The transfer function of a target limits the jitter detection threshold with signals of echolocating FM-bats

The delay jitter discrimination threshold in bats is a disputed subject. Some investigators have obtained results indicating that bats are able to discriminate alternations in delay down to 10 ns, which appears incredible for purely physical reasons. Using actual bat echolocation sequences recorded during an easy detection task to measure simulated delay jitter, it is shown here that jitter detection thresholds in the order of some tens of nanoseconds are actually physically realizable. However, if the transfer function of the target simulating apparatus is not perfect, the lowest thresholds are in the order of hundreds of nanoseconds and variable between individual bats. This phenomenon is shown to arise as a consequence of the variation in signal parameters from call to call. When the transfer function from a real jitter experiment was artificially applied to the echoes, the jitter detection thresholds again were several hundred nanoseconds. This is the first study to point out a limiting role of the transfer function of a system faced with variations in echolocation signal parameters, something that should be considered in evaluating all sonar systems with variable signal structure.     

Detection of jitter in intertarget spacing by the big brown bat Eptesicus fuscus

We trained bats to detect intertarget jitter, i.e., relative motion between two virtual (electronically synthesized) targets. Both targets were themselves moving with respect to nearby objects (e.g., the microphone and speaker used to create the virtual targets) so that the only reliable cue available to the bats was variation in intertarget spacing. Given a target at 80 cm and another at 95, 110 or 125 cm, the threshold for intertarget jitter (ITJ) of the two bats tested was <10 μs, corresponding to <1.7 mm of range. When, for one bat, we increased the range instability of the targets by adding varying amounts of random range shift to the target complex (while preserving the correct intertarget spacing), ITJ threshold worsened. When we presented three targets, one of which was jittering, the bat's threshold improved to 0.9 μs (equivalent to 0.16 mm). If no second target was presented, i.e., if the task was to detect jitter added to a single moving target, then bats' jitter threshold was very high (>200 μs). Eptesicus fuscus appears to be very good at detecting changes in intertarget spacing, which might prove valuable for detecting targets moving relative to the background or for constructing a spatial image of a complex environment. Accepted: 7 April 1997     

Phase sensitivity in bat sonar revisited

An echolocating bat produces echoes consisting of the convolution of echolocation call and the impulse response (IR) of the ensonified object. A crucial question in animal sonar is whether bats are able to extract this IR from the echo. The bat inner ear generates a frequency representation of call and echo and IR extraction in the frequency domain requires accurate analysis of both magnitude and phase information. Previous studies investigating the phase sensitivity of bats using a jitter paradigm reported a temporal acuity down to 10 ns, suggesting perfect sonar phase representation. In a phantom-target playback experiment, we investigate the perceptual phase sensitivity of the bat Phyllostomus discolor using a novel approach: instead of manipulating IR phase by changing IR delay (jitter paradigm), we randomized IR phase and thus lengthened the IR over time, leaving the magnitude spectrum unchanged. Our results show that phase sensitivity, as reflected in the analysis of signal duration, appears to be much lower than phase sensitivity, as reflected in the analysis of signal onset. The current data indicate that different temporal aspects of sonar processing are encoded with very different temporal resolution and thus an overall claim of “phase sensitivity” as such cannot be maintained.     

How greater mouse-eared bats deal with ambiguous echoic scenes

Echolocating bats have to assign the received echoes to the correct call that generated them. Failing to do so will result in the perception of virtual targets that are positioned where there is no actual target. The assignment of echoes to the emitted calls can be ambiguous especially if the pulse intervals between calls are short and kept constant. Here, we present first evidence that greater mouse-eared bats deal with ambiguity by changing the pulse interval more often, in particular by reducing the number of calls in the terminal group before landing. This strategy separates virtual targets from real ones according to their change in position. Real targets will always remain in a constant position, and virtual targets will jitter back and forth according to the change in the time interval.     

Rain increases the energy cost of bat flight

Similar to insects, birds and pterosaurs, bats have evolved powered flight. But in contrast to other flying taxa, only bats are furry. Here, we asked whether flight is impaired when bat pelage and wing membranes get wet. We studied the metabolism of short flights in Carollia sowelli, a bat that is exposed to heavy and frequent rainfall in neotropical rainforests. We expected bats to encounter higher thermoregulatory costs, or to suffer from lowered aerodynamic properties when pelage and wing membranes catch moisture. Therefore, we predicted that wet bats face higher flight costs than dry ones. We quantified the flight metabolism in three treatments: dry bats, wet bats and no rain, wet bats and rain. Dry bats showed metabolic rates predicted by allometry. However, flight metabolism increased twofold when bats were wet, or when they were additionally exposed to rain. We conclude that bats may not avoid rain only because of sensory constraints imposed by raindrops on echolocation, but also because of energetic constraints.     

Oceanic records of North American bats and implications for offshore wind energy development in the United States

Offshore wind energy is a growing industry in the United States, and renewable energy from offshore wind is estimated to double the country''s total electricity generation. There is growing concern that land‐based wind development in North America is negatively impacting bat populations, primarily long‐distance migrating bats, but the impacts to bats from offshore wind energy are unknown. Bats are associated with the terrestrial environment, but have been observed over the ocean. In this review, we synthesize historic and contemporary accounts of bats observed and acoustically recorded in the North American marine environment to ascertain the spatial and temporal distribution of bats flying offshore. We incorporate studies of offshore bats in Europe and of bat behavior at land‐based wind energy studies to examine how offshore wind development could impact North American bat populations. We find that most offshore bat records are of long‐distance migrating bats and records occur during autumn migration, the period of highest fatality rates for long‐distance migrating bats at land‐based wind facilities in North America. We summarize evidence that bats may be attracted to offshore turbines, potentially increasing their exposure to risk of collision. However, higher wind speeds offshore can potentially reduce the amount of time that bats are exposed to risk. We identify knowledge gaps and hypothesize that a combination of operational minimization strategies may be the most effective approach for reducing impacts to bats and maximizing offshore energy production.     

Network analysis of host–virus communities in bats and rodents reveals determinants of cross‐species transmission

Bats are natural reservoirs of several important emerging viruses. Cross‐species transmission appears to be quite common among bats, which may contribute to their unique reservoir potential. Therefore, understanding the importance of bats as reservoirs requires examining them in a community context rather than concentrating on individual species. Here, we use a network approach to identify ecological and biological correlates of cross‐species virus transmission in bats and rodents, another important host group. We show that given our current knowledge the bat viral sharing network is more connected than the rodent network, suggesting viruses may pass more easily between bat species. We identify host traits associated with important reservoir species: gregarious bats are more likely to share more viruses and bats which migrate regionally are important for spreading viruses through the network. We identify multiple communities of viral sharing within bats and rodents and highlight potential species traits that can help guide studies of novel pathogen emergence.     

Do predators influence the behaviour of bats?

Many aspects of animal behaviour are affected by real‐time changes in the risk of predation. This conclusion holds for virtually all taxa and ecological systems studied, but does it hold for bats? Bats are poorly represented in the literature on anti‐predator behaviour, which may reflect a lack of nocturnal predators specialized on bats. If bats actually experience a world with minimal anti‐predator concerns, then they will provide a unique contrast within the realm of vertebrate ecology. Alternatively, such predator‐driven behaviour in bats may not yet be fully understood, given the difficulties in working with these highly mobile and nocturnal animals. We provide a wide‐ranging exploration of these issues in bat behaviour. We first cover the basic predator‐prey information available on bats, both on potential predators and the ways in which bats might perceive predators and respond to attacks. We then cover work relevant to key aspects of bat behaviour, such as choice of daytime roosts, the nature of sleep and torpor, evening roost departures, moonlight avoidance, landscape‐related movement patterns, and habitat selection. Overall, the evidence in favour of a strong influence of predators on bat behaviour is equivocal, with the picture clouded by contradictory results and a lack of information on potential predators and the perception of risk by bats. It seems clear that day‐active bats run a considerable risk of being killed by diurnal raptors, which are able to capture bats with relative ease. Thus, bats taking advantage of a pulse of insects just prior to sunset are likely taking risks to gain much‐needed energy. Further, the choice of daytime roosts by bats is probably strongly influenced by roost safety. Few studies, however, have directly addressed either of these topics. As a group, insectivorous temperate‐zone bats show no clear tendency to avoid apparently risky situations, such as activity on moonlit nights. However, some observations are consistent with the idea that predation risk affects choice of movement paths and feeding areas by temperate‐zone bats, as well as the timing of roost departures. The behaviour of tropical bats, on the other hand, seems more generally influenced by predators; this is especially true for tropical nectarivores and frugivores, but also for insectivorous bats. Presumably there are more serious predators on bats in the tropics (e.g. specialized raptors or carnivorous bats), but the identity of these predators is unclear. More information is needed to assess fully the influence of predators on bat behaviour. There is much need for work on the ways in which bats perceive predators via auditory, visual, and olfactory cues, and whether bats have some knowledge of the risks posed by different predators. Also needed is information on how predators attack bats and how bats react to attacking predators. Difficult to obtain, but of critical value, will be information on the nature of the predation risk experienced by bats while away from roosts and during the full darkness of night.     

Alphacoronaviruses in New World bats: prevalence, persistence, phylogeny, and potential for interaction with humans

Bats are reservoirs for many different coronaviruses (CoVs) as well as many other important zoonotic viruses. We sampled feces and/or anal swabs of 1,044 insectivorous bats of 2 families and 17 species from 21 different locations within Colorado from 2007 to 2009. We detected alphacoronavirus RNA in bats of 4 species: big brown bats (Eptesicus fuscus), 10% prevalence; long-legged bats (Myotis volans), 8% prevalence; little brown bats (Myotis lucifugus), 3% prevalence; and western long-eared bats (Myotis evotis), 2% prevalence. Overall, juvenile bats were twice as likely to be positive for CoV RNA as adult bats. At two of the rural sampling sites, CoV RNAs were detected in big brown and long-legged bats during the three sequential summers of this study. CoV RNA was detected in big brown bats in all five of the urban maternity roosts sampled throughout each of the periods tested. Individually tagged big brown bats that were positive for CoV RNA and later sampled again all became CoV RNA negative. Nucleotide sequences in the RdRp gene fell into 3 main clusters, all distinct from those of Old World bats. Similar nucleotide sequences were found in amplicons from gene 1b and the spike gene in both a big-brown and a long-legged bat, indicating that a CoV may be capable of infecting bats of different genera. These data suggest that ongoing evolution of CoVs in bats creates the possibility of a continued threat for emergence into hosts of other species. Alphacoronavirus RNA was detected at a high prevalence in big brown bats in roosts in close proximity to human habitations (10%) and known to have direct contact with people (19%), suggesting that significant potential opportunities exist for cross-species transmission of these viruses. Further CoV surveillance studies in bats throughout the Americas are warranted.     

Body mass affects short-term heterothermy in Neotropical bats

Recent work in Australia and Africa has shown that heterothermy is widespread among phylogenetically diverse tropical and subtropical mammalian taxa. However, data on the use of heterothermy by Neotropical mammals are relatively scant, and those studies that exist focus on insect-eating bats. We investigated the capacity of fruit-eating Neotropical bats to use heterothermy when exposed to acute cold temperatures, and compared this to previous data focused on insect-eating bats sampled from the same region and time of year. We measured rectal temperatures prior to acute cold exposure (1 hr at an air temperature of 6, 7, or 10°C), and again after exposure. Our data show considerable variation in the thermoregulatory patterns of Neotropical bats, and generally, our results show that smaller bats cool quicker and to a greater extent than larger bats. Our results highlight the importance of energy conservation even in environments in which resources are relatively abundant.     

The role of frugivorous bats in tropical forest succession

Discussion of successional change has traditionally focused on plants. The role of animals in producing and responding to successional change has received far less attention. Dispersal of plant propagules by animals is a fundamental part of successional change in the tropics. Here we review the role played by frugivorous bats in successional change in tropical forests. We explore the similarities and differences of this ecological service provided by New and Old World seed-dispersing bats and conclude with a discussion of their current economic and conservation implications. Our review suggests that frugivorous New World phyllostomid bats play a more important role in early plant succession than their Old World pteropodid counterparts. We propose that phyllostomid bats have shared a long evolutionary history with small-seeded early successional shrubs and treelets while pteropodid bats are principally dispersers of the seeds of later successional canopy fruits. When species of figs (Ficus) are involved in the early stages of primary succession (e.g. in the river meander system in Amazonia and on Krakatau, Indonesia), both groups of bats are important contributors of propagules. Because they disperse and sometimes pollinate canopy trees, pteropodid bats have a considerable impact on the economic value of Old World tropical forests; phyllostomid bats appear to make a more modest direct contribution to the economic value of New World tropical forests. Nonetheless, because they critically influence forest regeneration, phyllostomid bats make an important indirect contribution to the economic value of these forests. Overall, fruit-eating bats play important roles in forest regeneration throughout the tropics, making their conservation highly desirable.     

蝙蝠抗病毒天然免疫的研究进展

近年来,健康的蝙蝠体内检测到了很多与人类传染病相关的病毒,包括狂犬病病毒(Rabies virus)、埃博拉病毒(Ebola virus)、严重急性呼吸综合征病毒(SARS-CoV)以及最近新出现的新型冠状病毒(SARS-CoV-2)等。与其他哺乳动物不同,蝙蝠在感染了这些病毒后不会表现出明显的临床症状。因此,人类可以通过研究蝙蝠的免疫系统获得抗病毒免疫的新知识。本文综述了蝙蝠抗病毒天然免疫研究的最新进展,指出了蝙蝠在天然免疫方面的特殊性:蝙蝠独有的飞行能力可能导致其演化出一套独特的抗病毒免疫响应机制,同时具有一套独特的机制限制炎症反应。蝙蝠物种的多样性丰富(超过1 400种),超过了哺乳动物的五分之一。因此对蝙蝠免疫基因的多样性研究,将促进对蝙蝠特殊免疫机制的理解,对人类传染病防治和畜牧业发展具有重要意义。     

Seed Dispersal in the Dark: Shedding Light on the Role of Fruit Bats in Africa

In spite of their recognized importance as seed dispersers in other parts of the tropics, seed dispersal by fruit bats has received scant research attention in Africa. To evaluate the role of African fruit bats in seed dispersal, we studied fruits and seeds below 480 bat feeding roosts in the East Usambara Mountains of Tanzania. We compared these findings to those reported in other African localities to place our results in a broader context. We found 49 plant species dispersed by bats: 28 species, 18 genera, and one family are novel reports of bat dispersal in Africa. Approximately 20 percent of the submontane tree flora of the East Usambaras is bat‐dispersed, including both widespread and endemic trees. African fruit bats are important seed dispersers at our study site because they move seeds of dozens of species tens or hundreds of meters, even seeds that are too large to ingest (greater than 5 mm in length). Fruit bats are likely important seed dispersers in other Afrotropical forests, as bats elsewhere in Africa are known to consume 20 genera and 16 species of plants reported here. Insights from studying remains under bat feeding roosts offer a simple method to further document and substantially increase our understanding of the role of African fruit bats in seed dispersal.     

Bats on a Budget: Torpor-Assisted Migration Saves Time and Energy

Bats and birds must balance time and energy budgets during migration. Migrating bats face similar physiological challenges to birds, but nocturnality creates special challenges for bats, such as a conflict between travelling and refueling, which many birds avoid by feeding in daylight and flying at night. As endothermic animals, bats and birds alike must expend substantial amounts of energy to maintain high body temperatures. For migratory birds refueling at stopovers, remaining euthermic during inactive periods reduces the net refuelling rate, thereby prolonging stopover duration and delaying subsequent movement. We hypothesized that bats could mitigate similar ambient-temperature dependent costs by using a torpor-assisted migration strategy. We studied silver-haired bats Lasionycteris noctivagans during autumn migration using a combination of respirometry and temperature-sensitive radiotelemetry to estimate energy costs incurred under ambient temperature conditions, and the energy that bats saved by using torpor during daytime roosting periods. All bats, regardless of sex, age, or body condition used torpor at stopover and saved up to 91% of the energy they would have expended to remain euthermic. Furthermore, bats modulated use of torpor depending on ambient temperature. By adjusting the time spent torpid, bats achieved a rate of energy expenditure independent of the ambient temperature encountered at stopover. By lowering body temperature during inactive periods, fuel stores are spared, reducing the need for refuelling. Optimal migration models consider trade-offs between time and energy. Heterothermy provides a physiological strategy that allows bats to conserve energy without paying a time penalty as they migrate. Although uncommon, some avian lineages are known to use heterothermy, and current theoretical models of migration may not be appropriate for these groups. We propose that thermoregulatory strategies should be an important consideration of future migration studies of both bats and birds.     

High activity enables life on a high-sugar diet: blood glucose regulation in nectar-feeding bats

High blood glucose levels caused by excessive sugar consumption are detrimental to mammalian health and life expectancy. Despite consuming vast quantities of sugar-rich floral nectar, nectar-feeding bats are long-lived, provoking the question of how they regulate blood glucose. We investigated blood glucose levels in nectar-feeding bats (Glossophaga soricina) in experiments in which we varied the amount of dietary sugar or flight time. Blood glucose levels increased with the quantity of glucose ingested and exceeded 25 mmol l(-1) blood in resting bats, which is among the highest values ever recorded in mammals fed sugar quantities similar to their natural diet. During normal feeding, blood glucose values decreased with increasing flight time, but only fell to expected values when bats spent 75 per cent of their time airborne. Either nectar-feeding bats have evolved mechanisms to avoid negative health effects of hyperglycaemia, or high activity is key to balancing blood glucose levels during foraging. We suggest that the coevolutionary specialization of bats towards a nectar diet was supported by the high activity and elevated metabolic rates of these bats. High activity may have conferred benefits to the bats in terms of behavioural interactions and foraging success, and is simultaneously likely to have increased their efficiency as plant pollinators.     

Some comments on the proposed perception of phase and nanosecond time disparities by echolocating bats

In a series of recent reports, Simmons and his colleagues propose that bats are able to accurately encode the spectral, temporal and phase information of their emitted calls and echoes. The information so encoded is then extracted by the networks of the auditory system with specialized processing. They propose that bats use this information to determine the distance to their target by crosscorrelating the entire structure of the emitted call with the structure of the echo. The idea is that slight deviations in the correlation function can be detected by the bat and the degree of mismatch provides an accurate measure of temporal disparity and hence range. The data in the reports purport to show that bats perceive the phase of ultrasonic signals and that they can resolve temporal disparities of about 10 ns, and thus can distinguish range differences as small as 2 m.The hypothesis also attempts to explain how a variety of acoustic cues are processed and represented in the auditory system and how they are combined to form a unitary percept of space and fine structure. The theory incorporates some time honored processes of extracting information, such as crosscorrelations. The implications of. the hypothesis, however, go far beyond a theory of neural processing and representation of information by ensembles of cells. The hypothesis requires some remarkable abilities, such as the phase coding of ultrasonic signals and a temporal acuity on the order of 10 ns. These features have never been seen in any neurophysiological study of any animal nor has its existence been implied in behavioral studies of other animals. If bats, in fact, detect and process those signals in the manner proposed by Simmons and his colleagues, it would suggest that bats are supermammals whose auditory systems have evolved new and extraordinary mechanisms not possessed by other animals.In view of the extraordinary implications of the hypothesis, it seems prudent to critically evaluate the data upon which the hypothesis is based. The purpose of this review is to point out a number of technical problems and deficiencies in those experiments which undermine the veracity of the purported demonstration of phase perception and nanosecond time resolution by bats.     

A comparison of bats and rodents as reservoirs of zoonotic viruses: are bats special?

Bats are the natural reservoirs of a number of high-impact viral zoonoses. We present a quantitative analysis to address the hypothesis that bats are unique in their propensity to host zoonotic viruses based on a comparison with rodents, another important host order. We found that bats indeed host more zoonotic viruses per species than rodents, and we identified life-history and ecological factors that promote zoonotic viral richness. More zoonotic viruses are hosted by species whose distributions overlap with a greater number of other species in the same taxonomic order (sympatry). Specifically in bats, there was evidence for increased zoonotic viral richness in species with smaller litters (one young), greater longevity and more litters per year. Furthermore, our results point to a new hypothesis to explain in part why bats host more zoonotic viruses per species: the stronger effect of sympatry in bats and more viruses shared between bat species suggests that interspecific transmission is more prevalent among bats than among rodents. Although bats host more zoonotic viruses per species, the total number of zoonotic viruses identified in bats (61) was lower than in rodents (68), a result of there being approximately twice the number of rodent species as bat species. Therefore, rodents should still be a serious concern as reservoirs of emerging viruses. These findings shed light on disease emergence and perpetuation mechanisms and may help lead to a predictive framework for identifying future emerging infectious virus reservoirs.     

Evaporative water loss is a plausible explanation for mortality of bats from white-nose syndrome

White-nose syndrome (WNS) has caused alarming declines of North American bat populations in the 5 years since its discovery. Affected bats appear to starve during hibernation, possibly because of disruption of normal cycles of torpor and arousal. The importance of hydration state and evaporative water loss (EWL) for influencing the duration of torpor bouts in hibernating mammals recently led to "the dehydration hypothesis," that cutaneous infection of the wing membranes of bats with the fungus Geomyces destructans causes dehydration which in turn, increases arousal frequency during hibernation. This hypothesis predicts that uninfected individuals of species most susceptible to WNS, like little brown bats (Myotis lucifugus), exhibit high rates of EWL compared to less susceptible species. We tested the feasibility of this prediction using data from the literature and new data quantifying EWL in Natterer's bats (Myotis nattereri), a species that is, like other European bats, sympatric with G. destructans but does not appear to suffer significant mortality from WNS. We found that little brown bats exhibited significantly higher rates of normothermic EWL than did other bat species for which comparable EWL data are available. We also found that Natterer's bats exhibited significantly lower rates of EWL, in both wet and dry air, compared with values predicted for little brown bats exposed to identical relative humidity (RH). We used a population model to show that the increase in EWL required to cause the pattern of mortality observed for WNS-affected little brown bats was small, equivalent to a solitary bat hibernating exposed to RH of ～95%, or clusters hibernating in ～87% RH, as opposed to typical near-saturation conditions. Both of these results suggest the dehydration hypothesis is plausible and worth pursuing as a possible explanation for mortality of bats from WNS.     

Vampire bats exhibit evolutionary reduction of bitter taste receptor genes common to other bats

The bitter taste serves as an important natural defence against the ingestion of poisonous foods and is thus believed to be indispensable in animals. However, vampire bats are obligate blood feeders that show a reduced behavioural response towards bitter-tasting compounds. To test whether bitter taste receptor genes (T2Rs) have been relaxed from selective constraint in vampire bats, we sampled all three vampire bat species and 11 non-vampire bats, and sequenced nine one-to-one orthologous T2Rs that are assumed to be functionally conserved in all bats. We generated 85 T2R sequences and found that vampire bats have a significantly greater percentage of pseudogenes than other bats. These results strongly suggest a relaxation of selective constraint and a reduction of bitter taste function in vampire bats. We also found that vampire bats retain many intact T2Rs, and that the taste signalling pathway gene Calhm1 remains complete and intact with strong functional constraint. These results suggest the presence of some bitter taste function in vampire bats, although it is not likely to play a major role in food selection. Together, our study suggests that the evolutionary reduction of bitter taste function in animals is more pervasive than previously believed, and highlights the importance of extra-oral functions of taste receptor genes.     

Survival of PIT-tagged lesser short-tailed bats (

Introduced mammalian predators are a major threat to New?Zealand?s wildlife, including bats. Controlling these predators using traps and poison baits can reduce their impact on bat populations. However, lesser short-tailed bats (Mystacina tuberculata) are potentially susceptible to toxins used for pest control in New?Zealand forests because of their broad diet and habit of feeding on the ground. Therefore, the risk of secondary poisoning should always be assessed before new toxins are used in areas inhabited by lesser short-tailed bats. We measured survivorship of a sample of lesser short-tailed bats monitored before, during and after deployment of the first-generation anticoagulant toxin pindone in the Eglinton Valley, Fiordland, during late winter and summer 2009?2010. Pindone-laced cereal baits were deployed in bait stations from September to late December 2009 in an effort to control rats (Rattus spp.). Communal roosts of the one lesser short-tailed bat colony in the valley are located entirely within or immediately adjacent to the area poisoned. Minimum number alive was determined for the sampled bat population after monitoring the occupancy of colonial roosts by individually PIT-tagged bats through the study period. Survivorship of bats was high throughout the monitoring period, with 319 of 322 bats (99%) recorded in the pre-monitoring period (August) known to be alive in October 2009 and 312 of 322 bats (97%) known to still be alive in January 2010. We conclude that lesser short-tailed bats did not consume pindone baits and that their survival was probably enhanced by rat control in the study area.