Bats as prey of diurnal birds: a global perspective

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Activity patterns of insectivorous bats and birds in northern Scandinavia (69° N), during continuous midsummer daylight

Previous studies suggest that many species of insectivorous bats are nocturnal, despite the relatively low availability of their insect prey at night, because of the risk of predation by diurnal predatory birds. We hypothesised that if this was the case bats living above the arctic circle would alter their feeding behaviour during midsummer because there would no longer be any benefit to restricting their activity to the period when their prey are least abundant. Alternatively, if bats were more influenced by competition from aerial insectivorous birds they would continue to feed at 'night' to avoid such competition. In northern Norway (69°  N), during continuous midsummer daylight, insectivorous sand martins ( Riparia riparia ) concentrated their aerial feeding activity when aerial insects were most abundant. The birds stopped feeding between 23:00 and 07:00 when aerial insects were least abundant. In contrast, northern bats ( Eptesicus nilssonii ), fed mostly between 22:00 and 02:00, coinciding with the lowest aerial insect availability, and with the period when light levels were lowest (ca 1000 lux). Bat activity patterns were closest to those predicted by the avian competition hypothesis. The low densities of both sand martins and Northern bats in the study area, however, were less consistent with this hypothesis. Possibly populations of both species were higher historically and the observed patterns reflected historical competition. Bat activity was most closely correlated to ambient light levels. This raised two alternative explanations that we could not eliminate. Perhaps there was differential predation risk, between the brightest and darkest parts of the day, because the visual capacities of falcons are strongly dependent on luminance. Alternatively the bats may have been entrained to emerge at given light levels by their behaviour at other times of year.     

Bats are Not Birds – Different Responses to Human Land‐use on a Tropical Mountain

Land‐use intensification has consequences for biodiversity and ecosystem functioning, with various taxonomic groups differing widely in their sensitivity. As land‐use intensification alters habitat structure and resource availability, both factors may contribute to explaining differences in animal species diversity. Within the local animal assemblages the flying vertebrates, bats and birds, provide important and partly complementary ecosystem functions. We tested how bats and birds respond to land‐use intensification and compared abundance, species richness, and community composition across a land‐use gradient including forest, traditional agroforests (home garden), coffee plantations and grasslands on Mount Kilimanjaro, Tanzania. Furthermore, we asked how sensitive different habitat and feeding guilds of bats and birds react to land‐use intensification and the associated alterations in vegetation structure and food resource availability. In contrast to our expectations, land‐use intensification had no negative effect on species richness and abundance of all birds and bats. However, some habitat and feeding guilds, in particular forest specialist and frugivorous birds, were highly sensitive to land‐use intensification. Although the habitat guilds of both, birds and bats, depended on a certain degree of vegetation structure, total bat and bird abundance was mediated primarily by the availability of the respective food resources. Even though the highly structured southern slopes of Mount Kilimanjaro are able to maintain diverse bat and bird assemblages, the sensitivity of avian forest specialists against land‐use intensification and the dependence of the bat and bird habitat guilds on a certain vegetation structure demonstrate that conservation plans should place special emphasis on these guilds.     

Effects of Wind Turbine Curtailment on Bird and Bat Fatalities

Bird and bat fatalities increase with wind energy expansion and the only effective fatality-reduction measure has been operational curtailment, which has been documented for bats but not for birds. We performed opportune before-after, control-impact (BACI) experiments of curtailment effects on bird and bat fatalities and nocturnal passage rates during fall migration at 2 wind projects, where 1 continued operating and the other shut down from peak migration to the study's end (study 1). We also performed BACI experiments during a 3-year study of curtailment and operational effects on bird fatalities among wind turbines of varying operational status (study 2). In study 1, wind turbine curtailment significantly reduced near-misses and rotor-disrupted flights of bats, and it significantly reduced fatalities of bats but not of birds. In study 2, converting wind turbines from inoperable to operable status did not significantly increase bird fatalities, and bird species of hole or sheltered-ledge nesters or roosters on human-made structures died in substantial numbers at vacant towers. Of bird species represented by fatalities in study 2, 79% were found at inoperable wind turbines. Because the migration season is relatively brief, seasonal curtailment would greatly reduce bat fatalities for a slight loss in annual energy generation, but it might not benefit many bird species. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

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.     

Landscape structure influences the use of social information in an insectivorous bat

In anthropogenic landscapes, aerial insectivores are often confronted with variable habitat complexity, which may influence the distribution of prey. Yet, high mobility may allow aerial insectivores to adjust their foraging strategy to different prey distributions. We investigated whether aerial-hunting common noctules Nyctalus noctula adjust their foraging strategy to landscapes with different habitat complexity and assumingly different prey distribution. We hypothesized that the movement behaviour of hunting common noctules and changes of movement behaviour in reaction towards conspecifics would depend on whether they hunt in a structurally poor cropland dominated landscape or a structurally rich forest dominated landscape. We tracked flight paths of common noctules in northeastern Germany using GPS loggers equipped with an ultrasonic microphone that recorded foraging events and presence of conspecifics. Above cropland, common noctules hunted mainly during bouts of highly tortuous and area restricted movements (ARM). Bats switched from straight flight to ARM after encountering conspecifics. In the forested landscape, common noctules hunted both during ARM and during straight flights. The onset of ARM did not correlate with the presence of conspecifics. Common noctules showed a lower feeding rate and encountered more conspecifics above the forested than above the cropland dominated landscape. We conjecture that prey distribution above cropland was patchy and unpredictable, thus making eavesdropping on hunting conspecifics crucial for bats during search for prey patches. In contrast, small scale structural diversity of the forested landscape possibly led to a more homogeneous prey distribution at the landscape scale, thus enabling bats to find sufficient food independent of conspecific presence. This suggests that predators depending on ephemeral prey can increase their foraging success in structurally poor landscapes by using social information provided by conspecifics. Hence, a minimum population density might be obligatory to enable successful foraging in simplified landscapes.     

Evolution of reversed sexual size dimorphism and role partitioning among predatory birds, with a size scaling of flight performance

To explain the adaptive significance of sex role partitioning and reversed sexual size dimorphism among raptors, owls and skuas, where females are usually larger than males, we combine several previous hypotheses with some new ideas. Owing to their structural and behavioural adaptations for prey capture, predatory birds have better prospects than other birds of defending their offspring against nest predators. This makes sex role partitioning advantageous; one parent guards the offspring while the other forages for the family. Further, among predators hunting alert prey such as vertebrates, two mates because of interference may not procur much more food than would one mate hunting alone. By contrast, two mates feeding on less alert prey may together obtain almost twice as much food as one mate hunting alone. For these reasons, partitioning of breeding labours might be adaptive only in predatory birds. An initial imbalance favours female nest guarding and male foraging: the developing eggs might be damaged if the female attacks prey; their mass might reduce her flight performance; she must visit the nest to lay; and the male feeds her before she lays (‘courtship feeding’). Increased female body size should enhance egg production, incubation, ability to tear apart prey for the young, and, in particular, offspring protection in predatory birds. Efficient foraging during the breeding period then becomes most important for the male. This imposes great demands on aerial agility in males, particularly among predators of agile prey. Flight performance decreases with increasing size in five of six aspects explored. The male must therefore not be too large in relation to the most important prey. For these reasons, he should be smaller than the female. Among predatory birds, size dimorphism increases with the proportion of birds in the diet, which may be explained as follows. Adult birds have mainly one type of predators: other predatory birds. Because almost only these specialists exploit adult birds, they carry out most of the cropping of this prey. A predator of easier prey competes with many other kinds of predators, which considerably reduce prey abundance in its territory. This is not so for predators of adult birds. Further, because birds are extremely agile, the specialized predator can hunt efficiently only within a limited size range of birds, whose flight skill it can match. Increased size dimorphism among these predators therefore should be particularly important for enlarging the combined food base of the pair. A bird specialist may consume much of the available prey in the suitable size range during the breeding period. When the predator's young are large enough to defend themselves, the female aids better by hunting than by guarding the chicks. It is advantageous among bird specialists if she hunts prey of other sizes than does the male, who has by then reduced prey abundance in his prey size class. But among predatory birds hunting easier prey the female gains little by hunting outside the male's prey spectrum, because other kinds of predators will have reduced the prey abundance outside as well as inside the male's preferred size range. Intra-pair food separation through large sexual size dimorphism therefore should be particularly advantageous among predators of birds. This may be the main reason why the degree of size dimorphism increases with the dietary proportion of birds.     

What you need is what you eat? Prey selection by the bat Myotis daubentonii

Optimal foraging theory predicts that predators are selective when faced with abundant prey, but become less picky when prey gets sparse. Insectivorous bats in temperate regions are faced with the challenge of building up fat reserves vital for hibernation during a period of decreasing arthropod abundances. According to optimal foraging theory, prehibernating bats should adopt a less selective feeding behaviour – yet empirical studies have revealed many apparently generalized species to be composed of specialist individuals. Targeting the diet of the bat Myotis daubentonii, we used a combination of molecular techniques to test for seasonal changes in prey selectivity and individual‐level variation in prey preferences. DNA metabarcoding was used to characterize both the prey contents of bat droppings and the insect community available as prey. To test for dietary differences among M. daubentonii individuals, we used ten microsatellite loci to assign droppings to individual bats. The comparison between consumed and available prey revealed a preference for certain prey items regardless of availability. Nonbiting midges (Chironomidae) remained the most highly consumed prey at all times, despite a significant increase in the availability of black flies (Simuliidae) towards the end of the season. The bats sampled showed no evidence of individual specialization in dietary preferences. Overall, our approach offers little support for optimal foraging theory. Thus, it shows how novel combinations of genetic markers can be used to test general theory, targeting patterns at both the level of prey communities and individual predators.     

Comparing aerodynamic efficiency in birds and bats suggests better flight performance in birds

Flight is one of the energetically most costly activities in the animal kingdom, suggesting that natural selection should work to optimize flight performance. The similar size and flight speed of birds and bats may therefore suggest convergent aerodynamic performance; alternatively, flight performance could be restricted by phylogenetic constraints. We test which of these scenarios fit to two measures of aerodynamic flight efficiency in two passerine bird species and two New World leaf-nosed bat species. Using time-resolved particle image velocimetry measurements of the wake of the animals flying in a wind tunnel, we derived the span efficiency, a metric for the efficiency of generating lift, and the lift-to-drag ratio, a metric for mechanical energetic flight efficiency. We show that the birds significantly outperform the bats in both metrics, which we ascribe to variation in aerodynamic function of body and wing upstroke: Bird bodies generated relatively more lift than bat bodies, resulting in a more uniform spanwise lift distribution and higher span efficiency. A likely explanation would be that the bat ears and nose leaf, associated with echolocation, disturb the flow over the body. During the upstroke, the birds retract their wings to make them aerodynamically inactive, while the membranous bat wings generate thrust and negative lift. Despite the differences in performance, the wake morphology of both birds and bats resemble the optimal wake for their respective lift-to-drag ratio regimes. This suggests that evolution has optimized performance relative to the respective conditions of birds and bats, but that maximum performance is possibly limited by phylogenetic constraints. Although ecological differences between birds and bats are subjected to many conspiring variables, the different aerodynamic flight efficiency for the bird and bat species studied here may help explain why birds typically fly faster, migrate more frequently and migrate longer distances than bats.     

Predation on bats by Great Kiskadees

ABSTRACT.   Great Kiskadees ( Pitangus sulphuratus ) are found in a variety of habitats from Argentina north to the United States and are the most generalist tyrant flycatcher in both foraging behavior and food habits. These kiskadees are known to occasionally prey on small vertebrates, but, to our knowledge, bats have never been reported as a prey item. We observed a breeding pair of Great Kiskadees preying on bats ( Myotis spp.) at the field station Base de Estudos do Pantanal (BEP) in the southern Pantanal, Brazil. At BEP, there are fissures under the building's floor slabs that allow two species of bats, black myotis ( Myotis nigricans ) and silver-tipped myotis ( M. albescens ), to access internal galleries and use them as day roosts. We found that bats, insects, and fruits were the most common food items fed to nestlings by adult kiskadees. Bats ( N = 10) were captured when kiskadees landed on ground below a building, looked up through a fissure, and then reached through the fissure and captured a bat in their bill. On one occasion, a kiskadee flew from a perch and captured a bat in flight. Our observations provide further evidence of the opportunistic feeding behavior of Great Kiskadees.     

Light enough to travel: migratory bats have smaller brains,but not larger hippocampi,than sedentary species

Migratory bird species have smaller brains than non-migratory species. The behavioural flexibility/migratory precursor hypothesis suggests that sedentary birds have larger brains to allow the behavioural flexibility required in a seasonally variable habitat. The energy trade-off hypothesis proposes that brains are heavy, energetically expensive and therefore, incompatible with migration. Here, we compared relative brain, neocortex and hippocampus volume between migratory and sedentary bats at the species-level and using phylogenetically independent contrasts. We found that migratory bats had relatively smaller brains and neocortices than sedentary species. Our results support the energy trade-off hypothesis because bats do not exhibit the same degree of flexibility in diet selection as sedentary birds. Our results also suggest that bat brain size differences are subtler than those found in birds, perhaps owing to bats'' shorter migration distances. Conversely, we found no difference in relative hippocampus volume between migratory and sedentary species, underscoring our limited understanding of the role of the hippocampus in bats.     

Do the evolutionary interactions between moths and bats promote niche partitioning between bats and birds?

Ecological theory suggests that the coexistence of species is promoted by the partitioning of available resources, as in dietary niche partitioning where predators partition prey. Yet, the mechanisms underlying dietary niche partitioning are not always clear. We used fecal DNA metabarcoding to investigate the diets of seven nocturnal insectivorous bird and bat species. Low diet overlap (2%–22%) supported resource partitioning among all species. Differences in diet corresponded with species identity, prey detection method, and foraging behavior of predators. Insects with ultrasonic hearing capabilities were consumed significantly more often by birds than bats, consistent with an evolved avoidance of echolocating strategies. In turn, bats consumed a greater proportion of noneared insects such as spruce budworms. Overall, our results suggest that evolutionary interactions among bats and moths translate to dietary niche partitioning and coexistence among bats and nocturnal birds.     

Echolocating bats cry out loud to detect their prey

Echolocating bats have successfully exploited a broad range of habitats and prey. Much research has demonstrated how time-frequency structure of echolocation calls of different species is adapted to acoustic constraints of habitats and foraging behaviors. However, the intensity of bat calls has been largely neglected although intensity is a key factor determining echolocation range and interactions with other bats and prey. Differences in detection range, in turn, are thought to constitute a mechanism promoting resource partitioning among bats, which might be particularly important for the species-rich bat assemblages in the tropics. Here we present data on emitted intensities for 11 species from 5 families of insectivorous bats from Panamá hunting in open or background cluttered space or over water. We recorded all bats in their natural habitat in the field using a multi-microphone array coupled with photographic methods to assess the bats' position in space to estimate emitted call intensities. All species emitted intense search signals. Output intensity was reduced when closing in on background by 4-7 dB per halving of distance. Source levels of open space and edge space foragers (Emballonuridae, Mormoopidae, Molossidae, and Vespertilionidae) ranged between 122-134 dB SPL. The two Noctilionidae species hunting over water emitted the loudest signals recorded so far for any bat with average source levels of ca. 137 dB SPL and maximum levels above 140 dB SPL. In spite of this ten-fold variation in emitted intensity, estimates indicated, surprisingly, that detection distances for prey varied far less; bats emitting the highest intensities also emitted the highest frequencies, which are severely attenuated in air. Thus, our results suggest that bats within a local assemblage compensate for frequency dependent attenuation by adjusting the emitted intensity to achieve comparable detection distances for prey across species. We conclude that for bats with similar hunting habits, prey detection range represents a unifying constraint on the emitted intensity largely independent of call shape, body size, and close phylogenetic relationships.     

Nectar-feeding bird and bat niches in two worlds: pantropical comparisons of vertebrate pollination systems

Aim We review several aspects of the structure of regional and local assemblages of nectar‐feeding birds and bats and their relationships with food plants to determine the extent to which evolutionary convergence has or has not occurred in the New and Old World tropics. Location Our review is pantropical in extent and also includes the subtropics of South Africa and eastern Australia. Within the tropics, it deals mostly with lowland forest habitats. Methods An extensive literature review was conducted to compile data bases on the regional and local species richness of nectar‐feeding birds and bats, pollinator sizes, morphology, and diets. Coefficients of variation (CVs) were used to quantify the morphospace occupied by the various families of pollinators. The extent to which plants have become evolutionarily specialized for vertebrate pollination was explored using several criteria: number and diversity of growth forms of plant families providing food for all the considered pollinator families; the most common flower morphologies visited by all the considered pollinator families; and the number of plant families that contain genera with both bird‐ and bat‐specialized species. Results Vertebrate pollinator assemblages in the New World tropics differ from those in the Old World in terms of their greater species richness, the greater morphological diversity of their most specialized taxa, and the greater degree of taxonomic and ecological diversity and morphological specialization of their food plants. Within the Old World tropics, Africa contains more specialized nectar‐feeding birds than Asia and Australasia; Old World nectar‐feeding bats are everywhere less specialized than their New World counterparts. Main conclusions We propose that two factors – phylogenetic history and spatio‐temporal predictability (STP) of flower resources – largely account for hemispheric and regional differences in the structure of vertebrate pollinator assemblages. Greater resource diversity and resource STP in the New World have favoured the radiation of small, hovering nectar‐feeding birds and bats into a variety of relatively specialized feeding niches. In contrast, reduced resource diversity and STP in aseasonal parts of Asia as well as in Australasia have favoured the evolution of larger, non‐hovering birds and bats with relatively generalized feeding niches. Tropical Africa more closely resembles the Neotropics than Southeast Asia and Australasia in terms of resource STP and in the niche structure of its nectar‐feeding birds but not its flower‐visiting bats.     

Dietary overlap and seasonality in three species of mormoopid bats from a tropical dry forest

Competing hypotheses explaining species’ use of resources have been advanced. Resource limitations in habitat and/or food are factors that affect assemblages of species. These limitations could drive the evolution of morphological and/or behavioural specialization, permitting the coexistence of closely related species through resource partitioning and niche differentiation. Alternatively, when resources are unlimited, fluctuations in resources availability will cause concomitant shifts in resource use regardless of species identity. Here, we used next‐generation sequencing to test these hypotheses and characterize the diversity, overlap and seasonal variation in the diet of three species of insectivorous bats of the genus Pteronotus. We identified 465 prey (MOTUs) in the guano of 192 individuals. Lepidoptera and Diptera represented the most consumed insect orders. Diet of bats exhibited a moderate level of overlap, with the highest value between Pteronotus parnellii and Pteronotus personatus in the wet season. We found higher dietary overlap between species during the same seasons than within any single species across seasons. This suggests that diets of the three species are driven more by prey availability than by any particular predator‐specific characteristic. P. davyi and P. personatus increased their dietary breadth during the dry season, whereas P. parnellii diet was broader and had the highest effective number of prey species in all seasons. This supports the existence of dietary flexibility in generalist bats and dietary niche overlapping among groups of closely related species in highly seasonal ecosystems. Moreover, the abundance and availability of insect prey may drive the diet of insectivores.     

Bird and bat predation services in tropical forests and agroforestry landscapes

Understanding distribution patterns and multitrophic interactions is critical for managing bat‐ and bird‐mediated ecosystem services such as the suppression of pest and non‐pest arthropods. Despite the ecological and economic importance of bats and birds in tropical forests, agroforestry systems, and agricultural systems mixed with natural forest, a systematic review of their impact is still missing. A growing number of bird and bat exclosure experiments has improved our knowledge allowing new conclusions regarding their roles in food webs and associated ecosystem services. Here, we review the distribution patterns of insectivorous birds and bats, their local and landscape drivers, and their effects on trophic cascades in tropical ecosystems. We report that for birds but not bats community composition and relative importance of functional groups changes conspicuously from forests to habitats including both agricultural areas and forests, here termed ‘forest‐agri’ habitats, with reduced representation of insectivores in the latter. In contrast to previous theory regarding trophic cascade strength, we find that birds and bats reduce the density and biomass of arthropods in the tropics with effect sizes similar to those in temperate and boreal communities. The relative importance of birds versus bats in regulating pest abundances varies with season, geography and management. Birds and bats may even suppress tropical arthropod outbreaks, although positive effects on plant growth are not always reported. As both bats and birds are major agents of pest suppression, a better understanding of the local and landscape factors driving the variability of their impact is needed.     

Selective Eavesdropping Behaviour in Three Neotropical Bat Species

Knowledge of interspecies information transfer in mammals is scarce compared with other taxa. We investigated whether eavesdropping on echolocation calls of bats may be used by sympatric bats with similar feeding ecology. We performed playback experiments with three free‐ranging neotropical bat species, broadcasting search phase calls or feeding buzzes of conspecifics and heterospecifics belonging either to the same or to another bat family. Both the greater fishing bat Noctilio leporinus and the lesser bulldog bat Noctilio albiventris (Noctilionidae) reacted with repeated approaches in response to playbacks of search phase calls and feeding buzzes from conspecifics and also to congeneric feeding buzzes. Noctilio leporinus also were attracted by search phase calls from its sister species N. albiventris. In contrast, the sac‐winged bat Saccopteryx bilineata (Emballonuridae) did not react to any playback sequences presented. Our results support the existence of eavesdropping behaviour for both species of Noctilio. We suggest that information transfer via eavesdropping may depend mainly on species‐specific traits, including foraging style and social behaviour (territoriality, group foraging), and on distribution and density of prey. Call design had only a minor influence on the reaction.     

Seasonal patterns of habitat selection in the insectivorous bat‐eared fox

Understanding how animals utilize their habitat provides insights about their ecological needs and is of importance for both theoretical and applied ecology. As changing seasons impact prey habitat selection and vegetation itself, it is important to understand how seasonality impacts microhabitat choice in optimal foragers and their prey. We followed habituated bat‐eared foxes (Otocyon megalotis) in the Kalahari, South Africa, to study their seasonal habitat selection patterns and relate them to the habitat preferences of their main prey, termites (Hodotermes mossambicus). We used Resource Selection Functions (RSFs) to study bat‐eared foxes’ 3rd‐ and 4th‐order habitat selection by comparing used locations to random ones within their home ranges. Third‐order habitat selection for habitat type and composition was weak and varied little between seasons. We found that patterns of fox habitat selection did not mirror habitat selection of Hodotermes (quantified using RSFs), even when feeding on them (4th‐order). Taken together, these results might indicate that bat‐eared foxes’ food resources are homogenously distributed across habitats and that prey other than Hodotermes play an important role in bat‐eared foxes’ space use.     

Evolution of nocturnality in bats: Potential competitors and predators during their early history

Despite their taxonomic and ecological diversity, modern bats (Order Chiroptera) are almost exclusively nocturnal. This behaviour is too ubiquitous to be explained by common patterns of temporal variation in availability of their diverse food sources or by the risk of hyperthermia when flying during the day. Other explanations for bat nocturnality include competition and increased predation risk from birds during the day. In the early and mid Eocene, the known bat fauna consisted of several insectivorous species of sizes similar to those of the modern European assemblage. This fauna was contemporaneous with several species of predatory birds, including owls (Strigiformes), hawks (Accipitridae), falcons (Falconidae) and rollers (Coraciiformes), which were the same size as modern predators on bats. Predation risk could therefore have been a significant factor preventing the early bats from becoming diurnal. Competition from aerial insectivorous birds, however, was less likely to have been significant for bats during the early Eocene, since very few such groups, mainly small Aegialornithidae, were present, with most of the major groups of aerial insectivores evolving later.     

Feeding behaviour of the bats Nycteris grandis and Nycteris thebaica (Nycteridae) in captivity

The feeding and hunting behaviour of Nycteris grandis and N. thebaica was observed in captivity at the Sengwa Wildlife Research Area in Zimbabwe in January and February 1982. Both species preferentially selected katydids and beetles over moths, and relied heavily on acoustic stimuli emanating from prey to detect targets. Nycteris grandis readily consumed frogs and bats and appeared not to use the calls of male frogs or the echolocation calls of other bats to locate prey. Both species produced echolocation calls during attacks on prey, increasing the rates of pulse repetition as they closed with targets and suggesting the use of echolocation in hunting. The echolocation calls of N. grandis are described along with general observations of the behaviour of both species.