Doppler detection triggers instantaneous escape behavior in scanning bats.

Animals must instantaneously escape from predators for survival, which requires quick detection of approaching threats. Although the neural mechanisms underlying the perception of looming objects have been extensively studied in the visual system, little is known about their auditory counterparts. Echolocating bats use their auditory senses to perceive not only the soundscape, but also the physical environment through active sensing. Although object movement induces both echo delay changes and Doppler shifts, the actual information required to perceive movement has been unclear. Herein, we addressed this question by playing back phantom echoes mimicking an approaching target to horseshoe bats and found that they relied only on Doppler shifts. This suggests that the bats do not perceive object motion in the spatiotemporal dimension (i.e., positional variation), as in vision, but rather take advantage of acoustic sensing by directly detecting velocity, thereby enabling them to respond instantaneously to approaching threats.

A glimpse into the foraging and movement behaviour of Nyctalus aviator; a complementary study by acoustic recording and GPS tracking.

Species of open-space bats that are relatively large, such as bats from the genus Nyctalus, are considered as high-risk species for collisions with wind turbines (WTs). However, important information on their behaviour and movement ecology, such as the locations and altitudes at which they forage, is still fragmentary, while crucial for their conservation in light of the increasing threat posed by progressing WT construction. We adopted two different methods of microphone array recordings and GPS-tracking capturing data from different spatio-temporal scales in order to gain a complementary understanding of the echolocation and movement ecology of Nyctalus aviator, the largest open-space bat in Japan. Based on microphone array recordings, we found that echolocation calls during natural foraging are adapted for fast flight in open-space optimal for aerial-hawking. In addition, we attached a GPS tag that can simultaneously monitor feeding buzz occurrence, and confirmed that foraging occurred at 300 m altitude and that the flight altitude in mountainous areas is consistent with the turbine conflict zone, suggesting that the birdlike noctule is a high-risk species in Japan. Further investigations on this species could provide valuable insights into their foraging and movement ecology, facilitating the development of a risk assessment regarding WTs.

Analysis of echolocation behavior of bats in "echo space" using acoustic simulation.

BACKGROUND: Echolocating bats use echo information to perceive space, control their behavior, and adjust flight navigation strategies in various environments. However, the echolocation behavior of bats, including echo information, has not been thoroughly investigated as it is technically difficult to measure all the echoes that reach the bats during flight, even with the conventional telemetry microphones currently in use. Therefore, we attempted to reproduce the echoes received at the location of bats during flight by combining acoustic simulation and behavioral experiments with acoustic measurements. By using acoustic simulation, echoes can be reproduced as temporal waveforms (including diffracted waves and multiple reflections), and detailed echo analysis is possible even in complex obstacle environments. RESULTS: We visualized the spatiotemporal changes in the echo incidence points detected by bats during flight, which enabled us to investigate the "echo space" revealed through echolocation for the first time. We then hypothesized that by observing the differences in the "echo space" before and after spatial learning, the bats' attentional position would change. To test this hypothesis, we examined how the distribution of visualized echoes concentrated at the obstacle edges after the bats became more familiar with their environment. The echo incidence points appeared near the edge even when the pulse direction was not toward the edge. Furthermore, it was found that the echo direction correlated with the turn rate of the bat's flight path, revealing for the first time the relationship between the echo direction and the bat's flight path. CONCLUSIONS: We were able to clarify for the first time how echoes space affects echolocation behavior in bats by combining acoustic simulations and behavioral experiments.

Discriminating predation attempt outcomes during natural foraging using the post-buzz pause in the Japanese large-footed bat, Myotis macrodactylus.

Bats emit a series of echolocation calls with an increasing repetition rate (the terminal buzz) when attempting to capture prey. This is often used as an acoustic indicator of prey-capture attempts. However, because it is directly linked to foraging efficiency, predation success is a more useful measure than predation attempts in ecological research. The characteristics of echolocation calls that consistently signify predation success across different situations have not been identified. Owing to additional influencing factors, identification of these characteristics is particularly challenging for wild bats foraging in their natural environment compared with those in flight chambers. This study documented the natural foraging behavior of wild Japanese large-footed bats (Myotis macrodactylus) using synchronized acoustic and video recordings. From the video recordings, we could assign 137 attacks to three outcome categories: prey captured (51.8%), prey dropped (29.2%) and failed attempt (19%). Based on previous indications from laboratory studies that the length of the silent interval following the terminal buzz (post-buzz pause) might reflect the prey-capture outcome, we compared post-buzz pause durations among categories of attack outcomes. The post-buzz pause was longest in the case of successful capture, suggesting that the length of the post-buzz pause is a useful acoustic indicator of predation success during natural foraging in M. macrodactylus. Our finding will advance the study of bat foraging behavior using acoustic data, including estimations of foraging efficiency and analyses of feeding habitat quality.

Dietary analysis reveals differences in the prey use of two sympatric bat species.

One mechanism for morphologically similar and sympatric species to avoid competition and facilitate coexistence is to feed on different prey items within different microhabitats. In the current study, we investigated and compared the diet of the two most common and similar-sized bat species in Japan-Murina ussuriensis (Ognev, 1913) and Myotis ikonnikovi (Ognev, 1912)-to gain more knowledge about the degree of overlap in their diet and their foraging behavior. We found that both bat species consumed prey from the orders of Lepidoptera and Diptera most frequently, while the proportion of Dipterans was higher in the diet of M. ikonnikovi. Furthermore, we found a higher prey diversity in the diet of M. ikonnikovi compared to that of M. ussuriensis that might indicate that the former is a more generalist predator than the latter. In contrast, the diet of M. ussuriensis contained many Lepidopteran families. The higher probability of prey items likely captured via gleaning to occur in the diet of M. ussuriensis in contrast to M. ikonnikovi indicates that M. ussuriensis might switch between aerial-hawking and gleaning modes of foraging behavior. We encourage further studies across various types of habitats and seasons to investigate the flexibility of the diet composition and foraging behavior of these two bat species.

Modulation of acoustic navigation behaviour by spatial learning in the echolocating bat Rhinolophus ferrumequinum nippon.

Using echolocation, bats receive acoustic information on their surroundings, which is assumed to help them sophisticatedly navigate complex environments. In this study, to understand spatial learning and acoustic sensing in bats, we investigated how flight and echolocation control changed in Rhinolophus ferrumequinum nippon as they learnt about their surroundings in an obstacle course that they flew through repeatedly. In these experiments, two testing environments (acoustically permeable and acoustically reflective) were prepared using chains and acrylic boards as obstacles to evaluate the interactive effects of spatial learning and flight environments. We found that bats reduced the meandering width of their flights and pulse emissions, and also seemed to reduce their shifts in pulse direction as they learnt more about their environments in both conditions. Throughout all our experiments, the bats with slower flight speeds tended to emit more pulses, which suggests that the number of pulse emissions reflects the echolocation tactics of each bat. The maximum flight speed was especially increased in the acoustically permeable condition, with frequent emissions of multiple pulses (≧triplets) in the early stages of flight, suggesting that bats adjust their flight plan based on how much of their surroundings they are able to sense in advance.

The effect of local land use and loss of forests on bats and nocturnal insects.

Land-use intensification at local and landscape level poses a serious threat to biodiversity and affects species interactions and ecosystem function. It is thus important to understand how interrelated taxa respond to land-use intensification and to consider the importance of different spatial scales. We investigated whether and how local land-use intensity and landscape features affect the predator-prey interaction of bats and insects. Bats and nocturnal insects were assessed on 50 grassland sites in the Schorfheide-Chorin. We analyzed the effect of local land use and distance to forested areas as a proxy for site accessibility on bats and insects and their biological interaction measured in bat's feeding activity. Insect abundance increased with higher land-use intensity, while size and diversity of insects decreased. In contrast, bat activity, diversity, and species composition were determined by the distance to forested areas and only slightly by land-use intensity. Feeding attempts of bats increased with higher insect abundance and diversity but decreased with insect size and distance to forested areas. Finally, our results revealed that near forested areas, the number of feeding attempts was much lower on grassland sites with high, compared to those with low land-use intensity. In contrast, far from forests, the feeding attempts did not differ significantly between intensively and extensively managed grassland sites. We conclude that the two interrelated taxa, bats and insects, respond to land-use intensification on very different scales. While insects respond to local land use, bats are rather influenced by surrounding landscape matrix. Hereby, proximity to forests reveals to be a prerequisite for higher bat species diversity and a higher rate of feeding attempts within the area. However, proximity to forest is not sufficient to compensate local high land-use intensity. Thus, local land-use intensification in combination with a loss of forest remnants weakens the interaction of bats and insects.

The Importance of Landscape Elements for Bat Activity and Species Richness in Agricultural Areas.

Landscape heterogeneity is regarded as a key factor for maintaining biodiversity and ecosystem function in production landscapes. We investigated whether grassland sites at close vicinity to forested areas are more frequently used by bats. Considering that bats are important consumers of herbivorous insects, including agricultural pest, this is important for sustainable land management. Bat activity and species richness were assessed using repeated monitoring from May to September in 2010 with acoustic monitoring surveys on 50 grassland sites in the Biosphere Reserve Schorfheide-Chorin (North-East Germany). Using spatial analysis (GIS), we measured the closest distance of each grassland site to potentially connecting landscape elements (e.g., trees, linear vegetation, groves, running and standing water). In addition, we assessed the distance to and the percent land cover of forest remnants and urban areas in a 200 m buffer around the recording sites to address differences in the local landscape setting. Species richness and bat activity increased significantly with higher forest land cover in the 200 m buffer and at smaller distance to forested areas. Moreover, species richness increased in proximity to tree groves. Larger amount of forest land cover and smaller distance to forest also resulted in a higher activity of bats on grassland sites in the beginning of the year during May, June and July. Landscape elements near grassland sites also influenced species composition of bats and species richness of functional groups (open, edge and narrow space foragers). Our results highlight the importance of forested areas, and suggest that agricultural grasslands that are closer to forest remnants might be better buffered against outbreaks of agricultural pest insects due to higher species richness and higher bat activity. Furthermore, our data reveals that even for highly mobile species such as bats, a very dense network of connecting elements within the landscape is beneficial to promote activity in open areas and thus assure vital ecosystem function in agricultural landscapes.