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.

The presence of ultrasonic harmonics in the calls of the rifleman (Acanthisitta chloris)

Ultrasonic vocalisations (frequencies > 20 kHz) have been extensively studied in the context of echolocation by bats and other mammals (Sales & Pye 1974; Wilson & Hare 2004). Ultrasonic calls have also been recorded from birds, including the blue-throated hummingbird ( Lampornis clemenciae ) (Pytte et al. 2004), where it was first thought that individuals made use of high pitch calls to avoid masking by background noise in a visually obscured environment. Similarly, city-dwelling great tits ( Parus major ) use song with a higher minimum frequency (although not ultrasonic) compared to woodland birds to communicate with conspecifics to avoid the predominantly low-frequency background noise in the city (Slabbekorn & Peet 2003). The theory that birds use ultrasound to avoid noise masking was discarded when it was discovered that there was no corresponding auditory brainstem response (i.e. sensory perception) to the ultrasonic calls in the hummingbirds producing those calls.

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.

Temporal and spatial patterns of seed dispersal of Musa acuminata by Cynopterus sphinx

The foraging behavior of greater short-nosed fruit bats (Cynopterus sphinx) on wild banana (Musa acuminata) and subsequent dispersal of seeds were studied in the Tropical Rainforest Conservation Area, Xishuangbanna Tropical Botanical Garden, Yunnan province, by direct observation of marked fruits, mist netting, and seed collection. The mean number (± SE) of individual C. sphinx captured by mist net were 2.2 ± 0.33/day and 1.4 ± 0.32/day in the rainy season (September to October) and dry season (November to December), respectively; the difference was not significant. The number of seed pellets expelled was 9.0 ± 1.12/day and 7.2 ± 1.37/day in the rainy and dry seasons respectively; again the difference was not significant. The removal curves for marked fruit were similar for 10 focal trees. Fruits were consumed heavily within two weeks after ripening and all the marked fruit were removed within one month. The difference in seed dispersal was significant between different feeding roosts indicating that patterns of seed dispersal may not be uniform. We found the seeds of M. acuminata can be dispersed by C. sphinx to a distance of about 200 m, and C. sphinx can be considered as an effective seed disperser of M. acuminata.

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.

Genetic divergence and echolocation call frequency in cryptic species of Hipposideros larvatus s.l. (Chiroptera : Hipposideridae) from the Indo-Malayan region

The intermediate leaf-nosed bat (Hipposideros larvatus) is a medium-sized bat distributed throughout the Indo-Malay region. In north-east India, bats identified as H. larvatus captured at a single cave emitted echolocation calls with a bimodal distribution of peak frequencies, around either 85 kHz or 98 kHz. Individuals echolocating at 85 kHz had larger ears and longer forearms than those echolocating at 98 kHz, although no differences were detected in either wing morphology or diet, suggesting limited resource partitioning. A comparison of mitochondrial control region haplotypes of the two phonic types with individuals sampled from across the Indo-Malay range supports the hypothesis that, in India, two cryptic species are present. The Indian 98-kHz phonic bats formed a monophyletic clade with bats from all other regional populations sampled, to the exclusion of the Indian 85-kHz bats. In India, the two forms showed 12–13% sequence divergence and we propose that the name Hipposideros khasiana for bats of the 85-kHz phonic type. Bats of the 98-kHz phonic type formed a monophyletic group with bats from Myanmar, and corresponded to Hipposideros grandis, which is suggested to be a species distinct from Hipposideros larvatus. Differences in echolocation call frequency among populations did not reflect phylogenetic relationships, indicating that call frequency is a poor indicator of evolutionary history. Instead, divergence in call frequency probably occurs in allopatry, possibly augmented by character displacement on secondary contact to facilitate intraspecific communication.

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.

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.

The effect of recording situation on the echolocation calls of the New Zealand Lesser short-tailed bat (Mystacina tuberculata Gray).

Using a broad‐band recording system (150 Hz‐100 kHz) the echolocation calls of the lesser short‐tailed bat (Mystacina tuberculata) were recorded under three very different situations: free‐flying, flying within a flight cage, and on release from the hand. Calls of bats landing and feeding on a platform in Wellington Zoo were also recorded. Both the lowest frequency and frequency of peak amplitude of calls were significantly affected by the situation under which calls were recorded. Although the calls of free‐flying bats are different from those produced by bats foraging on the ground, it is unlikely that M. tuberculata uses echolocation to locate prey on the ground. No significant differences could be found between the calls emitted by male and female bats, and no consistent relationships were obvious between temporal and spectral call characteristics. There was some evidence to suggest that individual bats could be identified by their echolocation calls.

Echolocation calls of the long tailed bat: a quantitative analysis of types of calls

The echolocation calls of long-tailed bats (Chalinolobus tuberculatus) were recorded in the Eglinton Valley, Fjordland, New Zealand, and digitized for analysis with the signal-processing software. Univariate and multivariate analyses of measure features facilitated a quantitative classification of the calls. Cluster analysis was used to categorize calls into two groups equating to search and terminal buzz calls described qualitatively for other species. When moving from search to terminal phases, the calls decrease in bandwidth, maximum and minimum frequency of call, and duration. Search calls begin with a steep-downward FM sweep followed by a short, less-modulated component. Buzz calls are FM sweeps. Although not found quantitatively, a broad pre-buzz group of calls also was identified. Ambiguity analysis of calls from the three groups shows that search-phrase calls are well suited to resolving the velocity of targets, and hence, identifying moving targets in a stationary clutter. Pre-buzz and buzz calls are better suited to resolving range, a feature that may aid the bats in capture of evasive prey after it has been identified.

Search-phase echolocation calls of the New Zealand lesser short-tailed bat (Mystacina tuberculata) and long-tailed bat (Chalinolobus tuberculatus).

This paper describes the search-phase echolocation calls of lesser short-tailed bats (Mystacina tuberculata) and long-tailed bats (Chalinolobus tuberculatus). Calls were recorded from all three subspecies of short-tailed bat and seven populations of long-tailed bat, three in Northland, two in the central North Island, and two in the lower South Island. The calls were recorded in the field and digitised, then three spectral components and one temporal component of the calls were measured. Calls of the lesser short-tailed bat could be loosely classified into subspecies by means of multivariate discriminant function analysis. Similarly, long-tailed bat calls showed regional variation, and discriminant function analysis was able to fit calls to regional groups with a high rate of success. The significance of the results presented is discussed in terms of the conservation of New Zealand bats and the unique ecology of the lesser short-tailed bat.

Acoustic identification of twelve 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.

Practical guidelines for planning and implementing fauna translocations

A successful translocation involves many complex factors, including a genetically appropriate source population that can sustain harvest, social and governmental support, assessment of disease transmission risk and a release site with appropriately secure habitat that can support population establishment and persistance. This information is typically discussed during staturory approval processes and can take considerable time. However, following approval, for translocations of most fauna, the initial critical step involves the inherently stressful process of capture, holding, transportation and release. This process is unpredictable and novel, and is especially challenging for wild animals when they are confined in close proximity to conspecifics and humans. In contrast, captive-reared animals have to cope with the unfamiliar challenges of finding food and shelter, along with coping with competition and predation. Little has been written in the scientific literature about the translocation process. This is unsurprising because this process has usually been the realm of skilled practioners, often with animal husbandry backgrounds, rather than research scientists. Highly skilled intuition, observation and the translocation practioner's equivalent of a 'green thumb' often guides the way. However, theory and experimentation, particularly on the effects of stress, is available and this work is invaluable for a successful translocation. Here, we provide a brief description of the translocation process, and discussion of what stress is and how it can be managed. We then provide practical guidelines for the successful translocation of invertebrates, lizards, turtles, passerine birds, marsupials and bats, using examples from Australia and New Zealand.

Adoption of alternative habitats by a threatened, "obligate" forest-dwelling bat in a fragmented landscape

While they are among the most ecologically important animals within forest ecosystems, little is known about how bats respond to habitat loss and fragmentation. The threatened lesser short-tailed bat (Mystacina tuberculata), considered to be an obligate deep-forest species, is one of only 2 extant land mammals endemic to New Zealand; it plays a number of important roles within native forests, including pollination and seed dispersal, and rarely occurs in modified forests. We used radiotelemetry to study the movements, roosting behavior, and habitat use of M. tuberculata within a fragmented landscape comprised of 3 main habitat types: open space (harvested forest and pastoral land), native forests, and exotic pine plantations. We found that the bats had smaller home-range areas and travelled shorter nightly distances than populations investigated previously from contiguous native forest. Furthermore, M. tuberculata occupied all 3 habitat types, with native forest being preferred overall. However, individual variation in habitat selection was high, with some bats preferring exotic plantation and open space over native forest. Roosting patterns were similar to those previously observed in contiguous forest; individual bats often switched between communal and solitary roosts. Our findings indicate that M. tuberculata exhibit some degree of behavioral plasticity that allows them to adapt to different landscape mosaics and exploit alternative habitats. To our knowledge, this is the first such documentation of plasticity in habitat use for a bat species believed to be an obligate forest-dweller.