 species, belonging to five families (Emballonuridae, Mormoopidae, Phyllostomidae, Molossidae and Vespertilionidae) were recorded. Quadratic discriminant function analysis (DFA) was used to obtain classification rates for each one of 11 individual species and for six guilds (based on diet, foraging mode and habitat) comprising 26 species. Overall classification rates were low compared to similar studies conducted in the Palaeotropics. We suggest that this may be due to a combination of ecological plasticity for certain species and a loose relationship between echolocation call shape, fine-grained resource partitioning and resource acquisition in phyllostomids.
Bats constitute a substantial proportion of mammal diversity within the Asian tropics and subtropics and are particularly susceptible to population losses associated with human activities. This poses a conservation concern in Asian karst areas which support high bat species diversity, yet are experiencing habitat loss and degradation and increasing pressure from tourism and extractive industries. As disturbance during crucial reproductive periods (late pregnancy, lactation and weaning) threatens reproductive success, we investigated the reproductive phenology of a bat assemblage at two karst sites in North Vietnam. Our results indicate that the timing of major reproductive events coincides among two cave-dwelling pteropodids, and among 26 cave and foliage dwelling rhinolophids, hipposiderids and vespertilionids. March-July is the primary reproductive period for all insectivorous species sampled, and protection of maternity roosts during this time is critical. Reproduction in cave-dwelling pteropodids spanned a greater period (March-December), due to two birth periods each year. Lactation in the three insectivorous families studied was positively correlated with rainfall and temperature, with weaning occurring during the peak wet season. The strong congruence in reproductive phenologies in our results and climatic homogeneity of North Vietnam (18-23°N) suggests that our study may have wider applicability within the region. Vietnamese caves support high bat diversity which is likely threatened by harvesting for consumption and tourism development nationwide. Studies to investigate and address these threats should be given high priority.
We studied the foraging behaviour of the dawn bat Eonycteris spelaea, a cave roosting nectarivore widespread in SE Asia, and principal pollinator of economically important crops. We radio-tracked 17 individuals for five to 19 nights over a three month period. The bats were from three cave colonies in agricultural habitats in southern Thailand. They traveled between one and 17.9 km (fi01_307.gif ± SD: 4.4 km ± 5.07, median = 2.34) from their roosting cave to food sources. The mean home-range size of the individuals varied with the method used in its calculation from 518.4 ha (100% Minimum Convex Polygon, MCP) to 564.5 ha (100% Local Convex Hull method, LoCoH) and 460.8 ha (95% Kernel density estimation, KDE). The mean size of foraging areas used by the bats also varied according to the method of calculation from 14.26 ha (100% MCP), 13.25 ha (100% LoCoH) and 38.52 ha (95% KDE) and accounted for 21.9%, 20.08% and 40.5% of the respective home-range size. The bats foraged in one to three foraging areas each night. The greatest distance between feeding trees varied between 0.25 and 8 km (mean 1.25 km ± 2.19). Those bats with multiple foraging areas moved from patch to patch of Durio zibethinus and did not return to a previously visited patch, whereas those feeding on Parkia repeated their visits to several patches in a single night. Ninety percent of foraging areas used by the radio-tagged individuals were in managed habitat such as fruit orchards and yards of houses to which the bats maintained strong site fidelity.
We analysed 890 faecal samples from 145 molossid bats in eastern Madagascar during the austral summer and winter. Coleoptera, Hemiptera, Lepidoptera and Diptera were the most important sources of food for Mops leucostigma, Mormopterus jugularis and Chaerephon pumilus. The percentage volume of Hemiptera and Lepidoptera were similar in the diet, pooled across season, for all species but significant differences were found for Diptera and Coleoptera. Mops leucostigma, however, had the highest volume of Diptera and M. jugularis of Coleoptera. Hemiptera were an important food source for all species during both seasons, whereas Coleoptera were prevalent in the diet only during the summer. Diptera were rarely eaten by M. jugularis but constituted a major source of food for the other two species during the winter. Although there was little evidence of strong interspecific dietary partitioning, M. jugularis appeared to have a more limited dietary composition at the ordinal level. Major differences in dietary composition were between season rather than species at the ordinal level. Further investigations are recommended to assess the potential role of molossids in consuming economic pests of cotton in Madagascar.
Bats may be vulnerable to predation during evening emergence and morning return to their roosts. Early emergence increases the risk of exposure to raptorial birds, but emerging late confers a risk of missing the dusk peak of aerial insects. Here, both emergence and return activity was studied in detail at the same roosts for the first time. We investigated six maternity colonies of pipistrelle bats (Pipistrellus pipistrellus and P. pygmaeus) in NE Scotland and recorded light levels and time of emergence and return of the bats with respect to sunset and sunrise on the same nights. Parameters of return activity generally occurred at lower light intensities than those of emergence. Therefore, the interval between dawn return and sunrise was generally longer than that between sunset and dusk emergence. Emergence and return were equal in duration. Bats clustered more on emergence in comparison with return during pregnancy and lactation, whereas during postlactation this trend was reversed.
The use of different roost types by Daubenton's bats (Myotis daubentonii) during reproduction was investigated in two adjacent river valleys in northeastern Scotland. Forty-six individuals from six colonies were radiotracked during the summers of 2004–2006. The frequency of roost switching varied with reproductive status, and was lowest in lactating females and highest in non-reproductive females, which changed roosts on average once every 5.0 and 1.5 days, respectively. Although Daubenton's bats regularly switched roosts, strong faithfulness to the roosting area was apparent regardless of whether they formed maternity colonies in trees or buildings. Although most roosts found at both study areas were in trees, lactating females and juveniles in one valley roosted only in trees whereas in the other they roosted exclusively in buildings, in which ambient temperatures were significantly higher. The implications of roosting habits for the transmission of European bat lyssavirus are discussed.
Here we expand our previous study to provide more detailed information on the relationship between the male eastern sucker-footed bat Myzopoda aurita and the traveler's tree Ravenala madagascariensis in south-eastern Madagascar, during six month-long field work sessions carried out over two years. We caught 593 bats, 229 newly caught and 364 recaptures, exclusively males, roosting in 37 day roosts in the partially unfurled central leaves of R. madagascariensis. No bats were found in any other roosting situation. To analyse potential roost availability, we monitored partially unfurled central leaves on R. madagascarienis on four transects and 12% appeared suitable as M. aurita roosts. These leaves took three to 25 days to unfurl, and roosts became available between one and 19 days after unfurling commenced. Day roosts were occupied for one to 12 days. Bats were more likely to occupy roosts in taller trees. The size of roosting groups varied between one and 36 individuals. Movements of bats between roosts were recorded on 35 occasions and between two and nine individuals of M. aurita found in one roost were subsequently found together in a different roost. Myzopoda aurita occurs in degraded forests and anthropogenic habitats of eastern Madagascar where it may be affected by loss of roosts since R. madagascariensis is used extensively for building and thatching houses.
Otomops madagascariensis is a large (24–27 g) molossid bat endemic to Madagascar. Unlike its congener O. martiensseni, in nearby mainland Africa, little is known about its ecology although it appears to roost only in caves. It is only known from a few sites in the west and occupies a small percentage of the available caves. We studied roosting colonies in seven vertical erosion domes in the roof of a cave in Parc National Tsingy de Bemaraha during July and October 2003. We also captured bats as they emerged from and returned to a roost cave in the south. Female bats examined in the west during October and in the south during November were pregnant. In the roosting colonies, one group contained 57 pregnant females and five adult males. Most other groups also consisted of both sexes but three male-only groups were encountered in October. Diet consisted mainly of Lepidoptera and Coleoptera and there was variation between sites and study locations in the contribution of these prey types. Otomops madagascariensis is an obligate cave dweller that appears to be rare within its known range and should be a target species for conservation and research.
Spectral and temporal features of echolocation calls produced by 15 insectivorous bat species in three families from Madagascar are described. In addition we provide a library of bat vocalizations that can be used for acoustic inventories involving heterodyne and time-expansion bat detectors. Time-expanded recordings of calls from 153 bats from 15 species were analyzed using five commonly used temporal and frequency variables measured from spectrographs. Echolocation calls for six species (Scotophilus tandrefana, S. marovaza, Emballonura tiavato, Neoromicia spp., N. malagasyensis and Triaenops auritus) are described for the first time. A discriminant function analysis revealed that a function based on the five measured variables provided a correct overall classification of 82.2%. Three groups of echolocation calls based upon the temporal and frequency characteristics of calls are recognized. The Constant Frequency group consists of hipposiderids and Emballonura spp., the Frequency Modulated/Quasi-Constant Frequency group is dominated by vespertilionids, and one species, Myotis goudoti, is in the Frequency Modulated group. Further we describe the utility of using acoustic sampling in inventory and monitoring studies, and in investigations of habitat use.
We used roost searches, mist netting and acoustic sampling to investigate the habitats used by bats in Parc National de Mantadia and the Réserve Spéciale d'Analamazaotra, eastern Madagascar. Four species were caught in relatively intact humid forest (Myotis goudoti, Miniopterus manavi, Miniopterus majori and Emballonura atrata) two in agricultural land, Neoromicia matroka and Neoromicia melckorum, and one, Rousettus madagascariensis, in Eucalyptus plantations. Mormopterus jugularis, Chaerephon pumilus and Mops leucostigma were found roosting in buildings ca. three km from the humid forest. Acoustic sampling revealed that Neoromicia spp. and molossids were ubiquitous and were recorded from intact and degraded humid forest, Eucalyptus plantations and agricultural land. Myotis goudoti showed the strongest association with intact humid forest. Taxon richness, determined by acoustic sampling, was highest in humid forest but activity was highest in plantations and agricultural land. Mixed-habitat landscapes that surround protected forests and consist of a mosaic of regenerating forest, agriculture, wetlands, villages and plantations are important for bats and promote chiropteran diversity because they provide roosting and foraging sites for species that rarely use intact forest. The humid forests of eastern Madagascar have lower bat diversity than the island's western deciduous karst forests.
The diet of E. spelaea was determined for bats captured monthly between June 2002-June 2003 at a cave entrance in Songkhla Province, Southern Thailand. Faecal analysis and pollen collected from the bats’ fur were used to identify the plant species ingested. From 1,155 diet records from 506 samples, at least eleven plant taxa were identified. Individual bats feed on flowers of up to six plant species each night. Parkia spp. (34%) and Musa spp. (28%) have the highest percentage frequency followed by Eugenia spp. (9.4%), Oroxylum indicum (6.4%), Durio zibethinus (6.2%), Ceiba pentandra (5.5%), Sonneratia spp. (5.2%), while Cocos nucifera and an unknown plant species, made up a minor proportion (<2.5%). Parkia and Musa were the main dietary items of E. spelaea in nearly every month, while the remaining components of the diet varied seasonally. Durio spp. is an important contributor to the diet during March–April (39–42%). The results from pollen collected from fur generally corresponded with those from faecal analysis, but Musa spp. had a higher percentage frequency on the fur (34%) than Parkia spp. (23%). The dawn return patterns of the bats to their roosts differ significantly between sexes. Most mature males return early in the night while most females return at dawn. Earlier returning males were significantly heavier than those returning later. This return pattern is similar to that reported during the breeding period for those polygynous fruit bats that maintain a harem. Thus, these results suggest that E. spelaea may exhibit a resource-defence polygynous mating system.
We investigated habitat use by the endemic Malagasy bat Hipposideros commersoni in evergreen littoral rainforest during the wet season in 2006, in order to better inform conservation guidelines. We used radiotracking to locate roosting and foraging sites. Roosts, typically 5.4 ± 0.2 m from the ground, were always occupied by single bats and were found on branches of trees with a diameter at breast height of 8.2 ± 0.7 cm. Home range size was 31.8 ± 9.2 ha for males and 41.7 ± 12.9 ha for females. Roosts were always located within the foraging areas and only five (5.4%) of the 91 located were situated outside the sheltered littoral forest. Foraging bats made greatest use of natural, sheltered littoral forest and relatively few foraging sorties occurred beyond the forest edge. Females were not trapped during January and may undergo local movements at that time. There are no known caves in the vicinity of the study area and H. commersoni roosted only on trees. Previous studies in Madagascar have highlighted the importance of caves for bats and we now extend this to include tree roosts, within the evergreen rainforest.
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