Climate and weather impact timing of emergence of bats

Interest in forecasting impacts of climate change have heightened attention in recent decades to how animals respond to variation in climate and weather patterns. One difficulty in determining animal response to climate variation is lack of long-term datasets that record animal behaviors over decadal scales. We used radar observations from the national NEXRAD network of Doppler weather radars to measure how group behavior in a colonially-roosting bat species responded to annual variation in climate and daily variation in weather over the past 11 years. Brazilian free-tailed bats (Tadarida brasiliensis) form dense aggregations in cave roosts in Texas. These bats emerge from caves daily to forage at high altitudes, which makes them detectable with Doppler weather radars. Timing of emergence in bats is often viewed as an adaptive trade-off between emerging early and risking predation or increased competition and emerging late which restricts foraging opportunities. We used timing of emergence from five maternity colonies of Brazilian free-tailed bats in south-central Texas during the peak lactation period (15 June-15 July) to determine whether emergence behavior was associated with summer drought conditions and daily temperatures. Bats emerged significantly earlier during years with extreme drought conditions than during moist years. Bats emerged later on days with high surface temperatures in both dry and moist years, but there was no relationship between surface temperatures and timing of emergence in summers with normal moisture levels. We conclude that emergence behavior is a flexible animal response to climate and weather conditions and may be a useful indicator for monitoring animal response to long-term shifts in climate.     

Ongoing changes in migration phenology and winter residency at Bracken Bat Cave

Bats play an important role in agroecology and are effective bioindicators of environmental conditions, but little is known about their fundamental migration ecology, much less how these systems are responding to global change. Some of the world's largest bat populations occur during the summer in the south‐central United States, when millions of pregnant females migrate from lower latitudes to give birth in communal maternity colonies. Despite a relatively large volume of research into these colonies, many fundamental questions regarding their abundance—including their intra‐ and interseasonal variability—remain unanswered, and even estimating the size of individual populations has been a long‐running challenge. Overall, monitoring these bat populations at high temporal resolution (e.g., nightly) and across long time spans (e.g., decades) has been impossible. Here, we show 22 continuous years of nightly population counts at Bracken Cave, a large bat colony in south‐central Texas, enabling the first climate‐scale phenological analysis. Using quantitative radar monitoring, we found that spring migration and the summer reproductive cycle have advanced by approximately 2 weeks over the study period. Furthermore, we quantify the ongoing growth of a newly—established overwintering population that indicates a system‐wide response to changing environmental conditions. Our observations reveal behavioral plasticity in bats’ ability to adapt to changing resource availability, and provide the first long‐term quantification of their response to a changing climate. As aerial insectivores, these changes in bat phenology and propensity for overwintering indicate probable shifts in prey availability, with clear implications for pest management across wider regional agrisystems.     

Impact of landscape fragmentation on a specialised woodland bat,Rhinolophus hipposideros

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.     

Roosting and Foraging Social Structure of the Endangered Indiana Bat (Myotis sodalis)

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.     

Does the mode of transmission between hosts affect the host choice strategies of parasites? Implications from a field study on bat fly and wing mite infestation of Bechstein's bats

In a two-year field study, we analyzed the distribution of two hematophagous ectoparasites, the bat fly Basilia nana and the wing mite Spinturnix bechsteini , within and among 14 female colonies and among 26 solitary male Bechstein's bats Myotis bechsteinii . Our goal was to investigate whether differences in the transmission mode of the parasites, which result from differences in their life cycle, affect their distribution between host colonies and among host individuals within colonies. Bat flies deposit puparia in bat roosts, allowing for the transmission of hatched flies via successively shared roosts, independent of body contact between hosts or of hosts occupying a roost at the same time. In contrast, wing mites stay on the bat's body and are transmitted exclusively by contact of bats that roost together. As expected in cases of higher inter-colony transmissibility, bat flies were more prevalent among the demographically isolated Bechstein's bat colonies and among solitary male bats, as compared to wing mites. Moreover, the prevalence and density of wing mites, but not of bat flies, was positively correlated with colony size, as expected in cases of low inter-colony transmissibility. Within colonies, bat flies showed higher abundance on host individuals in good body condition, which are likely to have high nutritional status and strong immunity. Wing mites showed higher abundance on hosts in medium body condition and on reproductive females and juveniles, which are likely to have relatively weak immunity. We suggest that the observed infestation patterns within host colonies reflect different host choice strategies of bat flies and wing mites, which may result from differences in their inter-colony transmissibility. Our data also indicate that infestation with wing mites, but not with bat flies, might be a cost of sociality in Bechstein's bats.     

‘Compromise’ in Echolocation Calls between Different Colonies of the Intermediate Leaf-Nosed Bat (Hipposideros larvatus)

Each animal population has its own acoustic signature which facilitates identification, communication and reproduction. The sonar signals of bats can convey social information, such as species identity and contextual information. The goal of this study was to determine whether bats adjust their echolocation call structures to mutually recognize and communicate when they encounter the bats from different colonies. We used the intermediate leaf-nosed bats (Hipposideros larvatus) as a case study to investigate the variations of echolocation calls when bats from one colony were introduced singly into the home cage of a new colony or two bats from different colonies were cohabitated together for one month. Our experiments showed that the single bat individual altered its peak frequency of echolocation calls to approach the call of new colony members and two bats from different colonies adjusted their call frequencies toward each other to a similar frequency after being chronically cohabitated. These results indicate that the ‘compromise’ in echolocation calls might be used to ensure effective mutual communication among bats.     

Risk factors associated with mortality from white-nose syndrome among hibernating bat colonies

White-nose syndrome (WNS) is a disease responsible for unprecedented mortality in hibernating bats. First observed in a New York cave in 2006, mortality associated with WNS rapidly appeared in hibernacula across the northeastern United States. We used yearly presence-absence data on WNS-related mortality among hibernating bat colonies in the Northeast to determine factors influencing its spread. We evaluated hazard models to test hypotheses about the association between the timing of mortality and colony-level covariates, such as distance from the first WNS-affected site, colony size, species diversity, species composition and type of hibernaculum (cave or mine). Distance to origin and colony size had the greatest effects on WNS hazard over the range of observations; the type of hibernaculum and species composition had weaker effects. The distance effect showed a temporal decrease in magnitude, consistent with the pattern of an expanding epizootic. Large, cave-dwelling bat colonies with high proportions of Myotis lucifugus or other species that seek humid microclimates tended to experience early mortality. Our results suggest that the timing of mortality from WNS is largely dependent on colony location, and large colonies tend to be first in an area to experience high mortality associated with WNS.     

Factors driving population recovery of the greater horseshoe bat (<Emphasis Type="Italic">Rhinolophus ferrumequinum</Emphasis>) in the UK: implications for conservation

Although populations of many bat species appear to be recovering in some European countries, the extrinsic and intrinsic factors driving these increases have not yet been assessed. Disentangling the benefits of conservation management from other factors such as climate change is a crucial step for improving evidence-based conservation strategies. We used the greater horseshoe bat (Rhinolophus ferrumequinum) as a case study for understanding the recovery of bat populations, as its north-western populations have increased substantially over the past two decades following severe population declines. Using summer roost count data from the UK National Bat Monitoring Programme spanning an 18 year period from 1997 to 2014, we investigated the effects of (i) landscape characteristics associated with the implementation of the agri-environment schemes on colony trends and size, and (ii) meteorological variables on annual colony growth rate. We also assessed the relationship between colony size and colony growth to investigate intrinsic factors such as an Allee effect. Our results indicated that colony size was positively related to a range of landscape features (e.g. amount of broadleaf woodland and grassland, and density of linear features) surrounding the roost, while the amount of artificial light at night had a significant negative effect. Spring temperatures and precipitation (the latter with a lag of one year) were associated with annual colony growth. We also identified a negative density-dependence effect within colonies. Though the conservation of essential landscape elements may have contributed to population increases in the long-term, we conclude that recent population recovery has also been driven by climate change. Finally we recommend that the conservation of photophobic bat species such as R. ferrumequinum should focus on both the improvement of foraging/commuting habitats and the creation of dark areas.     

Mitochondrial DNA (mtDNA) reveals that female Bechstein's bats live in closed societies

We present a microgeographic analysis of mitochondrial DNA (mtDNA) in Bechstein's bats using three sources of control region sequence variability, including a novel mtDNA microsatellite, to assess individual relatedness both within and among 10 maternity colonies. Comparison of marker variability among 268 adult females revealed little genetic variability within each colony. However, most colonies were clearly distinguished by colony-specific mitochondrial haplotypes (total n = 28). Low intracolony variability and strong haplotype segregation among colonies, was reflected by an extraordinary high FST of 0.68, indicating a very low intercolony dispersal rate of approximately one female in five generations. Haplotype distribution among 18 solitary males showed that males frequently disperse between colony locations, indicating the absence of dispersal barriers. Bechstein's bat maternity colonies are thus closed groups that comprise 20-40 females probably belonging to only one or, at most, two matrilines. The genetic population structure of Bechstein's bats is in agreement with the hypothesis that females seek familiar and, at least, partially related cooperation partners for raising their young. Alternatively strong philopatry might reflect the importance of profound roost or habitat knowledge for successful reproduction in female Bechstein's bats.     

Ecological and anthropogenic drivers of rabies exposure in vampire bats: implications for transmission and control

Despite extensive culling of common vampire bats in Latin America, lethal human rabies outbreaks transmitted by this species are increasingly recognized, and livestock rabies occurs with striking frequency. To identify the individual and population-level factors driving rabies virus (RV) transmission in vampire bats, we conducted a longitudinal capture-recapture study in 20 vampire bat colonies spanning four regions of Peru. Serology demonstrated the circulation of RV in vampire bats from all regions in all years. Seroprevalence ranged from 3 to 28 per cent and was highest in juvenile and sub-adult bats. RV exposure was independent of bat colony size, consistent with an absence of population density thresholds for viral invasion and extinction. Culling campaigns implemented during our study failed to reduce seroprevalence and were perhaps counterproductive for disease control owing to the targeted removal of adults, but potentially greater importance of juvenile and sub-adult bats for transmission. These findings provide new insights into the mechanisms of RV maintenance in vampire bats and highlight the need for ecologically informed approaches to rabies prevention in Latin America.     

Larger trees may support larger Indiana bat maternity colonies in a dynamic landscape

Indiana bats (Myotis sodalis), federally listed as endangered, are of management concern in eastern North America. While researchers quantified the habitat affinities of the species throughout the range, few studies have occurred in regions where populations are at high risk for wind energy development and changing climes. Central Illinois, USA, is a dynamic landscape where forest area has been increasing in recent decades (on public and private land) because of changing farming practices and increased habitat protections. The increasing availability of large diameter trees, increasing forest biomass, and changing forest compositions have the potential to influence Indiana bat roost habitat preferences. We assessed Indiana bat maternity roost selection at the tree and forest plot scale to characterize patterns of use in this region from 2017–2018. We predicted that large trees on the landscape would support large colonies of Indiana bats. We located bats in multiple species of trees including elm (Ulmus spp.), cottonwood (Populus deltoides), and shagbark hickory (Carya ovata). We documented larger maternity colonies sharing roosts than in previous studies from the 1980s in the same region. We suggest managers and regulatory agencies monitor Indiana bats in dynamic landscapes such as those with changing forest composition and biomass.     

Genetic population structure of Natterer's bats explained by mating at swarming sites and philopatry

During autumn 'swarming', large numbers of temperate bats chase each other in and around underground sites. Swarming has been proposed to be a mating event, allowing interbreeding between bats from otherwise isolated summer colonies. We studied the population structure of the Natterer's bat (Myotis nattereri), a swarming species in northern England, by sampling bats at seven sites in two swarming areas and at 11 summer colonies. Analysis of molecular variance (amova) and genetic assignment analyses showed that the swarming areas (60 km apart) support significantly different populations. A negative correlation was found between the distance of a summer colony from a swarming area and the assignment of bats to that area. High gene diversity was found at all sites (HE = 0.79) suggesting high gene flow. This was supported by a low FST (0.017) among summer colonies and the absence of isolation by distance or substructure among colonies which visit one swarming area. The FST, although low, was significantly different from zero, which could be explained by a combination of female philopatry and male-mediated gene flow through mating at swarming sites with bats from other colonies. Modelling suggested that if effective size of the summer colonies (Ne) was low to moderate (10-30), all mating must occur at the swarming sites to account for the observed FST. If the Ne was higher (50), in addition to random mating during swarming, there may be nonrandom mating at swarming sites or some within-colony mating. Conservation of swarming sites that support potentially large populations is discussed.     

Estimating density of a forest-dwelling bat: a predictive model for Rafinesque’s big-eared bat

Data on species distribution and abundance are the foundation of population ecology. However, due to difficulties in surveying bats, abundance estimates for tree-roosting microchiropterans are non-existent. Therefore, our objective was to develop methods for estimating colony abundance and density, taking as our model Rafinesque’s big-eared bat Corynorhinus rafinesquii, a species of conservation concern found in cypress-gum swamps of the southeastern United States. We searched 123 transects at eight study sites in the Coastal Plain of Georgia, USA to locate and characterize diurnal summer roosts of C. rafinesquii. We modeled the relationship between the number of bat colonies and landscape-scale habitat variables with zero-inflated negative binomial regression and used Akaike’s information criterion to select the most parsimonious models. We generated a predictive density map to identify areas of high colony density and to estimate overall abundance. Colony density was predicted by the duration of wetland flooding, wetland width, and study site. Application of the regression model to a GIS indicated there were 3,734 colonies containing 6,910 adult bats on the eight study sites. Predicted density ranged from 0.07 colonies/ha and 0.07 adult bats/ha in saturated wetlands to 0.47 colonies/ha and 1.18 adult bats/ha in semi-permanently flooded wetlands. This study is the first to estimate density and abundance of forest-dwelling microchiropterans over a large area. Such data can serve as a baseline for future work on population trends in C. rafinesquii. In addition, our approach could be replicated for other bat species with moderately cryptic roosts.     

Seasonal bat activity related to insect emergence at three temperate lakes

Knowledge of aquatic food resources entering terrestrial systems is important for food web studies and conservation planning. Bats, among other terrestrial consumers, often profit from aquatic insect emergence and their activity might be closely related to such events. However, there is a lack of studies which monitor bat activity simultaneously with aquatic insect emergence, especially from lakes. Thus, our aim was to understand the relationship between insect emergence and bat activity, and investigate whether there is a general spatial or seasonal pattern at lakeshores. We assessed whole‐night bat activity using acoustic monitoring and caught emerging and aerial flying insects at three different lakes through three seasons. We predicted that insect availability and seasonality explain the variation in bat activity, independent of the lake size and characteristics. Spatial (between lakes) differences of bat activity were stronger than temporal (seasonal) differences. Bat activity did not always correlate to insect emergence, probably because other factors, such as habitat characteristics, or bats’ energy requirements, play an important role as well. Aerial flying insects explained bat activity better than the emerged aquatic insects in the lake with lowest insect emergence. Bats were active throughout the night with some activity peaks, and the pattern of their activity also differed among lakes and seasons. Lakes are important habitats for bats, as they support diverse bat communities and activity throughout the night and the year when bats are active. Our study highlights that there are spatial and temporal differences in bat activity and its hourly nocturnal pattern, that should be considered when investigating aquatic–terrestrial interactions or designing conservation and monitoring plans.     

Bat activity before and after a severe wildfire in Pilliga forests: Resilience influenced by fire extent and landscape mobility?

Wildfires are becoming increasingly severe and common in many parts of the world, yet their effects on certain taxa remain poorly studied. In November 2006, a major wildfire occurred in the Pilliga forests of north‐west New South Wales, Australia. This fire was severe and extensive, burning more than 120 000 ha. Prior to the fire, we conducted a study in the affected area assessing bat activity in relation to flyway size and riparian zones, allowing us to opportunistically compare before – and – after effects of wildfire on bats in burnt and unburnt areas. We assessed how bat activity and bat composition changed from 1 year pre‐fire to 1 year post‐fire and 4 years post‐fire, accounting for the effects of flyway size. A BACI analysis showed that there was an effect of time (year) on bat activity, with activity markedly reduced in both burnt and unburnt sites in the year post‐ fire, before recovering 4‐years post‐fire to levels that approached pre‐burn activity. However, there was no treatment or interaction effect, indicating fire alone did not affect bat activity at our sites. Activity was 50% lower at supplementary sites more remote (~5 km) from the fire edge. Multi‐dimensional scaling and permanova described a strong effect of year but no evidence of fire or stream size‐mediated effects on species composition. The drop in activity post‐fire coincided with a period of drought that subsequently broke in the year prior to our survey 4 years post‐fire. Some species‐specific responses to the fire were evident and depended on the proximity to unburnt forest. We suggest the potential for landscape re‐assortment, whereby some species may move from the unburnt landscape into the burnt landscape, possibly in response to a resource pulse of insects. A landscape pattern of movement could obscure the detection of direct fire effects on mobile bats. We suggest fire extent, proximity to unburnt vegetation and severity are key considerations when interpreting the response of bats to wildfire.     

A landscape perspective on bat foraging ecology along rivers: does channel confinement and insect availability influence the response of bats to aquatic resources in riverine landscapes?

River and riparian areas provide an important foraging habitat for insectivorous bats owing to high insect availability along waterways. However, structural characteristics of the riverine landscape may also influence the location of foraging bats. We used bat detectors to compare bat activity longitudinally along river reaches with contrasting channel confinement, ratio of valley floor width to active channel width, and riparian vegetation, and laterally with distance from the river along three different reach types. We measured rates of insect emergence from the river and aerial insect availability above the river and laterally up to 50-m into the riparian habitat in order to assess the relationship between food resources and insectivorous bat activity. Longitudinally, bat activity was concentrated along confined reaches in comparison to unconfined reaches but was not related to insect availability. Laterally, bats tracked exponential declines in aquatic insects with distance from the river. These data suggest that along the lateral dimension bats track food resources, but that along the longitudinal dimension channel shape and landscape structure determine bat distributions more than food resources.     

雄性菲菊头蝠夏季集群大小的变化及其影响因素

本研究以栖息于废弃防空洞的雄性菲菊头蝠(Rhinolophus pusillus)作为研究对象,探究其夏季集群大小变化及其潜在影响因素.应用红外热成像仪监测菲菊头蝠在6月到8月期间的集群大小.利用单因素方差分析和一般线性回归模型,检验菲菊头蝠集群大小的月变化及其与洞内微气候的关系.研究发现,菲菊头蝠集群大小从6月至8月...     

Bats are able to maintain long-term social relationships despite the high fission-fusion dynamics of their groups

Elephants, dolphins, as well as some carnivores and primates maintain social links despite their frequent splitting and merging in groups of variable composition, a phenomenon known as fission-fusion. Information on the dynamics of social links and interactions among individuals is of high importance to the understanding of the evolution of animal sociality, including that of humans. However, detailed long-term data on such dynamics in wild mammals with fully known demography and kin structures are scarce. Applying a weighted network analysis on 20,500 individual roosting observations over 5 years, we show that in two wild Bechstein's bat colonies with high fission-fusion dynamics, individuals of different age, size, reproductive status and relatedness maintain long-term social relationships. In the larger colony, we detected two stable subunits, each comprising bats from several family lineages. Links between these subunits were mainly maintained by older bats and persisted over all years. Moreover, we show that the full details of the social structure become apparent only when large datasets are used. The stable multi-level social structures in Bechstein's bat colonies resemble that of elephants, dolphins and some primates. Our findings thus may shed new light on the link between social complexity and social cognition in mammals.     

Summer tree roost selection by Rafinesque's big-eared bat

Roost requirements of most North American forest bats are well-documented, but questions remain regarding the ultimate mechanisms underlying roost selection. Hypotheses regarding roost selection include provision of a stable microclimate, space for large colonies, protection from predators, and proximity to foraging habitat, among others. Although several hypotheses have been proposed, specific mechanisms likely vary by species and geographic region. Rafinesque's big-eared bat (Corynorhinus rafinesquii) commonly roosts in trees with large basal hollows in the Coastal Plain of the southeastern United States. Our objective was to weigh evidence for hypotheses regarding selection of diurnal summer roosts by Rafinesque's big-eared bat at 8 study sites across the Coastal Plain of Georgia, USA. We used transect searches and radiotelemetry to locate roosts and measured 22 characteristics of trees, tree cavities, and surrounding vegetation at all occupied roosts and for randomly selected unoccupied trees. We evaluated 10 hypotheses using single-season occupancy models and used Akaike's information criterion to select the most parsimonious models. We located 170 tree roosts containing approximately 870 bats for our analysis. The best supported model predicted bat presence from cavity size, interior wall texture, and number of entrances. Because large cavities allow bats to fly and smooth walls impede attacks by terrestrial predators, our results are consistent with the hypothesis that bats select roosts that allow them to evade predators. However, data on predation rates are needed for a conclusive determination. Because trees suitable as roosts for Rafinesque's big-eared bat are rare in the landscape, protection of suitable forested wetland habitat is essential to provide current and long-term roost tree availability. © 2012 The Wildlife Society.     

Secretions of the interaural gland contain information about individuality and colony membership in the Bechstein's bat

Mammals use chemical signals for individual and kin recognition, to establish social hierarchies, mark territories and choose mates. The nocturnal and social lifestyle of bats suggests that, besides acoustic signals, they also use scent to communicate. We investigated in the communally breeding Bechstein's bat, Myotis bechsteinii, whether secretions of the facial interaural gland contain information that can be used for individual and colony recognition. Since female Bechstein's bats live in closed societies and show cooperative behaviour, we predicted they would recognize colony members. We analysed interaural gland secretions, which we repeatedly sampled from 85 females belonging to four free-ranging colonies. Gas chromatography/mass spectrometry profiles were individually specific and differed between colonies. Comparing odour profiles between colonies we found a relation between chemical similarity and the mitochondrial haplotype of colony members. Within colonies there was no correlation between mass spectrometer profile similarity and genetic relatedness. Our results suggest that female Bechstein's bats may use interaural gland secretions for individual and colony recognition but not to infer kinship directly.Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.