Identification of sympatric bat species by the echolocation calls

One hundred and thirty-eight echolocation calls of 63 free-flying individuals of five bat species (Rhinolophus ferrumequinum,Myotis formosus,Myotis ikonnikovi,Myotis daubentoni and Murina leucogaster)were recorded (by ultrasonic bat detector (D980)) in Zhi'an village of Jilin Province,China.According to the frequency-time spectra,these calls were categorized into two types:FM/CF (constant frequency) / FM (R.ferrumequinum) and FM (frequency modulated)(M.formosus,M.ikonnikovi,M.daubentoni and M.leucogaster).Sonograms of the calls of R.ferrumequinum could easily be distinguished from those of the other four species.For the calls of the remaining four species,six echolocation call parameters,including starting frequency,ending frequency,peak frequency duration,longest inter-pulse interval and shortest inter-pulse interval,were examined by stepwise discriminant analysis.The results show that 84.1% of calls were correctly classified,which indicates that these parameters of echolocation calls play an important role in identifying bat species.These parameters can be used to test the accuracy of general predictions based on bats' morphology in the same forest and can provide essential information for assessing patterns of bat habitat use.     

Identification of sympatric bat species by the echolocation calls

One hundred and thirty-eight echolocation calls of 63 free-flying individuals of five bat species (Rhinolophus ferrumequinum, Myotis formosus, Myotis ikonnikovi, Myotis daubentoni and Murina leucogaster) were recorded (by ultrasonic bat detector (D980)) in Zhi’an village of Jilin Province, China. According to the frequency-time spectra, these calls were categorized into two types: FM/CF (constant frequency) / FM (R. ferrumequinum) and FM (frequency modulated) (M. formosus, M. ikonnikovi, M. daubentoni and M. leucogaster). Sonograms of the calls of R. ferrumequinum could easily be distinguished from those of the other four species. For the calls of the remaining four species, six echolocation call parameters, including starting frequency, ending frequency, peak frequency duration, longest inter-pulse interval and shortest inter-pulse interval, were examined by stepwise discriminant analysis. The results show that 84.1% of calls were correctly classified, which indicates that these parameters of echolocation calls play an important role in identifying bat species. These parameters can be used to test the accuracy of general predictions based on bats’ morphology in the same forest and can provide essential information for assessing patterns of bat habitat use. __________ Translated from Journal of Northeast Normal University (Natural Science Edition), 2006, 38 (3): 109–114 [译自: 东北师范大学学报(自然科学版)]     

Plasticity in echolocation calls of <Emphasis Type="Italic">Myotis macrodactylus</Emphasis> (Chiroptera: Vespertilionidae): implications for acoustic identification

Poor knowledge of the intraspecific variability in echolocation calls is recognized as an important limiting factor for the accurate acoustic identification of bats. We studied the echolocation behaviors of an ecologically poorly known bat species, Myotis macrodactylus, while they were commuting in three types of habitats differing significantly in the amount of background clutter, as well as searching for prey above the water surface in a river. Results showed that M. macrodactylus altered their echolocation call structure in the same way during commuting as foraging bats do in relation to the changing level of clutter. With increasing level of clutter, M. macrodactylus generally produced echolocation calls with higher start, end, and peak frequencies; wider bandwidth; and shorter pulse duration. Compared to commuting, bats emitted significantly lower frequency calls with narrower bandwidth while searching for prey. Discriminant function analysis indicated that 79.8% of the calls from the three commuting habitats were correctly grouped, and 87% of the calls were correctly classified to the commuting and foraging contexts. Our finding has implications for those who would identify species by their calls.     

Flight performance, foraging tactics and echolocation in the trawling insectivorous bat Myotis adversus (Chiroptera: Vespertilionidae)

The bat Myotis adversus hunts for prey by aerial hawking and by taking prey from the water surface with its feet (trawling). The flight performance and echolocation of this species were studied in Queensland, Australia, and comparisons were made with Myotis daubentoni , a bat filling a similar ecological niche in the Palaearctic Region. The bats foraged in very similar ways, using the same foraging tactics and feeding in similar habitats, yet they were not geometrically similar in shape. The slightly larger Myotis adversus had relatively larger wings than M. daubentoni , conferring a slightly lower wing-loading. Nevertheless, M. adversus flew faster than M. daubentoni during the searching phase of foraging. Myotis daubentoni turned in tighter circles than M. adversus . Both species used short frequency-modulated (FM) echolocation calls of a characteristic sigmoidal structure, and nulls typically observed in the calls were an observational artefact. Myotis adversus also adopted an unusual 'long'FM call while foraging. The relations between echolocation frequencies and body size were explored in male M. adversus . Specialized morphological and acoustic adaptations for prey capture by trawling in insectivorous bats are discussed.     

Characteristics of echolocating bats’auditory stereocilia length, compared with other mammals

The stereocilia of the Organ of Corti in 4 different echolocating bats, Myotis adversus, Murina leuco-gaster, Nyctalus plancyi (Nyctalus velutinus), and Rhinolophus ferrumequinum were observed by using scanning electron microscopy (SEM). Stereocilia lengths were estimated for comparison with those of non-echolocating mammals. The specialized lengths of outer hair cells (OHC) stereocilia in echolocating bats were shorter than those of non-echolocating mammals. The specialized lengths of inner hair cells (IHC) stereocilia were longer than those of outer hair cells stereocilia in the Organ of Corti of echolocating bats. These characteristics of the auditory stereocilia length of echolocating bats represent the fine architecture of the electromotility process, helping to adapt to high frequency sound and echolocation.     

Calls in the Forest: A Comparative Approach to How Bats Find Tree Cavities

Although tree cavities are a particularly critical resource for forest bats, how bats search for and find new roosts is still poorly known. Building on a recent study on the sensory basis of roost finding in the noctule (Ruczynski et al. 2007), here we take a comparative approach to how bats find roosts. We tested the hypothesis that species' flight abilities and echolocation call characteristics play important roles in how well and by which cues bats find new tree roosts. We used the very manoeuvrable, faintly echolocating brown long-eared bat ( Plecotus auritus ) and the less manoeuvrable, louder Daubenton's bat ( Myotis daubentonii ) as study species. The species are sympatric in European temperate forests and both roost in tree cavities. We trained bats in short-term captivity to find entrances to tree cavities and experimentally manipulated the sensory cues available to them. In both species, cue type influenced the search time for successful cavity detection. Visual, olfactory and temperature cues did not improve the bats' performance over the performance by echolocation alone. Eavesdropping on conspecific echolocation calls played back from inside the cavity decreased search time in Daubenton's bat ( M. daubentonii ), underlining the double function of echolocation signals – orientation and communication. This was not so in the brown long-eared bat ( P. auritus ) that has low call amplitudes. The highly manoeuvrable P. auritus found cavities typically from flight and the less manoeuvrable M. daubentonii found more entrances during crawling. Comparison with the noctule data from Ruczyński et al. (2007) indicates that manoeuvrability predicts the mode of cavity search. It further highlights the importance of call amplitude for eavesdropping and cavity detection in bats.     

湖南省蝙蝠新纪录——大足鼠耳蝠

于湖南省永州市铜岩洞采集到14只蝙蝠,经鉴定为大足鼠耳蝠(Myotis ricketti),为湖南省翼手目新纪录.本文报道了该物种体型和头骨特征,并与浙江和安徽样本进行比较.大足鼠耳蝠发出典型的下扫调频声波,其低峰频、短时程的声波特征以及强壮的后足和锋利的爪等形态特征与其在水面"拖网式(trawhng)"捕食鱼类的捕食...     

Support for the allotonic frequency hypothesis in an insectivorous bat community

The allotonic frequency hypothesis proposes that certain insectivorous bat species can prey upon moths that can hear bat echolocation calls by using echolocation frequencies outside the sensitivity range of moth ears. The hypothesis predicts that the peak frequencies of bat echolocation calls are correlated with the incidence of moths in the diets of these bats. The aim of this study was to test this prediction on a bat community dominated by bats using low duty cycle echolocation calls, i.e. aerial foraging, insectivorous species using frequency modulated calls. The community consisted of nine species, two molossids, Sauromys petrophillus and Tadarida aegyptiaca, five vespertilionids, Eptesicus capensis, Eptesicus hottentotus, Miniopteris schreibersii, Myotis tricolor, and Myotis lesueuri, one rhinolophid, Rhinolophus clivosus, and one nycterid, Nycteris thebaica. The insect fauna in the habitat used by the bat community was suited to the testing of the allotonic frequency hypothesis because more than 90% of the moths comprising the insect fauna were tympanate. These included Pyralidae (3.8%), Geometridae (44.9%), Notodontidae (3.8%), Arctiidae (4.6%), Lymantriidae (0.8%) and Noctuidae (32.4%). As predicted, peak echolocation frequency was correlated with the incidence of moths in the diets of these nine species (r=0.98, df=7, P<0.01). Furthermore, multivariate analysis revealed that echolocation frequency (t=9.91, n=129, P<0.001) was a better predictor of diet than forearm length (t=5.51, n=129, P<0.001) or wing area (t=-3.41, n=129, P<0.001). This suggests that the selection pressure exerted by moth hearing might have acted directly on call frequency and secondarily on body size and wing morphology, as part of the same adaptive complex. It is unlikely that dietary differences were due to temporal and spatial differences in the availability of prey because the pattern of differences in skull morphology of the nine species supported our dietary analyses. The skull morphology of a bat represents a historical record of the kind of diet it has become adapted to over its evolutionary history. These results suggest that prey defences may mediate other factors structuring bat communities, e.g. competition. Competition may be reduced for those species of bats that can circumvent prey defences.     

广东省南岭新纪录种中管鼻蝠的形态测量、核型及超声波数据

2009年9月在广东省南岭采集到5只森林型蝙蝠,其鼻孔突出成短管状,背部毛棕褐色,前臂长34.3～36.8mm;核型为2n=44,FN=50.经鉴定为蝙蝠科管鼻蝠亚科的中管鼻蝠(Murina huttonii),为广东省翼手类分布新纪录.用蝙蝠超声波接收器(Anabat Ⅱ)录制并分析其回声定位声波,为FM型.中管鼻蝠...     

Use of foraging habitats by bats in a Mediterranean area determined by acoustic surveys: conservation implications

We determined habitat use by foraging bats by broad-band acoustic surveys in 10 habitat types from a Mediterranean area (southern Italy). We applied discriminant functions to identify time-expanded echolocation calls from free-flying bats.
Moon phase and cloud cover had no effect on bat activity. Only Hypsugo savii was influenced by temperature, and activity of Myotis daubentonii and Myotis capaccinii was reduced at higher wind speeds. Both total numbers of bat passes and feeding buzzes were highest over rivers and lakes. Pipistrellus kuhlii and H. savii were most frequently recorded. Pipistrellus kuhlii , Pipistrellus pipistrellus and Tadarida teniotis proved generalists in using foraging habitats.
Water sites and conifer plantations were respectively the most and the least used habitats by H. savii . Rivers were especially important to Myotis bats, Miniopterus schreibersii and Pipistrellus pygmaeus . Unlike P. kuhlii , P. pipistrellus was frequent in beech woodlands; P. pygmaeus made a considerable use of chestnut woodlands and Myotis spp. were moderately active in both these woodland types.
A large number of endangered or vulnerable species featured in riparian habitats, broadleaved woodlands and olive groves. Riparian and woodland habitats constitute an important target for conservation. Typical land use forms such as woodlands used for chestnut production and traditionally managed olive groves should be encouraged in conservation plans. The negative impact of urbanisation on bats might be counteracted by fostering trees, gardens and small cultivated patches. Farmland practices should encourage landscape complexity and limit the use of pesticides.     

Use of Forest Edges by Bats in a Managed Pine Forest Landscape

ABSTRACT Forest edges often have increased species richness and abundance (edge effect) and affect spatial behaviors of species and dynamics of species interactions. Landscapes of intensively managed pine (Pinus spp.) stands are characterized by a mosaic of patches and linear forest edges. Managed pine forests are a primary landscape feature of the southeastern United States, but the effects of intensive management on bat communities are poorly understood. Insectivorous bats are important top predators in nocturnal forest food webs. We examined bat foraging behavior along forest edges and in 4 structurally distinct stand types (open-canopy pine, prethinned pine, thinned pine, and unmanaged forest) within a managed pine forest in the coastal plain of North Carolina, USA. During May-August, 2006 and 2007, we recorded echolocation calls using Pettersson D240X bat detectors linked to digital recorders at 156 sites. We also sampled nocturnal flying insects at each site using Malaise insect traps. We used negative binomial count regression models to describe bat foraging behavior relative to forest edges, stand types, and prey availability. Although some species showed affinities for certain stand types and prey items, bat activity patterns were most strongly related to forest edges. Edges were used extensively by 6 aerial-hunting bat species, but avoided by Myotis species. Forest edges function similarly to natural forest gaps, by providing foraging opportunities for aerial-hunting bat species. Therefore, the maintenance of forest edges in managed pine landscapes may enhance foraging habitat for aerial-hunting bat species.     

马铁菊头蝠(Rhinolophus ferrumequinum)回声定位声波与生境类型和环境因子的关系

2007年在吉林省罗通山自然保护区,利用超声波探测仪（Avisoft-SASLAB PRO）录制并分析不同生境中马铁菊头蝠的回声定位声波。结果显示马铁菊头蝠在不同类型生境中活动;各生境中回声定位声波参数存在显著差异（one-way ANOVA,P〈0.05）。从环境因子中通过主成分分析筛选出与其回声定位声波相关的植被、气候和地形因子,探讨回声定位声波与这些因子的相关性。结果显示FM1和FM2带宽与乔木高（r=-0.948,-0.825;P〈0.05）、FM1起始频率和FM2终止频率与林冠面积（r=-0.967,-0.958;P〈0.05）、FM1起始频率、FM2终止频率和峰频与湿度（r=-0.776、-0.875和-0.794,P〈0.05）、脉冲持续时间和脉冲间隔与平均灌木高均呈显著负相关（r=-0.911,-0.990;P〈0.05）,峰频与植被株数（r=0.756,P〈0.05）、脉冲持续时间与冠下高呈显著正相关（r=0.870,P〈0.05）。表明各种环境因子（植被因子、气候因子和地形因子）都在一定程度上影响回声定位声波,回声定位声波具有表型可塑性和生境适应性,这些特性决定了马铁菊头蝠生境利用的程度和可利用的资源。     

Female greater wax moths reduce sexual display behavior in relation to the potential risk of predation by echolocating bats

Female greater wax moths Galleria mellonella display by wing fanning in response to bursts of ultrasonic calls produced bymales. The temporal and spectral characteristics of these callsshow some similarities with the echolocation calls of batsthat emit frequency-modulated (FM) signals. Female G. mellonellatherefore need to distinguish between the attractive signalsof male conspecifics, which may lead to mating opportunities,and similar sounds made by predatory bats. We therefore predictedthat (1) females would display in response to playbacks of male calls; (2) females would not display in response to playbacksof the calls of echolocating bats (we used the calls of Daubenton'sbat Myotis daubentonii as representative of a typical FM echolocatingbat); and (3) when presented with male calls and bat callsduring the same time block, females would display more whenperceived predation risk was lower. We manipulated predationrisk in two ways. First, we varied the intensity of bat callsto represent a nearby (high risk) or distant (low risk) bat.Second, we played back calls of bats searching for prey (lowrisk) and attacking prey (high risk). All predictions weresupported, suggesting that female G. mellonella are able todistinguish conspecific male mating calls from bat calls, andthat they modify display rate in relation to predation risk.The mechanism (s) by which the moths separate the calls ofbat and moth must involve temporal cues. Bat and moth signalsdiffer considerably in duration, and differences in durationcould be encoded by the moth's nervous system and used in discrimination.     

Echolocation calls of Rhinolophus ferrumequinum in relation to habitat type and environmental factors

The present experiment was carried out in Luotong Mountain Natural Reserve in Jilin Province of China in 2007. We recorded and analyzed the echolocation calls of Rhinolophus ferrumequinum in different habitats by using Avisoft Bioacoustics USG 116 and Avisoft-SASLAB PRO (Avisoft Bioacoustics, Berlin, Germany). Our results showed that R. ferrumequinum foraged in diverse habitats in the study area, and their echolocation calls were significantly variable in different habitats (One-Way ANOVA, P < 0.05). Vegetative, climatic and topographical factors were selected by using the principal component analysis and the correlations between the parameters of echolocation calls and these environmental factors were analyzed. The results indicated that although R. ferrumequinum always emitted FM/CF/FM echolocation calls in different habitats, the parameters of echolocation calls varied with variable environmental factors. Significant negative correlation existed between FM1 bandwidth and arbor height (r = ?0.948, P < 0.05), FM₂ bandwidth and arbor height (r = ?0.825; P < 0.05), FM1 starting frequency and canopy area (r = ?0.967, P < 0.05), FM₂ ending frequency and canopy area (r = ?0.958, P < 0.05), FM1 starting frequency and air relative humidity (r = ?0.776, P < 0.05), FM₂ ending frequency and air relative humidity (r = ?0.875, P < 0.05), peak frequency and air relative humidity (r = ?0.794, P < 0.05), pulse duration and average shrub height (r = ?0.911, P < 0.05), and inter-pulse interval and average shrub height (r = ?0.990, P < 0.05). Significant positive correlation existed between peak frequency and number of plants (r = 0.756, P < 0.05), and pulse duration and height below the canopy (r = 0.870, P < 0.05). Our results suggested that many kinds of ecological factors (such as vegetation factor, climatic factor and topographical factor) affected the structure of echolocation calls and made them diverse in different habitats, i.e., echolocation calls of bats had phenotypic flexibility and eco-adaptability. These characteristics determined the degree of available habitats and natural resources for R. ferrumequinum.     

Echolocation calls produced by Kuhl's pipistrelles in different flight situations

Flexibility in the echolocation call structure of bats can improve their performances, because, in some situations, some signal designs are better than others. Hence, at least some bats should adjust their echolocation calls according to the setting in which they are operating but also to the specific task at hand, that is their behavioral intention. We studied variation in the echolocation calls of Pipistrellus kuhlii emitted during four flight situations that were similar in setting but differed in behavioral context: emergence from a roost, commuting to and from foraging sites, foraging and returning to a roost. Echolocation calls produced by P. kuhlii differed significantly according to the flight situation. Call types differed most distinctly between foraging and commuting. We also found a high variance in the emergence calls we recorded, perhaps reflecting pre- and post-takeoff calls. Discriminant function analysis on calls emitted while foraging, commuting or returning to the roost classified the calls to the correct group 73.3% of the time. The differences between bats' echolocation calls in different flight situations might indicate an intrinsic change in the bat's behavior. Recognizing these differences could be crucial when using call variables to identify bat species.     

Weather conditions determine attenuation and speed of sound: Environmental limitations for monitoring and analyzing bat echolocation

Echolocating bats are regularly studied to investigate auditory‐guided behaviors and as important bioindicators. Bioacoustic monitoring methods based on echolocation calls are increasingly used for risk assessment and to ultimately inform conservation strategies for bats. As echolocation calls transmit through the air at the speed of sound, they undergo changes due to atmospheric and geometric attenuation. Both the speed of sound and atmospheric attenuation, however, are variable and determined by weather conditions, particularly temperature and relative humidity. Changing weather conditions thus cause variation in analyzed call parameters, limiting our ability to detect, and correctly analyze bat calls. Here, I use real‐world weather data to exemplify the effect of varying weather conditions on the acoustic properties of air. I then present atmospheric attenuation and speed of sound for the global range of weather conditions and bat call frequencies to show their relative effects. Atmospheric attenuation is a nonlinear function of call frequency, temperature, relative humidity, and atmospheric pressure. While atmospheric attenuation is strongly positively correlated with call frequency, it is also significantly influenced by temperature and relative humidity in a complex nonlinear fashion. Variable weather conditions thus result in variable and unknown effects on the recorded call, affecting estimates of call frequency and intensity, particularly for high frequencies. Weather‐induced variation in speed of sound reaches up to about ±3%, but is generally much smaller and only relevant for acoustic localization methods of bats. The frequency‐ and weather‐dependent variation in atmospheric attenuation has a threefold effect on bioacoustic monitoring of bats: It limits our capability (1) to monitor bats equally across time, space, and species, (2) to correctly measure frequency parameters of bat echolocation calls, particularly for high frequencies, and (3) to correctly identify bat species in species‐rich assemblies or for sympatric species with similar call designs.     

BOOK REVIEW

Echolocation calls of four species of insectivorous bats of central Chile were recorded and characterized to determine vocal signatures that allow their identification in the field. Pulses of Tadarida brasiliensis were characterized by the highest duration and the lowest values for all frequencies, which do not overlap those of the remaining species. Tadarida emits narrowband, shallow frequency-modulated search calls. All three vespertilionid species studied (Histiotus montanus, Lasiurus varius and Myotis chiloensis) showed similar echolocation design to one another, consisting of a downward frequency modulation at the beginning of the signal followed by a narrowband quasi-constant frequency component; however, their calls differ by their spectral characteristics. Discriminant function analysis of six acoustic parameters (duration, initial frequency, slope frequency modulation, peak frequency, minimal and maximal frequencies) gave an overall classification of 87.4%, suggesting species could be correctly classified based on echolocation calls. Call duration and minimal frequency were the variables most important for species identification.     

Moth hearing in response to bat echolocation calls manipulated independently in time and frequency

We measured the auditory responses of the noctuid moth Noctua pronuba to bat echolocation calls which were manipulated independently in time and frequency. Such manipulations are important in understanding how insect hearing influences the evolution of echolocation call characteristics. We manipulated the calls of three bat species (Rhinolophus hipposideros, Myotis nattereri and Pipistrellus pipistrellus) that use different echolocation call features by doubling their duration or reducing their frequency, and measured the auditory thresholds from the A1 cells of the moths. Knowing the auditory responses of the moth we tested three predictions. (i) The ranking of the audibility of unmanipulated calls to the moths should be predictable from their temporal and/or frequency structure. This was supported. (ii) Doubling the duration of the calls should increase their audibility by ca. 3 dB for all species. Their audibility did indeed increase by 2.1-3.5 dB. (iii) Reducing the frequency of the calls would increase their audibility for all species. Reducing the frequency had small effects for the two bat species which used short duration (2.7-3.6 ms) calls. However, the relatively long-duration (50 ms), largely constant-frequency calls of R. hipposideros increased in audibility by 21.6 dB when their frequency was halved. Time and frequency changes influence the audibility of calls to tympanate moths in different ways according to call design. Large changes in frequency and time had relatively small changes on the audibility of calls for short, largely broadband calls. Channelling energy into the second harmonic of the call substantially decreased the audibility of calls for bats which use long-duration, constant-frequency components in echolocation calls. We discuss our findings in the contexts of the evolution of both bat echolocation call design and the potential responses of insects which hear ultrasound.     

Acoustic characterization of bats from Malta: setting a baseline for monitoring and conservation of bat populations

Bioacoustic research has made several advancements in developing systems to record extensive acoustic data and classify bat echolocation calls to species level using automated classifiers. These systems are useful as echolocation calls give valuable information on bat behaviour and ecology and hence are widely used for research and conservation of bat populations. Despite the challenges associated with automated classifiers, due to the interspecific differences in call characteristics of bat species found in the Maltese Islands, the use of a quantitative and automated approach is investigated. The sound analysis pipeline involved the use of an algorithm to clean sound files from background noise and measure temporal and spectral parameters of bat echolocation calls. These parameters were then fed to a trained and validated artificial neural network using a bat call library built from reference bat calls from Malta. The automatic classifier achieved an overall correct classification rate of 98%. This high correct classification rate for reliable species identification may have benefitted from the absence of typically problematic species, such as species in the genus Myotis, in the analyses. This study’s results pave the way for efficient and reliable bat acoustic surveys in Malta in aid of necessary monitoring and conservation by providing an updated bat species list and their echolocation characteristics.     

Noctuid moths show neural and behavioural responses to sounds made by some bat-marking rings

Coloured rings are often used for marking bats so that specific individuals can be recognized. We noticed that the rings of mouse-eared bats, Myotis myotis and Myotis blythii, in a combination of one plastic-split and one metallic ring on the same forearm, emitted sounds that were largely ultrasonic each time the rings met in flight. We recorded the ring sounds and the echolocation calls produced by the bats, and played them back to neural preparations of lesser yellow underwing moths, Noctua comes, while making extracellular recordings from the moths' A1 auditory receptors. The peak energy of the ring sounds occurred much closer in frequency to the moth's best auditory frequency (the frequency at which the moth has the lowest auditory threshold) than the peak energy of the calls, for both bat species, and the ring sounds were detected at a threshold 5-6 dB peSPL lower than the calls. Moths performed evasive manoeuvres to playbacks of ring sounds more frequently than they did to control (tape noise) sequences. These neural and behavioural responses imply that certain bats should not be marked with two rings on one wing, as this may make the bat more apparent to tympanate insects, and may therefore reduce its foraging success. Copyright 1999 The Association for the Study of Animal Behaviour.