Lipid composition of the stratum corneum in different regions of the body of Kuhl's pipistrelle from the Negev Desert, Israel.

The most superficial epidermal layer in endotherms is the stratum corneum (SC), which is composed of dead corneocytes embedded in a lipid matrix with free fatty acids, cholesterol, ceramides, and cerebrosides; the lipid composition of the SC determines its permeability to water vapor. Lipids that are more polar, have longer hydrocarbon chains, and are less bulky are often packed in more ordered phase states to slow cutaneous evaporative water loss (CEWL); these lipids also resist transitions to more disordered phases at high ambient temperatures (Ta). In bats, wing and tail membranes (wing patagia and tail uropatagium, respectively) allow powered flight, but increase surface area, and hence CEWL, with implications for survival in arid environments. We captured Pipistrellus kuhlii from an arid habitat and measured the lipid composition of the SC of the plagiopatagium in the wing, the uropatagium, and the non-membranous region (NMR) of the body using thin layer chromatography and reversed phase high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. The patagia contained more cholesterol and shorter-chained ceramides, and fewer cerebrosides than the NMR, indicating that the lipid phase transition temperature in the patagia is lower than in the NMR. Thus, at moderate Ta the lipids in the SC in all body regions will remain in an ordered phase state, allowing water conservation; but as Ta increases, the lipids in the SC of the patagia will more easily transition into a disordered phase, resulting in increased CEWL from the patagia facilitating efficient heat dissipation in hot environments.

An appetite for pests: Synanthropic insectivorous bats exploit cotton pest irruptions and consume various deleterious arthropods.

Conservation biological control (CBC) seeks to minimize the deleterious effects of agricultural pests by enhancing the efficiency of natural enemies. Despite the documented potential of insectivorous bats to consume pests, many synanthropic bat species are still underappreciated as beneficial species. We investigated the diet of Kuhl's pipistrelle (Pipistrellus kuhlii), a common synanthropic insectivorous bat that forages in urban and agricultural areas, to determine whether it may function as a natural enemy in CBC. Faecal samples of P. kuhlii were collected throughout the cotton-growing season from five roost sites near cotton fields located in a Mediterranean agroecosystem, Israel, and analyzed using DNA metabarcoding. Additionally, data on estimated abundance of major cotton pests were collected. We found that the diet of P. kuhlii significantly varied according to sites and dates and comprised 27 species of agricultural pests that were found in 77.2% of the samples, including pests of key economic concern. The dominant prey was the widespread cotton pest, the pink bollworm, Pectinophora gossypiella, found in 31% of the samples and in all the roosts. Pink bollworm abundance was positively correlated with its occurrence in the bat diet. Furthermore, the bats' dietary breadth narrowed, while temporal dietary overlap increased, in relation to increasing frequencies of pink bollworms in the diet. This suggests that P. kuhlii exploits pink bollworm irruptions by opportunistic feeding. We suggest that synanthropic bats provide important pest suppression services, may function as CBC agents of cotton pests and potentially contribute to suppress additional deleterious arthropods found in their diet in high frequencies.

Evaporative water loss in Kuhl's pipistrelles declines along an environmental gradient, from mesic to hyperarid.

Intraspecific variation in animal energy and water balances may play an important role in local adaptation of populations to specific habitats such as deserts. We examined Kuhl's pipistrelle (Pipistrellus kuhlii), a common bat in Israel that ranges in distribution from mesic Mediterranean to hyperarid desert habitats, for intraspecific differences in metabolic rate (MR) and evaporative water loss (EWL) among populations along a climatic gradient. We tested the prediction that EWL, especially at high ambient temperatures is lower in Kuhl's pipistrelles from desert habitats than from mesic habitats. We measured MR and total evaporative water loss (TEWL) at four ambient temperatures (10 °C, 20 °C, 30 °C and 35 °C) in three groups of bats using open-flow respirometry. We fitted the bats with a mask to separate cutaneous water loss (CWL) from respiratory water loss (RWL) at 35 °C. At 35 °C, mean TEWL in the southernmost group, from the hyperarid location, was significantly lower than in the other two groups, with no apparent difference in mean MR. The source of difference TEWL was that the southern group had significantly lower CWL than the other two groups; RWL did not differ among them. This suggests that there are mechanisms that reduce EWL from the skin of the bats; a likely candidate is modification of the lipids in the outer layer of the dermis that make the skin possibly less permeable to water as has been described in birds and a few other species of bat.

Ultrasound avoidance by flying antlions (Myrmeleontidae).

The acoustic arms race between insectivorous bats and their invertebrate prey has led to the convergent evolution of ultrasound hearing in seven orders of nocturnal insects. Upon hearing the echolocation calls of an approaching bat, such insects take defensive action. Here, we document a previously unknown sense of ultrasound hearing and phonotactic flight behaviour in the neuropteran family Myrmeleontidae (antlions). The antlion Myrmeleon hyalinus was presented with sound pulses at ultrasonic frequencies used by echolocating bats and its response thresholds in tethered flight determined. Behaviours included abdominal twitches, wing flicks, brief pauses in flight and flight cessation. Such behaviours create erratic evasive flight manoeuvres in other eared insects, particularly mantids and lacewings. Antlions responded best to ultrasound between 60 and 80 kHz (75 dB peSPL at 80 kHz), showing response thresholds similar to those of the related lacewings (Neuroptera, Chrysopidae). Yet, at lower ultrasonic frequencies (20-50 kHz), antlions were far less sensitive than lacewings. Based on calculated response distances, we conclude that antlions respond only after having been detected by bats rather than using early evasive flights. We argue that the high response threshold for low-frequency ultrasound is adaptive for an insect that is mainly active close to and within vegetation, because a behavioural response to the lower ultrasonic frequencies used by high-flying bats would result in evasive action in the absence of actual predation risk.

The effect of water contamination and host-related factors on ectoparasite load in an insectivorous bat.

We examined the effects of sex, age, and reproductive state of the insectivorous bat Pipistrellus kuhlii on the abundance and prevalence of arthropod ectoparasites (Macronyssidae and Cimicidae) in habitats with either sewage-polluted or natural bodies of water, in the Negev Desert, Israel. We chose water pollution as an environmental factor because of the importance of water availability in desert environments, particularly for P. kuhlii, which needs to drink on a daily basis. We predicted that parasite infestation rates would be affected by both environment and demographic cohort of the host. We found that female bats in the polluted site harbored significantly more mites than female bats in the natural site and that juveniles in the polluted site harbored significantly more cimicid individuals than juveniles in the natural site. We further found that age and sex (host-related factors) affected ectoparasite prevalence and intensity (i.e., the abundance of parasites) in the polluted site. Our results may suggest that the interaction between host-related and environment-related factors affected parasite infestations, with females and young bats being more susceptible to ectoparasites when foraging over polluted water. This effect may be particularly important for bats that must drink or forage above water for other wildlife that depend on drinking water for survival.

The cutaneous lipid composition of bat wing and tail membranes: a case of convergent evolution with birds.

The water vapour permeability barrier of mammals and birds resides in the stratum corneum (SC), the outermost layer of the epidermis. The molar ratio and molecular arrangement of lipid classes in the SC determine the integrity of this barrier. Increased chain length and polarity of ceramides, the most abundant lipid class in mammalian SC, contribute to tighter packing and thus to reduced cutaneous evaporative water loss (CEWL). However, tighter lipid packing also causes low SC hydration, making it brittle, whereas high hydration softens the skin at the cost of increasing CEWL. Cerebrosides are not present in the mammalian SC; their pathological accumulation occurs in Gaucher's disease, which leads to a dramatic increase in CEWL. However, cerebrosides occur normally in the SC of birds. We tested the hypothesis that cerebrosides are also present in the SC of bats, because they are probably necessary to confer pliability to the skin, a quality needed for flight. We examined the SC lipid composition of four sympatric bat species and found that, as in birds, their SC has substantial cerebroside contents, not associated with a pathological state, indicating convergent evolution between bats and birds.

Metabolic rate, evaporative water loss and thermoregulatory state in four species of bats in the Negev desert.

Life in deserts is challenging for bats because of their relatively high energy and water requirements; nevertheless bats thrive in desert environments. We postulated that bats from desert environments have lower metabolic rates (MR) and total evaporative water loss (TEWL) than their mesic counterparts. To test this idea, we measured MR and TEWL of four species of bats, which inhabit the Negev desert in Israel, one species mainly restricted to hyper-arid deserts (Otonycteris hemprichii), two species from semi-desert areas (Eptesicus bottae and Plecotus christii), and one widespread species (Pipistrellus kuhlii). We also measured separately, in the same individuals, the two components of TEWL, respiratory water loss (RWL) and cutaneous evaporative water loss (CEWL), using a mask. In all the species, MR and TEWL were significantly reduced during torpor, the latter being a consequence of reductions in both RWL and CEWL. Then, we evaluated whether MR and TEWL in bats differ according to their geographic distributions, and whether those rates change with Ta and the use of torpor. We did not find significant differences in MR among species, but we found that TEWL was lowest in the species restricted to desert habitats, intermediate in the semi-desert dwelling species, and highest in the widespread species, perhaps a consequence of adaptation to life in deserts. Our results were supported by a subsequent analysis of data collected from the literature on rates of TEWL for 35 bat species from desert and mesic habitats.

A whispering bat that screams: bimodal switch of foraging guild from gleaning to aerial hawking in the desert long-eared bat.

Echolocating bats have historically been classified as either loud aerial hawkers or whispering gleaners. Some bat species can forage in multiple ways and others have demonstrated limited flexibility in the amplitude of their echolocation calls. The desert long-eared bat, Otonycteris hemprichii, has been said to be a passive gleaning whispering bat preying on terrestrial arthropods such as scorpions. Using an acoustic tracking system, we recorded individuals flying at foraging and drinking sites and compared their flight height, flight speed, call duration, pulse interval and source levels with those of gleaning individuals previously recorded using the same setup. We found differences in all variables with the strongest difference in source levels, where bats called at a mean of 119 dB peSPL (compared with 75 dB peSPL when gleaning). Bat faecal analysis indicated that their diet differed from previous studies and that prey species were capable of flight. We conclude that the bats switched from passive gleaning to capturing airborne insects (aerial hawking). Although whispering bats have been known to opportunistically catch insects on the wing, in the present study we show a full bimodal switch between foraging guilds with the respective changes in source level to those typical of a true aerial hawker.

Climatic gradients and forest composition shape bat communities in Eastern Mediterranean pine plantations.

Biotic and abiotic factors can act as filters for determining the species composition of biological communities. We aimed to identify abiotic factors driving the assembly of bat communities in Eastern Mediterranean pine plantations along a north-south climatic gradient, as they are crucial forest habitats for the assessment and conservation of these communities. We expected that bat communities are predominantly shaped by environmental filtering. We conducted acoustic sampling in 35 pine plantations in Israel and analyzed recordings for species identification. We used the ESLTP analysis, an extension of the three-table ordination (RLQ analysis), to explore relationships between environmental characteristics, species occurrences, and functional traits of species while accounting for phylogenetic relationships between species and spatial distribution of the communities. Communities showed phylogenetic and trait clustering. Climatic conditions and forest vegetation composition shaped communities of bats, affecting the distribution of traits related to foraging behaviors, vegetation clutter, and the ability of bats to maneuver in it. Maneuverable species were associated with the northern Mediterranean climatic zone, with a scarce cover of drought-tolerant small shrubs and grassland. Fast flyers were associated with the center-south semi-arid area, with abundant drought-tolerant small shrubs and grassland. These forces might have a predominant role in the assembly of these communities, presumably due to the stressful climatic conditions of the study area. The ESLTP approach can be extended to other taxa and environments to predict species responses to disturbance and environmental changes and give insights into environmental management.

Waking to drink: rates of evaporative water loss determine arousal frequency in hibernating bats.

Bats hibernate to cope with low ambient temperatures (T(a)) and low food availability during winter. However, hibernation is frequently interrupted by arousals, when bats increase body temperature (T(b)) and metabolic rate (MR) to normothermic levels. Arousals account for more than 85% of a bat's winter energy expenditure. This has been associated with variation in T(b), T(a) or both, leading to a single testable prediction, i.e. that torpor bout length (TBL) is negatively correlated with T(a) and T(b). T(a) and T(b) were both found to be correlated with TBL, but correlations alone cannot establish a causal link between arousal and T(b) or T(a). Because hydration state has also been implicated in arousals from hibernation, we hypothesized that water loss during hibernation creates the need in bats to arouse to drink. We measured TBL of bats (Pipistrellus kuhlii) at the same T(a) but under different conditions of humidity, and found an inverse relationship between TBL and total evaporative water loss, independent of metabolic rate, which directly supports the hypothesis that hydration state is a cue to arousal in bats.

A species-level trait dataset of bats in Europe and beyond.

Knowledge of species' functional traits is essential for understanding biodiversity patterns, predicting the impacts of global environmental changes, and assessing the efficiency of conservation measures. Bats are major components of mammalian diversity and occupy a variety of ecological niches and geographic distributions. However, an extensive compilation of their functional traits and ecological attributes is still missing. Here we present EuroBaTrait 1.0, the most comprehensive and up-to-date trait dataset covering 47 European bat species. The dataset includes data on 118 traits including genetic composition, physiology, morphology, acoustic signature, climatic associations, foraging habitat, roost type, diet, spatial behaviour, life history, pathogens, phenology, and distribution. We compiled the bat trait data obtained from three main sources: (i) a systematic literature and dataset search, (ii) unpublished data from European bat experts, and (iii) observations from large-scale monitoring programs. EuroBaTrait is designed to provide an important data source for comparative and trait-based analyses at the species or community level. The dataset also exposes knowledge gaps in species, geographic and trait coverage, highlighting priorities for future data collection.

Effects of host diet and thermal state on feeding performance of the flea Xenopsylla ramesis.

We examined feeding performance of the flea Xenopsylla ramesis on three different hosts: its natural, granivorous, rodent host, Sundevall's jird (Meriones crassus); the frugivorous Egyptian fruit bat (Rousettus aegyptiacus); and an insectivorous bat, Kuhl's pipistrelle (Pipistrellus kuhlii). Because these fleas are not known to occur on bats, we hypothesized that the fleas' feeding performance (i.e. feeding and digestion rates) would be higher when feeding on their natural host than on either of the bats that they do not naturally parasitize. We found that mass-specific blood-meal size of both male and female fleas was significantly lower when feeding on Kuhl's pipistrelles than on the other two species, but was not different in female fleas feeding on fruit bats or on jirds at all stages of digestion. However, more male fleas achieved higher levels of engorgement if they fed on Sundevall's jirds than if they fed on Egyptian fruit bats. The fleas digested blood of fruit bats and jirds significantly faster than blood of Kuhl's pipistrelle. In addition, after a single blood meal, the survival time of fleas fed on normothermic Kuhl's pipistrelles was significantly shorter than that of fleas fed on Sundevall's jirds and even lower when male fleas fed on Egyptian fruit bats. Thus, our prediction was partially supported: normothermic Kuhl's pipistrelles were inferior hosts for fleas compared with Sandevall's jirds and Egyptian fruit bats. Interestingly, the proportion of engorged fleas that fed on torpid Kuhl's pipistrelles was significantly higher than the proportion of the fleas that fed on normothermic individuals, indicating that becoming torpid might be a liability, rather than an effective defense against parasites.

Habitat aridity as a determinant of the trade-off between water conservation and evaporative heat loss in bats.

The maintenance of water balance in arid environments might represent a formidable challenge for Chiroptera, since they have high surface-to-volume ratios. In deserts, bats conserve water, for example, using daily torpor, but they also might experience episodic heat bouts, when they may need to increase total evaporative water loss (TEWL) to thermoregulate. We hypothesized that in bats, habitat aridity and its variability determine a trade-off between water conservation and thermoregulation via evaporative means. To test this hypothesis, we collated data from the literature of 22 species of bats on TEWL, body temperature and resting metabolic rate, in torpor and euthermy. We also collected data on ambient temperature (Ta) and precipitation of the locations where bats were captured, calculated an aridity index, and built an index of variability of the environment. After correcting for phylogeny, we found that, as aridity and variability of the environment increased, bats had lower values of TEWL, but the rate at which TEWL increases with Ta was higher, supporting our hypothesis. These results suggest that at high Ta there is a trade-off between water conservation and evaporative heat loss in bats. The evolution of physiological mechanisms that allow water conservation and tolerance to conditions of high Ta without access to free water might thus be crucial to explain the distribution of desert bats.

Hemprich's long-eared bat (Otonycteris hemprichii) as a predator of scorpions: whispering echolocation, passive gleaning and prey selection.

Over 70% of the droppings of the gleaning bat Otonycteris hemprichii can contain scorpion fragments. Yet, some scorpions found in its desert habitat possess venom of the highest known toxicity, rendering them a very dangerous prey. In this study, we describe how O. hemprichii catches and handles scorpions, quantify its flight and echolocation behaviour in the field, investigate what sensory modality it uses to detect scorpions, and test whether it selects scorpions according to their size or toxicity. We confirmed that O. hemprichi is a whispering bat (approx. 80 dB peSPL) with short, multi-harmonic calls. In a flight room we also confirmed that O. hemprichii detects scorpions by their walking noises. Amplitudes of such noises were measured and they reach the flying bat at or below the level of echoes of the loess substrate. Bats dropped straight onto moving scorpions and were stung frequently even straight in their face. Stings did not change the bats' behaviour and caused no signs of poisoning. Scorpions were eaten including poison gland and stinger. Bats showed no preference neither for any of the scorpion species nor their size suggesting they are generalist predators with regard to scorpions.

Does interspecific competition drive patterns of habitat use in desert bat communities?

Bodies of water are a key foraging habitat for insectivorous bats. Since water is a scarce and limiting resource in arid environments, bodies of open water may have a structuring effect on desert bat communities, resulting in temporal or spatial partitioning of bat activity. Using acoustic monitoring, we studied the spatial and temporal activity patterns of insectivorous bats over desert ponds, and hypothesised that sympatric bat species partition the foraging space above ponds based on interspecific competitive interactions. We used indirect measures of competition (niche overlap and competition coefficients from the regression method) and tested for differences in pond habitat selection and peak activity time over ponds. We examined the effect of changes in the activity of bat species on their potential competitors. We found that interspecific competition affects bat community structure and activity patterns. Competing species partitioned their use of ponds spatially, whereby each species was associated with different pond size and hydroperiod (the number of months a pond holds water) categories, as well as temporally, whereby their activity peaked at different hours of the night. The drying out of temporary ponds increased temporal partitioning over permanent ponds. Differences in the activity of species over ponds in response to the presence or absence of their competitors lend further support to the role of interspecific competition in structuring desert bat communities. We suggest that habitat use and night activity pattern of insectivorous bats in arid environments reflect the trade-offs between selection of preferred pond type or activity time and constraints posed by competitive interactions.

Context-dependent flight speed: evidence for energetically optimal flight speed in the bat Pipistrellus kuhlii?

1. Understanding the causes and consequences of animal flight speed has long been a challenge in biology. Aerodynamic theory is used to predict the most economical flight speeds, minimizing energy expenditure either per distance (maximal range speed, Vmr) or per time (minimal power speed, Vmp). When foraging in flight, flight speed also affects prey encounter and energy intake rates. According to optimal flight speed theory, such effects may shift the energetically optimal foraging speed to above Vmp. 2. Therefore, we predicted that if energetic considerations indeed have a substantial effect on flight speed of aerial-hawking bats, they will use high speed (close to Vmr) to commute from their daily roost to the foraging sites, while a slower speed (but still above Vmp) will be preferred during foraging. To test these predictions, echolocation calls of commuting and foraging Pipistrellus kuhlii were recorded and their flight tracks were reconstructed using an acoustic flight path tracking system. 3. Confirming our qualitative prediction, commuting flight was found to be significantly faster than foraging flight (9.3 vs. 6.7 m s(-1)), even when controlling for its lower tortuosity. 4. In order to examine our quantitative prediction, we compared observed flight speeds with Vmp and Vmr values generated for the study population using two alternative aerodynamic models, based on mass and wing morphology variables measured from bats we captured while commuting. The Vmp and Vmr values generated by one of the models were much lower than our measured flight speed. According to the other model used, however, measured foraging flight was faster than Vmp and commuting flight slightly slower than Vmr, which is in agreement with the predictions of optimal flight speed theory. 5. Thus, the second aerodynamic model we used seems to be a reasonable predictor of the different flight speeds used by the bats while foraging and while commuting. This supports the hypothesis that bats fly at a context-dependent, energetically optimal flight speed.

The influence of ethanol on feeding in the frugivorous yellow-vented bulbul (Pycnonotus xanthopygos).

The sugars in fleshy fruits provide a rich source of energy to frugivorous animals. However, these carbohydrates also serve as a substrate for alcoholic fermentation by yeasts, ethanol being the main by-product of this process. Ethanol ingestion via frugivory thus occurs in a diverse assemblage of invertebrate and vertebrate taxa, including numerous species of birds. We tested the roles of ethanol as an odor cue for resource location by adult yellow-vented bulbuls (Pycnonotus xanthopygos) and as a possible appetite stimulant in feeding trials with artificial food. We hypothesized (1) that the odor of ethanol does not serve as a food-locating cue in diurnal frugivorous passerine birds, and predicted that the choice of food source and the mass of food eaten by such birds will not be influenced by the odor of ethanol. We further hypothesized (2) that food intake in passerine birds is affected by ingestion of ethanol according to its concentration [EtOH], and predicted that food intake will follow a bell-shaped curve in relation to [EtOH]. In accord with hypothesis (1) and its prediction, we found that the odor of ethanol did not affect food preferences, in either ethanol-naïve or ethanol-experienced yellow-vented bulbuls, when presented at concentrations found in naturally ripe fruit (0.0-1.0%); this suggests that the odor of ethanol is not a food-locating cue for the bulbuls. Hypothesis (2) was partially supported, namely at low [EtOH] (0-3%), food intake was constant and at high [EtOH] (3%) food intake decreased, following only the right half of the predicted bell-shaped response. Ethanol-naïve birds showed no preference towards any [EtOH] presented in two-way choice trials. However, daily food intake in ethanol-experienced bulbuls in single option trials decreased by an average of 36% when the artificial food contained the highest tested concentration of ethanol (3.0%). We suggest that decreasing food intake when food ethanol concentration is relatively high may be a means of avoiding intoxication and is related to the ethanol-metabolizing ability of the bird.

Energy metabolism and evaporative water loss in the European free-tailed bat and Hemprich's long-eared bat (Microchiroptera): species sympatric in the Negev Desert.

We compared the thermoregulatory abilities of two insectivorous bat species, Tadarida teniotis (mean body mass 32 g) and Otonycteris hemprichii (mean body mass 25 g), that are of different phylogenetic origins and zoogeographic distributions but are sympatric in the Negev Desert. At night, both were normothermic. By day, both were torpid when exposed to ambient temperatures (T(a)) below 25 degrees Celsius, with concomitant adjustments in metabolic rate (MR). Otonycteris hemprichii entered torpor at higher T(a) than T. teniotis, and, when torpid, their body temperatures (T(b)) were 1 degrees -2 degrees Celsius and 5 degrees -8 degrees Celsius above T(a), respectively; MR was correspondingly reduced. At night, the lower critical temperature of T. teniotis was 31.5 degrees Celsius, and that of O. hemprichii was 33 degrees Celsius. Mean nocturnal thermoneutral MR of T. teniotis was 37% greater than that of O. hemprichii. At high T(a), evaporative water loss (EWL) increased markedly in both species, but it was significantly higher in T. teniotis above 38 degrees Celsius. In both species, the dry heat transfer coefficient (thermal conductance) followed the expected pattern for small mammals, by day and by night. Total EWL was notably low in normothermic and torpid animals of both species, much lower than values reported for other bats, indicating efficient water conservation mechanisms in the study species. Comparing thermoregulatory abilities suggests that O. hemprichii is better adapted to hot, arid environments than T. teniotis, which may explain its wider desert distribution. By both standard and phylogenetically informed ANCOVA, we found no differences in basal metabolic rate (BMR) between desert and nondesert species of insectivorous bats, substantiating previous studies suggesting that low BMR is a characteristic common to insectivorous bats in general.

The influence of ambient temperature and the energy and protein content of food on nitrogenous excretion in the Egyptian fruit bat (Rousettus aegyptiacus).

The diets of frugivorous and nectarivorous vertebrates contain much water and generally have high energy but low protein contents. Therefore, we tested the prediction that to save energy under conditions of high energy demands and high water intake, frugivorous Egyptian fruit bats (Rousettus aegyptiacus) will increase both the absolute quantity and the proportion of ammonia in their urine. We also examined whether such changes occur when protein intake is low and water intake is high. We did three feeding trials. In trials 1 and 2, bats were fed one of four liquid diets containing constant soy protein concentrations but varying in sucrose concentration and were kept at ambient temperatures (T(a)) of 30 degrees Celsius and 12 degrees Celsius, respectively. In trial 3, bats were kept at Ta=12 degrees Celsius and fed one of four liquid diets with equal sucrose concentrations but varying protein concentrations. In trial 1, food intake at a sucrose concentration of 256 mmol/kg H(2)O was initially high but decreased to a constant rate with further increases in sucrose concentration, while in trial 2, food intake decreased exponentially with increasing sucrose concentration. As predicted, at 12 degrees Celsius with varying sucrose concentration, both the absolute quantity and the fraction of ammonia in the bats' urine increased significantly with food intake (P<0.02), while the absolute quantity of urea and the fraction of urea nitrogen excreted decreased significantly with food intake (P<0.03). Varying sucrose concentration had no significant effect on nitrogen excretion at Ta=30 degrees Celsius. Varying protein concentration had no significant effect on nitrogen excretion at Ta=12 degrees Celsius. We suggest that Egyptian fruit bats can increase ammonia excretion in response to increased energetic demands, and we calculate that they can save energy equal to approximately 2% of their daily metabolic rate by doing so.

Dynamics of jamming avoidance in echolocating bats.

Animals using active sensing systems such as echolocation or electrolocation may experience interference from the signals of neighbouring conspecifics, which can be offset by a jamming avoidance response (JAR). Here, we report JAR in one echolocating bat (Tadarida teniotis: Molossidae) but not in another (Taphozous perforatus: Emballonuridae) when both flew and foraged with conspecifics. In T. teniotis, JAR consisted of shifts in the dominant frequencies of echolocation calls, enhancing differences among individuals. Larger spectral overlap of signals elicited stronger JAR. Tadarida teniotis showed two types of JAR: (i) for distant conspecifics: a symmetric JAR, with lower- and higher-frequency bats shifting their frequencies downwards and upwards, respectively, on average by the same amount; and (ii) for closer conspecifics: an asymmetric JAR, with only the upper-frequency bat shifting its frequency upwards. In comparison, 'wave-type' weakly electric fishes also shift frequencies of discharges in a JAR, but unlike T. teniotis, the shifts are either symmetric in some species or asymmetric in others. We hypothesize that symmetric JAR in T. teniotis serves to avoid jamming and improve echolocation, whereas asymmetric JAR may aid communication by helping to identify and locate conspecifics, thus minimizing chances of mid-air collisions.