Effects of clear-fell harvest on bat home range.

We investigated effects of roost loss due to clear-fell harvest on bat home range. The study took place in plantation forest, inhabited by the New Zealand long-tailed bat (Chalinolobus tuberculatus), in which trees are harvested between the ages 26-32 years. We determined home ranges by radiotracking different bats in areas that had and had not been recently clear-fell harvested. Home ranges were smaller in areas that had been harvested. Adult male bats selected 20-25 year old stands within home ranges before and after harvest. Males selected edges with open unplanted areas when harvest had not occurred but no longer selected these at proportions greater than their availability post harvest, probably because they were then readily available. This is the first radiotracking study to demonstrate a change in home range size and selection concomitant with felling of large areas of plantation forest, and thus quantify negative effects of forestry operations on this speciose group. The use of smaller home ranges post-harvest may reflect smaller colony sizes and lower roost availability, both of which may increase isolation of colonies and vulnerability to local extinction.

Bat colony size reduction coincides with clear-fell harvest operations and high rates of roost loss in plantation forest

Clear-fell harvest of forest concerns many wildlife biologists because of loss of vital resources such as roosts or nests, and effects on population viability. However, actual impact has not been quantified. Using New Zealand long-tailed bats (Chalinolobus tuberculatus) as a model species we investigated impacts of clear-fell logging on bats in plantation forest. C. tuberculatus roost within the oldest stands in plantation forest so it was likely roost availability would decrease as harvest operations occurred. We predicted that post-harvest: (1) roosting range sizes would be smaller, (2) fewer roosts would be used, and (3) colony size would be smaller. We captured and radiotracked C. tuberculatus to day-roosts in Kinleith Forest, an exotic plantation forest, over three southern hemisphere summers (Season 1 October 2006–March 2007; Season 2 November 2007–March 2008; and Season 3 November 2008–March 2009). Individual roosting ranges (100% MCPs) post harvest were smaller than those in areas that had not been harvested, and declined in area during the 3 years. Following harvest, bats used fewer roosts than those in areas that had not been harvested. Over 3 years 20.7% of known roosts were lost: 14.5% due to forestry operations and 6.2% due to natural tree fall. Median colony size was 4.0 bats (IQR = 2.0–8.0) and declined during the study, probably because of locally high levels of roost loss. Post harvest colonies were smaller than colonies in areas that had not been harvested. Together, these results suggest the impact of clear-fell harvest on long-tailed bat populations is negative.

Home range and habitat selection by a threatened bat in exotic plantation forest

Individuals' home ranges are constrained by resource distribution and density, population size, and energetic requirements. Consequently, home ranges and habitat selection may vary between individuals of different sex and reproductive conditions. Whilst home ranges of bats are well-studied in native habitats, they are often not well understood in modified landscapes, particularly exotic plantation forests. Although Chalinolobus tuberculatus (Vespertilionidae, Chiroptera) are present in plantation forests throughout New Zealand their home ranges have only been studied in native forest and forest-agricultural mosaic and no studies of habitat selection that included males had occurred in any habitat type. Therefore, we investigated C. tuberculatus home range and habitat selection within exotic plantation forest. Home range sizes did not differ between bats of different reproductive states. Bats selected home ranges with higher proportions of relatively old forest than was available. Males selected edges with open unplanted areas within their home ranges, which females avoided. We suggest males use these edges, highly profitable foraging areas with early evening peaks in invertebrate abundance, to maintain relatively low energetic demands. Females require longer periods of invertebrate activity to fulfil their needs so select older stands for foraging, where invertebrate activity is higher. These results highlight additional understanding gained when data are not pooled across sexes. Mitigation for harvest operations could include ensuring that areas suitable for foraging and roosting are located within a radius equal to the home range of this bat species.

Echolocation call production during aerial and terrestrial locomotion by New Zealand's enigmatic lesser short-tailed bat, Mystacina tuberculata

Linkage of echolocation call production with contraction of flight muscles has been suggested to reduce the energetic cost of flight with echolocation, such that the overall cost is approximately equal to that of flight alone. However, the pattern of call production with limb movement in terrestrially agile bats has never been investigated. We used synchronised high-speed video and audio recordings to determine patterns of association between echolocation call production and limb motion by Mystacina tuberculata Gray 1843 as individuals walked and flew, respectively. Results showed that there was no apparent linkage between call production and limb motion when bats walked. When in flight, two calls were produced per wingbeat, late in the downstroke and early in the upstroke. When bats walked, calls were produced at a higher rate, but at a slightly lower intensity, compared with bats in flight. These results suggest that M. tuberculata do not attempt to reduce the cost of terrestrial locomotion and call production through biomechanical linkage. They also suggest that the pattern of linkage seen when bats are in flight is not universal and that energetic savings cannot necessarily be explained by contraction of muscles associated with the downstroke alone.

The importance of exotic plantation forest for the New Zealand long-tailed bat (Chalinolobus tuberculatus)

Environmental certification schemes have stimulated increasing interest in biodiversity and its management within exotic plantation forests. These schemes expect management to be scientifically-based, even though little is known about how often, or which, native species use exotic plantation forests. Greater knowledge of the ecology of native species within exotic plantation forests is required to advise management and reduce risks to native species, particularly those that are rare, such as the New Zealand long-tailed bat (Chalinolobus tuberculatus). Long-tailed bats use exotic plantation forests throughout New Zealand but need protection from the impacts of forest management, and particularly clear-fell harvest, that is achievable only through a better understanding of their biology. The consequences of the current reduced re-planting, and the conversion of plantation forests into pasture resulting in smaller forested areas, should not be ignored because they may be associated with reductions in long-tailed bat populations. We review the current knowledge of long-tailed bats' use of exotic plantation forests, and report for the first time which exotic plantations long-tailed bats are known to use. We make recommendations for the design of monitoring programmes to detect long-tailed bats within plantation forests, and for research into the effects of forest management, especially logging, and comment on the likely impacts of reductions in forested areas on long-tailed bats.

Classification of echolocation calls from 14 species of bat by Support Vector Machines and Ensembles of Neural Networks

Calls from 14 species of bat were classified to genus and species using discriminant function analysis (DFA), support vector machines (SVM) and ensembles of neural networks (ENN). Both SVMs and ENNs outperformed DFA for every species while ENNs (mean identification rate – 97%) consistently outperformed SVMs (mean identification rate – 87%). Correct classification rates produced by the ENNs varied from 91% to 100%; calls from six species were correctly identified with 100% accuracy. Calls from the five species of Myotis, a genus whose species are considered difficult to distinguish acoustically, had correct identification rates that varied from 91 – 100%. Five parameters were most important for classifying calls correctly while seven others contributed little to classification performance.

Human vs. machine : identification of bat species from their echolocation calls by humans and by artificial neural networks

Automated remote ultrasound detectors allow large amounts of data on bat presence and activity to be collected. Processing of such data involves identifying bat species from their echolocation calls. Automated species identification has the potential to provide more consistent, predictable, and potentially higher levels of accuracy than identification by humans. In contrast, identification by humans permits flexibility and intelligence in identification, as well as the incorporation of features and patterns that may be difficult to quantify. We compared humans with artificial neural networks (ANNs) in their ability to classify short recordings of bat echolocation calls of variable signal to noise ratios; these sequences are typical of those obtained from remote automated recording systems that are often used in large-scale ecological studies. We presented 45 recordings (1–4 calls) produced by known species of bats to ANNs and to 26 human participants with 1 month to 23 years of experience in acoustic identification of bats. Humans correctly classified 86% of recordings to genus and 56% to species; ANNs correctly identified 92% and 62%, respectively. There was no significant difference between the performance of ANNs and that of humans, but ANNs performed better than about 75% of humans. There was little relationship between the experience of the human participants and their classification rate. However, humans with <1 year of experience performed worse than others. Currently, identification of bat echolocation calls by humans is suitable for ecological research, after careful consideration of biases. However, improvements to ANNs and the data that they are trained on may in future increase their performance to beyond those demonstrated by humans.

Fruit-feeding behaviour and use of olfactory cues by the fruit bat Rousettus leschenaulti: an experimental study

This paper describes the feeding behaviour ofRousettus leschenaulti Desmarest, 1820 on lychees, the preferred cultivated food of this bat in captive conditions. We found that feeding comprised 25–30% of the total activity of these animals in a flight cage and that feeding durations were not significantly different between two sexes. To evaluate the role of odor and vision in foraging behaviour, we provided animals with artificial lychees, real lychees and artificial lychees soaked in the juice of real lychees and we recorded the number of feeding approaches to the different “fruit” types. The results indicated that bats approached real fruit significantly more than artificial fruit, and that the number of approaches to the soaked artificial fruit was also significantly higher than to the unsoaked artificial fruit. There were no significant differences between sexes in approach rates to any “fruit” type. We discuss the role of different sensory cues in the foraging behaviour of these bats and emphasize that the olfactory cue is important in detecting food resources and discriminating between different kinds of food items.

Morphology, echolocation and foraging behaviour in two sympatric sibling species of bat (Tylonycteris pachypus and Tylonycteris robustula) (Chiroptera : Vespertilionidae)

We studied the wing morphology, echolocation calls, foraging behaviour and flight speed of Tylonycteris pachypus and Tylonycteris robustula in Longzhou County, South China during the summer (June–August) of 2005. The wingspan, wing loading and aspect ratio of the two species were relatively low, and those of T. pachypus were lower compared with T. robustula. The echolocation calls of T. pachypus and T. robustula consist of a broadband frequency modulated (FM) sweep followed by a short narrowband FM sweep. The dominant frequency of calls of T. pachypus was 65.1 kHz, whereas that of T. robustula was 57.7 kHz. The call frequencies (including highest frequency of the call, lowest frequency of the call and frequency of the call that contained most energy) of T. pachypus were higher than those of T. robustula, and the pulse duration of the former was longer than that of the latter. The inter-pulse interval and bandwidth of the calls were not significantly different between the two species. Tylonycteris pachypus foraged in more complex environments than T. robustula, although the two species were both netted in edge habitats (around trees or houses), along pathways and in the tops of trees. Tylonycteris pachypus flew slower (straight level flight speed, 4.3 m s−1) than T. robustula (straight level flight speed, 4.8 m s−1). We discuss the relationship between wing morphology, echolocation calls, foraging behaviour and flight speed, and demonstrate resource partitioning between these two species in terms of morphological and behavioural factors.

Echolocation calls, wing shape, diet and phylogenetic diagnosis of the endemic Chinese bat Myotis pequinius

We describe the echolocation calls, flight morphology and diet of the endemic Chinese bat Myotis pequinius Thomas, 1908. Orientation calls are broadband, and reach low terminal frequencies. Diet comprised 80% beetles by volume. Wing shape and call design suggest that the bats fly in cluttered habitats, and the possession of moderately long ears and the dietary composition imply they forage at least sometimes by gleaning. Myotis pequinius resembles a larger Oriental version of the western Palaearctic species M. nattereri. Phylogenetic analysis based on sequences of the cytochrome b gene of mitochondrial DNA (1,140 base pairs) from a range of Palaearctic Myotis species confirmed that M. pequinius is close to the nattereri group, and is a sister-species to the eastern Palaearctic M. bombinus. One bat sequenced from China could not be identified from available species descriptions. It was smaller than M. pequinius, and also differed from it in sequence divergence by 6.7%, suggesting the existence of additional, cryptic taxonomic diversity in this group. Our phylogenetic analysis also supports the recognition of M. schaubi as a species distinct from M. nattereri in Transcaucasia and south-western Asia. Myotis nattereri tschuliensis is more closely related to M. schaubi than to M. nattereri, and is best considered either as a subspecies of M. schaubi, or possibly as a distinct species.

Terrestrial locomotion of the New Zealand short-tailed bat Mystacina tuberculata and the common vampire bat Desmodus rotundus

Bats (Chiroptera) are generally awkward crawlers, but the common vampire bat (Desmodus rotundus) and the New Zealand short-tailed bat (Mystacina tuberculata) have independently evolved the ability to manoeuvre well on the ground. In this study we describe the kinematics of locomotion in both species, and the kinetics of locomotion in M. tuberculata. We sought to determine whether these bats move terrestrially the way other quadrupeds do, or whether they possess altogether different patterns of movement on the ground than are observed in quadrupeds that do not fly. Using high-speed video analyses of bats moving on a treadmill, we observed that both species possess symmetrical lateral-sequence gaits similar to the kinematically defined walks of a broad range of tetrapods. At high speeds, D. rotundus use an asymmetrical bounding gait that appears to converge on the bounding gaits of small terrestrial mammals, but with the roles of the forelimbs and hindlimbs reversed. This gait was not performed by M. tuberculata. Many animals that possess a single kinematic gait shift with increasing speed from a kinetic walk (where kinetic and potential energy of the centre of mass oscillate out of phase from each other) to a kinetic run (where they oscillate in phase). To determine whether the single kinematic gait of M. tuberculata meets the kinetic definition of a walk, a run, or a gait that functions as a walk at low speed and a run at high speed, we used force plates and high-speed video recordings to characterize the energetics of the centre of mass in that species. Although oscillations in kinetic and potential energy were of similar magnitudes, M. tuberculata did not use pendulum-like exchanges of energy between them to the extent that many other quadrupedal animals do, and did not transition from a kinetic walk to kinetic run with increasing speed. The gait of M. tuberculata is kinematically a walk, but kinetically run-like at all speeds.

Echolocation call intensity in the aerial hawking bat Eptesicus bottae (Vespertilionidae) studied using stereo videogrammetry

Aerial hawking bats use intense echolocation calls to search for insect prey. Their calls have evolved into the most intense airborne animal vocalisations. Yet our knowledge about call intensities in the field is restricted to a small number of species. We describe a novel stereo videogrammetry method used to study flight and echolocation behaviour, and to measure call source levels of the aerial hawking bat Eptesicus bottae (Vespertilionidae). Bats flew close to their predicted minimum power speed. Source level increased with call duration; the loudest call of E. bottae was at 133 dB peSPL. The calculated maximum detection distance for large flying objects (e.g. large prey, conspecifics) was up to 21 m. The corresponding maximum echo delay is almost exactly the duration of one wing beat in E. bottae and this also is its preferred pulse interval. These results, obtained by using videogrammetry to track bats in the field, corroborate earlier findings from other species from acoustic tracking methods.

Effects of different surfaces on the perception of prey-generated noise by the Indian false vampire bat Megaderma lyra.

The low- and high-frequency components of a rustling sound, created when prey (freshly killed frog) was jerkily pulled on dry and wet sandy floors and asbestos, were recorded and played back to individual Indian false vampire bats (Megaderma lyra). Megaderma lyra responded with flight toward the speakers and captured dead frogs, that were kept as reward. The spectral peaks were at 8.6, 7.1 and 6.8 kHz for the low-frequency components of the sounds created at the dry, asbestos and wet floors, respectively. The spectral peaks for the high-frequency sounds created on the respective floors were at 36.8,27.2 and 23.3 kHz. The sound from the dry floor was more intense than that of from the other two substrata. Prey movements that generated sonic or ultrasonic sounds were both sufficient and necessary for the bats to detect and capture prey. The number of successful prey captures was significantly greater for the dry floor sound, especially to its high-frequency components. Bat-responses were low to the wet floor and moderate to the asbestos floor sounds. The bats did not respond to the sound of unrecorded parts of the tape. Even though the bats flew toward the speakers when the prey generated sounds were played back and captured the dead frogs we cannot rule out the possibility of M. lyra using echolocation to localize prey. However, the study indicates that prey that move on dry sandy floor are more vulnerable to predation by M. lyra.

Acoustic identification of 12 species of echolocating bat by discriminant function analysis and artificial neural networks

We recorded echolocation calls from 14 sympatric species of bat in Britain. Once digitised, one temporal and four spectral features were measured from each call. The frequency-time course of each call was approximated by fitting eight mathematical functions, and the goodness of fit, represented by the mean-squared error, was calculated. Measurements were taken using an automated process that extracted a single call from background noise and measured all variables without intervention. Two species of Rhinolophus were easily identified from call duration and spectral measurements. For the remaining 12 species, discriminant function analysis and multilayer back-propagation perceptrons were used to classify calls to species level. Analyses were carried out with and without the inclusion of curve-fitting data to evaluate its usefulness in distinguishing among species. Discriminant function analysis achieved an overall correct classification rate of 79% with curve-fitting data included, while an artificial neural network achieved 87%. The removal of curve-fitting data improved the performance of the discriminant function analysis by 2 %, while the performance of a perceptron decreased by 2 %. However, an increase in correct identification rates when curve-fitting information was included was not found for all species. The use of a hierarchical classification system, whereby calls were first classified to genus level and then to species level, had little effect on correct classification rates by discriminant function analysis but did improve rates achieved by perceptrons. This is the first published study to use artificial neural networks to classify the echolocation calls of bats to species level. Our findings are discussed in terms of recent advances in recording and analysis technologies, and are related to factors causing convergence and divergence of echolocation call design in bats.