Bats Actively Use Leaves as Specular Reflectors to Detect Acoustically Camouflaged Prey.

Filtering relevant signals from noisy sensory input is a crucial challenge for animals [1, 2]. Many bats are acoustic specialists relying on sound to find prey. They discern salient acoustic signals from irrelevant background masking noise. It has long been considered a sensory impossibility for bats to use solely echolocation for the detection of silent and motionless prey resting directly on foliage due to the masking effects of background echoes [3, 4]. Some bats, however, do successfully perform this seemingly impossible task [5], raising the question-what underlying acoustic and behavioral mechanisms do bats use to solve this conundrum? To address this question, we used biomimetic sonar to record high-resolution measurements of echoes from insects resting on leaves. Based on our echo recordings, we predicted optimal approach angles from which masking echoes can best be avoided. In behavioral experiments, we put these predictions to test. We recorded the prey approach behavior of wild bats in a flight cage equipped with an ultrasonic microphone synchronized with two high-speed cameras for 3D flightpath reconstructions. Bats approached prey from our predicted optimal oblique angles, using the leaf as a specular reflector to uncover previously acoustically hidden prey. Our findings disclose key behavioral and acoustic mechanisms enabling the detection of prey echoes that background clutter would otherwise mask. This work adds to the fundamental understanding of how bat echolocation strategies can override acoustic camouflage by silent, motionless prey, thus providing new insights into the evolutionary arms race between predators and their prey.

Nocturnal scent in a 'bird-fig': A cue to attract bats as additional dispersers?

The plant genus Ficus is a keystone resource in tropical ecoystems. One of the unique features of figs is the diversity of fruit traits, which in many cases match their various dispersers, the so-called fruit syndromes. The classic example of this is the strong phenotypic differences found between figs with bat and bird dispersers (color, size, presentation, and scent). The 'bird-fig' Ficus colubrinae represents an exception to this trend since it attracts the small frugivorous bat species Ectophylla alba at night, but during the day it attracts bird visitors. Here we investigate day to night changes in fruit scent as a possible mechanism by which this 'bird-fig' could attract bats despite its fruit traits, which should appeal solely to birds. Analyses of odor bouquets from the bat- and bird-dispersal phases (i.e. day and night) differed significantly in their composition of volatiles. We observed a significant increase in relative amounts of sesquiterpene and aromatic compounds at night while relative amounts of two compounds of the fatty acid pathway were significantly higher during day. This finding raises the question whether Ficus colubrinae, a phenotypically classic 'bird-fig', might be able to attract bat dispersers by an olfactory signal at night. Preliminary observations from feeding experiments which indicate that Ectophylla alba is capable of finding ripe figs by scent alone point in this direction. However, additional behavioral experiments on whether bats prefer the 'night-bouquet' over the 'day-bouquet' will be needed to unequivocally answer this question.

Land use history and population dynamics of free-standing figs in a maturing forest.

Figs (Ficus sp.) are often considered as keystone resources which strongly influence tropical forest ecosystems. We used long-term tree-census data to track the population dynamics of two abundant free-standing fig species, Ficus insipida and F. yoponensis, on Barro Colorado Island (BCI), a 15.6-km2 island in Lake Gatún, Panama. Vegetation cover on BCI consists of a mosaic of old growth (>400 years) and maturing (about 90-150 year old) secondary rainforest. Locations and conditions of fig trees have been mapped and monitored on BCI for more than 35 years (1973-2011), with a focus on the Lutz Catchment area (25 ha). The original distribution of the fig trees shortly after the construction of the Panama Canal was derived from an aerial photograph from 1927 and was compared with previous land use and forest status. The distribution of both fig species (~850 trees) is restricted to secondary forest. Of the original 119 trees observed in Lutz Catchment in 1973, >70% of F. insipida and >90% of F. yoponensis had died by 2011. Observations in other areas on BCI support the trend of declining free-standing figs. We interpret the decline of these figs on BCI as a natural process within a maturing tropical lowland forest. Senescence of the fig trees appears to have been accelerated by severe droughts such as the strong El Niño event in the year 1982/83. Because figs form such an important food resource for frugivores, this shift in resource availability is likely to have cascading effects on frugivore populations.

Predictors of elevational biodiversity gradients change from single taxa to the multi-taxa community level.

The factors determining gradients of biodiversity are a fundamental yet unresolved topic in ecology. While diversity gradients have been analysed for numerous single taxa, progress towards general explanatory models has been hampered by limitations in the phylogenetic coverage of past studies. By parallel sampling of 25 major plant and animal taxa along a 3.7 km elevational gradient on Mt. Kilimanjaro, we quantify cross-taxon consensus in diversity gradients and evaluate predictors of diversity from single taxa to a multi-taxa community level. While single taxa show complex distribution patterns and respond to different environmental factors, scaling up diversity to the community level leads to an unambiguous support for temperature as the main predictor of species richness in both plants and animals. Our findings illuminate the influence of taxonomic coverage for models of diversity gradients and point to the importance of temperature for diversification and species coexistence in plant and animal communities.

Finding flowers in the dark: nectar-feeding bats integrate olfaction and echolocation while foraging for nectar.

Nectar-feeding bats depend mainly on floral nectar to fulfil their energetic requirements. Chiropterophilous flowers generally present strong floral scents and provide conspicuous acoustic echoes to attract bats. While floral scents are assumed to attract bats over long distances, acoustic properties of flower structures may provide detailed information, thus supporting the localization of a flower at close ranges. So far, to our knowledge, there is no study trying to understand the relative importance as well as the combination of these generally coupled cues for detection (presence) and localization (exact position) of open flowers in nature. For a better comprehension of the significance of olfaction and echolocation in the foraging behaviour of nectar-feeding bats, we conducted two-choice experiments with Leptonycteris yerbabuenae. We tested the bats' behaviour in three experimental scenarios with different cues: (i) olfaction versus echolocation, (ii) echolocation versus echolocation and olfaction, and (iii) olfaction versus echolocation and olfaction. We used the floral scent of the bat-pollinated cactus Pachycereus pringlei as olfactory cue and an acrylic paraboloid as acoustic cue. Additionally, we recorded the echolocation behaviour of the bats and analysed the floral scent of P. pringlei. When decoupled cues were offered, bats displayed no preference in choice for any of the two cues. However, bats reacted first to and chose more often the coupled cues. All bats echolocated continuously and broadcast a long terminal group before a successful visit. The floral scent bouquet of P. pringlei is composed of 20 compounds, some of which (e.g. methyl benzoate) were already reported from chiropterophilous plants. Our investigation demonstrates for the first time to our knowledge, that nectar-feeding bats integrate over different sensory modes for detection and precise localization of open flowers. The combined information from olfactory and acoustic cues allows bats to forage more efficiently.

How Nectar-Feeding Bats Localize their Food: Echolocation Behavior of Leptonycteris yerbabuenae Approaching Cactus Flowers.

Nectar-feeding bats show morphological, physiological, and behavioral adaptations for feeding on nectar. How they find and localize flowers is still poorly understood. While scent cues alone allow no precise localization of a floral target, the spatial properties of flower echoes are very precise and could play a major role, particularly at close range. The aim of this study is to understand the role of echolocation for classification and localization of flowers. We compared the approach behavior of Leptonycteris yerbabuenae to flowers of a columnar cactus, Pachycereus pringlei, to that to an acrylic hollow hemisphere that is acoustically conspicuous to bats, but has different acoustic properties and, contrary to the cactus flower, present no scent. For recording the flight and echolocation behaviour we used two infrared video cameras under stroboscopic illumination synchronized with ultrasound recordings. During search flights all individuals identified both targets as a possible food source and initiated an approach flight; however, they visited only the cactus flower. In experiments with the acrylic hemisphere bats aborted the approach at ca. 40-50 cm. In the last instant before the flower visit the bats emitted a long terminal group of 10-20 calls. This is the first report of this behaviour for a nectar-feeding bat. Our findings suggest that L. yerbabuenae use echolocation for classification and localization of cactus flowers and that the echo-acoustic characteristics of the flower guide the bats directly to the flower opening.

Sensory challenges for trawling bats: Finding transient prey on water surfaces.

Bats are able to identify obstacles and prey objects based exclusively on acoustic information acquired via echolocation. To assess the echo information potentially available to the trawling bat Noctilio leporinus, prey objects were ensonified with artificial bat calls and deduced echo target strengths (TS) of the reflected signals. The artificial calls consisted either of constant frequency (CF) or frequency modulated (FM) sounds. Detection distances were calculated for call intensities of N. leporinus emitted in the field and in confined space. Measurements of a transient target consisting of a brief water splash and subsequently expanding water ripples revealed that concentrically expanding water ripples can provide sufficiently loud echoes to be detected by trawling bats. Experiments with stationary targets revealed differences in TS depending on the type of signal used (CF or FM). A calculated maximum detection distance between 4.5 and 13.7 m for all measured targets indicates that prey detection in this very loud calling species occurs much earlier than suggested by estimations based on modifications in echolocation or flight behavior.

Spatial Scales of Genetic Structure in Free-Standing and Strangler Figs (Ficus, Moraceae) Inhabiting Neotropical Forests.

Wind-borne pollinating wasps (Agaonidae) can transport fig (Ficus sp., Moraceae) pollen over enormous distances (> 100 km). Because of their extensive breeding areas, Neotropical figs are expected to exhibit weak patterns of genetic structure at local and regional scales. We evaluated genetic structure at the regional to continental scale (Panama, Costa Rica, and Peru) for the free-standing fig species Ficus insipida. Genetic differentiation was detected only at distances > 300 km (Jost´s Dest = 0.68 ± 0.07 & FST = 0.30 ± 0.03 between Mesoamerican and Amazonian sites) and evidence for phylogeographic structure (RST>>permuted RST) was only significant in comparisons between Central and South America. Further, we assessed local scale spatial genetic structure (SGS, d ≤ 8 km) in Panama and developed an agent-based model parameterized with data from F. insipida to estimate minimum pollination distances, which determine the contribution of pollen dispersal on SGS. The local scale data for F. insipida was compared to SGS data collected for an additional free-standing fig, F. yoponensis (subgenus Pharmacosycea), and two species of strangler figs, F. citrifolia and F. obtusifolia (subgenus Urostigma) sampled in Panama. All four species displayed significant SGS (mean Sp = 0.014 ± 0.012). Model simulations indicated that most pollination events likely occur at distances > > 1 km, largely ruling out spatially limited pollen dispersal as the determinant of SGS in F. insipida and, by extension, the other fig species. Our results are consistent with the view that Ficus develops fine-scale SGS primarily as a result of localized seed dispersal and/or clumped seedling establishment despite extensive long-distance pollen dispersal. We discuss several ecological and life history factors that could have species- or subgenus-specific impacts on the genetic structure of Neotropical figs.

Sensory-based niche partitioning in a multiple predator - multiple prey community.

Many predators and parasites eavesdrop on the communication signals of their prey. Eavesdropping is typically studied as dyadic predator-prey species interactions; yet in nature, most predators target multiple prey species and most prey must evade multiple predator species. The impact of predator communities on prey signal evolution is not well understood. Predators could converge in their preferences for conspicuous signal properties, generating competition among predators and natural selection on particular prey signal features. Alternatively, predator species could vary in their preferences for prey signal properties, resulting in sensory-based niche partitioning of prey resources. In the Neotropics, many substrate-gleaning bats use the mate-attraction songs of male katydids to locate them as prey. We studied mechanisms of niche partitioning in four substrate-gleaning bat species and found they are similar in morphology, echolocation signal design and prey-handling ability, but each species preferred different acoustic features of male song in 12 sympatric katydid species. This divergence in predator preference probably contributes to the coexistence of many substrate-gleaning bat species in the Neotropics, and the substantial diversity in the mate-attraction signals of katydids. Our results provide insight into how multiple eavesdropping predator species might influence prey signal evolution through sensory-based niche partitioning.

Cheating on the mutualistic contract: nutritional gain through seed predation in the frugivorous bat Chiroderma villosum (Phyllostomidae).

Most frugivorous bats are efficient seed dispersers, as they typically do not damage seeds and transport them over long distances. In contrast, bats of the phyllostomid genus Chiroderma cheat fig trees by acting more as seed predators than as seed dispersers. The bats initially separate seeds from fruit pulp in the mouth. After extracting the juice from the fruit pulp, they thoroughly chew the seeds and spit out small seed fragments in a pellet. Consequently, the faeces contain almost no viable seeds. We compared the nutrient content of intact fig seeds with ejecta and faecal samples from both Chiroderma villosum and the 'conventional' frugivorous bat Artibeus watsoni. We show that C. villosum can extract nutrients from the seeds, especially protein and fat. The processing time of figs showed no significant difference between the two bat species. Food-choice experiments showed that C. villosum preferred fig species with more seeds over those with fewer seeds. This preference, in combination with the specialized seed-chewing behaviour, leads to an increased nutrient intake per fig. This unique strategy enables C. villosum to satisfy its nutritional requirements with a lower number of figs than other species, which decreases the amount of energy necessary for foraging flights as well as the predation risk during foraging.

Frugivorous bats maintain functional habitat connectivity in agricultural landscapes but rely strongly on natural forest fragments.

Anthropogenic changes in land use threaten biodiversity and ecosystem functioning by the conversion of natural habitat into agricultural mosaic landscapes, often with drastic consequences for the associated fauna. The first step in the development of efficient conservation plans is to understand movement of animals through complex habitat mosaics. Therefore, we studied ranging behavior and habitat use in Dermanura watsoni (Phyllostomidae), a frugivorous bat species that is a valuable seed disperser in degraded ecosystems. Radio-tracking of sixteen bats showed that the animals strongly rely on natural forest. Day roosts were exclusively located within mature forest fragments. Selection ratios showed that the bats foraged selectively within the available habitat and positively selected natural forest. However, larger daily ranges were associated with higher use of degraded habitats. Home range geometry and composition of focal foraging areas indicated that wider ranging bats performed directional foraging bouts from natural to degraded forest sites traversing the matrix over distances of up to three hundred meters. This behavior demonstrates the potential of frugivorous bats to functionally connect fragmented areas by providing ecosystem services between natural and degraded sites, and highlights the need for conservation of natural habitat patches within agricultural landscapes that meet the roosting requirements of bats.

Species undersampling in tropical bat surveys: effects on emerging biodiversity patterns.

Undersampling is commonplace in biodiversity surveys of species-rich tropical assemblages in which rare taxa abound, with possible repercussions for our ability to implement surveys and monitoring programmes in a cost-effective way. We investigated the consequences of information loss due to species undersampling (missing subsets of species from the full species pool) in tropical bat surveys for the emerging patterns of species richness (SR) and compositional variation across sites. For 27 bat assemblage data sets from across the tropics, we used correlations between original data sets and subsets with different numbers of species deleted either at random, or according to their rarity in the assemblage, to assess to what extent patterns in SR and composition in data subsets are congruent with those in the initial data set. We then examined to what degree high sample representativeness (r ≥ 0·8) was influenced by biogeographic region, sampling method, sampling effort or structural assemblage characteristics. For SR, correlations between random subsets and original data sets were strong (r ≥ 0·8) with moderate (ca. 20%) species loss. Bias associated with information loss was greater for species composition; on average ca. 90% of species in random subsets had to be retained to adequately capture among-site variation. For nonrandom subsets, removing only the rarest species (on average c. 10% of the full data set) yielded strong correlations (r > 0·95) for both SR and composition. Eliminating greater proportions of rare species resulted in weaker correlations and large variation in the magnitude of observed correlations among data sets. Species subsets that comprised ca. 85% of the original set can be considered reliable surrogates, capable of adequately revealing patterns of SR and temporal or spatial turnover in many tropical bat assemblages. Our analyses thus demonstrate the potential as well as limitations for reducing survey effort and streamlining sampling protocols, and consequently for increasing the cost-effectiveness in tropical bat surveys or monitoring programmes. The dependence of the performance of species subsets on structural assemblage characteristics (total assemblage abundance, proportion of rare species), however, underscores the importance of adaptive monitoring schemes and of establishing surrogate performance on a site by site basis based on pilot surveys.

Temporal changes in randomness of bird communities across Central Europe.

Many studies have examined whether communities are structured by random or deterministic processes, and both are likely to play a role, but relatively few studies have attempted to quantify the degree of randomness in species composition. We quantified, for the first time, the degree of randomness in forest bird communities based on an analysis of spatial autocorrelation in three regions of Germany. The compositional dissimilarity between pairs of forest patches was regressed against the distance between them. We then calculated the y-intercept of the curve, i.e. the 'nugget', which represents the compositional dissimilarity at zero spatial distance. We therefore assume, following similar work on plant communities, that this represents the degree of randomness in species composition. We then analysed how the degree of randomness in community composition varied over time and with forest management intensity, which we expected to reduce the importance of random processes by increasing the strength of environmental drivers. We found that a high portion of the bird community composition could be explained by chance (overall mean of 0.63), implying that most of the variation in local bird community composition is driven by stochastic processes. Forest management intensity did not consistently affect the mean degree of randomness in community composition, perhaps because the bird communities were relatively insensitive to management intensity. We found a high temporal variation in the degree of randomness, which may indicate temporal variation in assembly processes and in the importance of key environmental drivers. We conclude that the degree of randomness in community composition should be considered in bird community studies, and the high values we find may indicate that bird community composition is relatively hard to predict at the regional scale.

High local diversity of Trypanosoma in a common bat species, and implications for the biogeography and taxonomy of the T. cruzi clade.

The Trypanosoma cruzi clade is a group of parasites that comprises T. cruzi sensu lato and its closest relatives. Although several species have been confirmed phylogenetically to belong to this clade, it is uncertain how many more species can be expected to belong into this group. Here, we present the results of a survey of trypanosome parasites of the bat Artibeus jamaicensis from the Panamá Canal Zone, an important seed disperser. Using a genealogical species delimitation approach, the Poisson tree processes (PTP), we tentatively identified five species of trypanosomes - all belonging to the T. cruzi clade. A small monophyletic group of three putative Trypanosoma species places at the base of the clade phylogeny, providing evidence for at least five independent colonization events of these parasites into the New World. Artibeus jamaicensis presents a high diversity of these blood parasites and is the vertebrate with the highest number of putative trypanosome species reported from a single locality. Our results emphasize the need for continued efforts to survey mammalian trypanosomes.

Movement and ranging patterns of the Common Chaffinch in heterogeneous forest landscapes.

The partitioning of production forests into discretely managed forest stands confronts animals with diversity in forest attributes at scales from point-level tree assemblages to distinct forest patches and range-level forest cover. We have investigated the movement and ranging patterns of male Common Chaffinches, Fringilla coelebs, in heterogeneous forest production landscapes during spring and summer in south-western Germany. We radio-tracked a total of 15 adult males, each for up to six days, recording locations at 10-min intervals. We then performed point-level tree surveys at all tracking locations and classified forest stand attributes for the areal covering of birds' ranges. Movement distances were shortest in beech forest stands and longer in spruce-mixed and non-spruce conifer stands. Movement distances increased with stand age in beech stands but not in others, an effect that was only detectable in a multilevel hierarchical model. We found negligible effects of point-level tree assemblages and temperature on movement distances. Daily range estimates were from 0.01 to 8.0 hectare (median of 0.86 ha) with no evident impact of forest attributes on ranging patterns but considerable intra-individual variation in range sizes over consecutive days. Most daily ranges covered more than one forest stand type. Our results show that forest management impacts the movement behaviour of chaffinches in heterogeneous production forest. Although point-level effects of movement distances are weak compared with stand-level effects in this study, the hierarchical organization of forest is an important aspect to consider when analysing fine-scale movement and might exert more differentiated effects on bird species that are more sensitive to habitat changes than the chaffinch.

Biosonar resolving power: echo-acoustic perception of surface structures in the submillimeter range.

The minimum distance for which two points still can be separated from each other defines the resolving power of a visual system. In an echo-acoustic context, the resolving power is usually measured as the smallest perceivable distance of two reflecting surfaces on the range axis and is found to be around half a millimeter for bats employing frequency modulated (FM) echolocation calls. Only few studies measured such thresholds with physical objects, most often bats were trained on virtual echoes i.e., echoes generated and played back by a computer; moreover, bats were sitting while they received the stimuli. In these studies differences in structure depth between 200 and 340 µm were found. However, these low thresholds were never verified for free-flying bats and real physical objects. Here, we show behavioral evidence that the echo-acoustic resolving power for surface structures in fact can be as low as measured for computer generated echoes and even lower, sometimes below 100 µm. We found this exceptional fine discrimination ability only when one of the targets showed spectral interferences in the frequency range of the bats' echolocation call while the other target did not. This result indicates that surface structure is likely to be perceived as a spectral quality rather than being perceived strictly in the time domain. Further, it points out that sonar resolving power directly depends on the highest frequency/shortest wavelength of the signal employed.

Complementary ecosystem services provided by pest predators and pollinators increase quantity and quality of coffee yields.

Wild animals substantially support crop production by providing ecosystem services, such as pollination and natural pest control. However, the strengths of synergies between ecosystem services and their dependencies on land-use management are largely unknown. Here, we took an experimental approach to test the impact of land-use intensification on both individual and combined pollination and pest control services in coffee production systems at Mount Kilimanjaro. We established a full-factorial pollinator and vertebrate exclosure experiment along a land-use gradient from traditional homegardens (agroforestry systems), shaded coffee plantations to sun coffee plantations (total sample size = 180 coffee bushes). The exclusion of vertebrates led to a reduction in fruit set of ca 9%. Pollinators did not affect fruit set, but significantly increased fruit weight of coffee by an average of 7.4%. We found no significant decline of these ecosystem services along the land-use gradient. Pest control and pollination service were thus complementary, contributing to coffee production by affecting the quantity and quality of a major tropical cash crop across different coffee production systems at Mount Kilimanjaro.

Driving factors for the evolution of species-specific echolocation call design in new world free-tailed bats (molossidae).

Phylogeny, ecology, and sensorial constraints are thought to be the most important factors influencing echolocation call design in bats. The Molossidae is a diverse bat family with a majority of species restricted to tropical and subtropical regions. Most molossids are specialized to forage for insects in open space, and thus share similar navigational challenges. We use an unprecedented dataset on the echolocation calls of 8 genera and 18 species of New World molossids to explore how habitat, phylogenetic relatedness, body mass, and prey perception contribute to echolocation call design. Our results confirm that, with the exception of the genus Molossops, echolocation calls of these bats show a typical design for open space foraging. Two lines of evidence point to echolocation call structure of molossids reflecting phylogenetic relatedness. First, such structure is significantly more similar within than among genera. Second, except for allometric scaling, such structure is nearly the same in congeneric species. Despite contrasting body masses, 12 of 18 species call within a relatively narrow frequency range of 20 to 35 kHz, a finding that we explain by using a modeling approach whose results suggest this frequency range to be an adaptation optimizing prey perception in open space. To conclude, we argue that the high variability in echolocation call design of molossids is an advanced evolutionary trait allowing the flexible adjustment of echolocation systems to various sensorial challenges, while conserving sender identity for social communication. Unraveling evolutionary drivers for echolocation call design in bats has so far been hampered by the lack of adequate model organisms sharing a phylogenetic origin and facing similar sensorial challenges. We thus believe that knowledge of the echolocation call diversity of New World molossid bats may prove to be landmark to understand the evolution and functionality of species-specific signal design in bats.

The third dimension of bat migration: evidence for elevational movements of Miniopterus natalensis along the slopes of Mount Kilimanjaro.

Bats are important ecosystem service providers, and therefore most relevant for both lowland and highland habitats, particularly in the tropics. Yet, it is poorly understood to what extent they perform large-scale movements, especially movements along mountain slopes. Here, we studied the movement ecology of the potentially migratory species Miniopterus natalensis at Mount Kilimanjaro in Tanzania. We analysed stable isotope ratios of C (δ(13)C), N (δ(15)N) and H (δ(2)H) in keratin of sedentary frugivorous and insectivorous bats captured between 800 and 2,400 m above sea level to establish elevational gradients of stable isotope ratios in consumer tissues. We expected correlations between stable isotope ratios of the non-exchangeable portion of H in fur keratin and the elevation of capture site, but not necessarily for δ(13)C and δ(15)N. Yet, in bats of both feeding ensembles, we found δ(15)N of fur keratin to correlate positively with the elevation of capture sites but not δ(2)H. In frugivorous bats, δ(13)C increased with increasing capture elevation as well. By looking at intra-individual variation of δ(13)C and δ(15)N in fur keratin and wing membrane tissues of sedentary Rhinolophus cf. clivosus and of the potentially migratory species M. natalensis, we gathered evidence that M. natalensis migrates seasonally between low and high elevations along the slopes of Mount Kilimanjaro. Finally, based on an isoscape origin model we estimated that M. natalensis captured before and after the cold period at around 1,800 m above sea level originated from around 1,400 m a.s.l. or lower. Thus, we received convergent results in support of seasonal elevational movements of M. natalensis, probably in search for cold hibernacula at higher elevations of Mount Kilimanjaro.

Behavioral flexibility of the trawling long-legged bat, Macrophyllum macrophyllum (Phyllostomidae).

We assessed the behavioral flexibility of the trawling long-legged bat, Macrophyllum macrophyllum (Phyllostomidae) in flight cage experiments by exposing it to prey suspended from nylon threads in the air and to food placed onto the water surface at varying distances to clutter-producing background (water plants). The bat revealed flexibility in foraging mode and caught prey in the air (aerial hawking) and from the water surface (trawling). M. macrophyllum was constrained in finding food very near to and within clutter. As echolocation was the prime sensory mode used by M. macrophyllum for detection and localization of food, the bat might have been unable to perceive sufficient information from prey near clutter as background echoes from the water plant increasingly overlapped with echoes from food. The importance of echolocation for foraging is reflected in a stereotypic call pattern of M. macrophyllum that resembles other aerial insectivorous and trawling bats with a pronounced terminal phase (buzz) prior to capture attempts. Our findings contrast studies of other phyllostomid bats that glean prey very near or from vegetation, often using additional sensory cues, such as prey-produced noise, to find food and that lack a terminal phase in echolocation behavior. In M. macrophyllum, acoustic characteristics of its foraging habitat have shaped its sonar system more than phylogeny.