Temporal and spatial distribution and habitat associations of an urban population of New Zealand long-tailed bats (Chalinolobus tuberculatus)

Two ultrasound survey methods were used to determine the presence and activity patterns of New Zealand long-tailed bats (Chalinolobus tuberculatus) in the city of Hamilton. First, 13 monthly surveys conducted at 18 green spaces found C. tuberculatus in only one urban forest reserve, Hammond Bush, where they were found consistently throughout the year. Bat activity was strongly related to temperature. Second, twice-yearly citywide surveys conducted over 2 years determined the distribution and habitat associations of C. tuberculatus. Bats were found only in the southern part of the city and were strongly associated with the Waikato River. Bat activity was negatively correlated with housing and street light density and positively correlated with topographical complexity. In Hamilton, topographical complexity indicates the presence of gullies. Gullies probably provide foraging and roosting opportunities and connect the river to distant forest patches. These results suggest that urban habitats can be useful for bats if gullies can link these to distant habitat fragments.

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.

Evidence of homing following translocation of long‐tailed bats (Chalinolobus tuberculatus) at Grand Canyon Cave, New Zealand

Homing behaviour in the New Zealand long-tailed bat (Chalinolobus tuberculatus), a temperate insectivorous species, was investigated at Grand Canyon Cave, central North Island. A pilot study of nine adult male bats was conducted to determine whether use of the cave was regular enough for a homing study. Eight bats returned to the cave over the 3 week monitoring period, six on the night of the following release. Nine additional bats carrying radio transmitters were then released at three sites (three at each site) c.5, 10 and 20km due east of the border of, and outside the population's known familiar area respectively. All but one of these nine was subsequently detected at the cave. Results suggest that adult long-tailed bats are able to return home following displacement both inside and outside their familiar area. Implications of these findings for translocations of bats and the possessions of a potential long distance navigation system by this species are discussed.

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.

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.

Sex-specific roost selection by bats in clearfell harvested plantation forest : improved knowledge advises management

To maximize energetic savings, female bats often roost communally whilst pregnant or with non-volant dependents, whereas male bats more often roost alone; however, differences in selection of roosts by sex have not often been investigated. Better understanding of female colony locations could focus management to protect the majority of bats. New Zealand's long-tailed bat (Chalinolobus tuberculatus) roost in exotic plantation forest, where sex-specific roost selection has not been investigated, and therefore such management is not possible. We investigated sex-specific roost selection by long-tailed bats for the first time. Roosts and paired nonroosts were characterized testing predictions that males and females select roosts that differ from non-roosts, and males and females select different roosts. Females and males chose Pinus radiata roosts that differed from non-roost trees. Results suggest each sex chose roosts that maximized energetic savings. Female bats used roosts closer to water sources, that warmed earlier in the day, which allowed maintenance of high temperatures. Males appeared to choose roosts that allowed torpor use for long periods of the day. Males may be less selective with their roost locations than females, as they roosted further from water sources. This could allow persistence of male bats in marginal habitat. As all female long-tailed bats chose roosts within 150 m of waterways, management to protect bats could be focused here. To protect bats least able to escape when roosts are harvested, harvest of forest stands selected by female bats as roost sites should be planned when bats are not heavily pregnant nor have non-volant dependents.

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.

Long-tailed bats' use of a Pinus radiata stand in Kinleith Forest : Recommendations for monitoring

Targeted monitoring of threatened species within plantations is becoming more important due to forest certification programmes’ requirement to consider protection of threatened species, and to increase knowledge of the distribution of species. To determine patterns of long-tailed bat (Chalinolobus tuberculatus) activity in different habitat structures, with the aim of improving the likelihood of detection by targeting monitoring, we monitored one stand of 26 year-old Pinus radiata over seven months between December 2007 and June 2008 in Kinleith Forest, an exotic plantation forest centred around Tokoroa, South Waikato, New Zealand. Activity was determined by acoustic recording equipment, which is able to detect and record bats’ echolocation calls. We monitored activity from sunset to sunrise along a road through the stand, along stand edges, and in the interior of the stand. Bats were recorded on 80% of the 35 nights monitored. All activity throughout the monitoring period was detected on the edge of the stand or along the road. No bats were detected within the interior of the stand. Bat activity was highest along the road through the stand (40.4% of all passes), followed by an edge with stream running alongside (35.2%), along the road within a skidsite (19.8%), and along an edge without a stream (4.6%). There was a significant positive relationship between bat pass rate (bat passes h-1) and the feeding buzz rate (feeding buzzes h-1) indicating that bat activity was associated with feeding and not just commuting. Bat feeding activity was also highest along the road through the stand (59.2% of feeding buzzes), followed by the road within the skidsite (30.6%), and along the stream-side edge (10.2%). No feeding buzzes were recorded in either the interior or along the edge without the stream. Differences in overall feeding activity were significant only between the road and edge and between edges with and without a stream. Bat activity was detected each month and always by the second night of monitoring, and in this stand was highest during April. We recommend targeted monitoring for long-tailed bats be focused on road-side and stand edge habitat, and along streams, and that monitoring take place for at least three nights to maximise probability of detection.

Identification of New Zealand bats in flight from analysis of echolocation calls by artificial neural networks.

Time-expanded and heterodyned echolocation calls of the New Zealand long-tailed Chalinolobus tuberculatus and lesser short-tailed bat Mystacina tuberculata were recorded and digitally analysed. Temporal and spectral parameters were measured from time-expanded calls and power spectra generated for both time-expanded and heterodyned calls. Artificial neural networks were trained to classify the calls of both species using temporal and spectral parameters and power spectra as input data. Networks were then tested using data not previously seen. Calls could be unambiguously identified using parameters and power spectra from time-expanded calls. A neural network, trained and tested using power spectra of calls from both species recorded using a heterodyne detector set to 40 kHz (the frequency with the most energy of the fundamental of C. tuberculatus call), could identify 99% and 84% of calls of C. tuberculatus and M. tuberculata, respectively. A second network, trained and tested using power spectra of calls from both species recorded using a heterodyne detector set to 27 kHz (the frequency with the most energy of the fundamental of M. tuberculata call), could identify 34% and 100% of calls of C. tuberculatus and M. tuberculata, respectively. This study represents the first use of neural networks for the identification of bats from their echolocation calls. It is also the first study to use power spectra of time-expanded and heterodyned calls for identification of chiropteran species. The ability of neural networks to identify bats from their echolocation calls is discussed, as is the ecology of both species in relation to the design of 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.

The conservation status of New Zealand bats, 2009

The New Zealand Threat Classification System (NZTCS) is a national system used to assess the risk of extinction faced by New Zealand plants, animals and fungi. The system is specifically designed to be relevant to New Zealand's unusual ecological and geographic conditions. We undertook a re-evaluation of the status of seven bat taxa based on our knowledge of New Zealand bats using revised NZTCS criteria. Five taxa were listed as Threatened or At Risk: one as Nationally Critical (long-tailed bat Chalinolobus tuberculatus ‘South Island’), one as Nationally Endangered (southern lesser short-tailed bat Mystacina tuberculata tuberculata), two as Nationally Vulnerable (long-tailed bat ‘North Island’ and northern lesser short-tailed bat M. t. aupourica) and one as Declining (central lesser short-tailed bat M. t. rhyacobia). One taxon was assessed as Data Deficient (greater short-tailed bat M. robusta) and one (little red flying fox Pteropus scapulatus) as Vagrant. We suspect declines result primarily from predation and competition from introduced mammals, habitat degradation, and disturbance.

Applying network analysis to the conservation of habitat trees in urban environments: a case study from Brisbane, Australia

In Australia more than 300 vertebrates, including 43 insectivorous bat species, depend on hollows in habitat trees for shelter, with many species using a network of multiple trees as roosts, We used roost-switching data on white-striped freetail bats (Tadarida australis; Microchiroptera: Molossidae) to construct a network representation of day roosts in suburban Brisbane, Australia. Bats were caught from a communal roost tree with a roosting group of several hundred individuals and released with transmitters. Each roost used by the bats represented a node in the network, and the movements of bats between roosts formed the links between nodes. Despite differences in gender and reproductive stages, the bats exhibited the same behavior throughout three radiotelemetry periods and over 500 bat days of radio tracking: each roosted in separate roosts, switched roosts very infrequently, and associated with other bats only at the communal roost This network resembled a scale-free network in which the distribution of the number of links from each roost followed a power law. Despite being spread over a large geographic area (> 200 km(2)), each roost was connected to others by less than three links. One roost (the hub or communal roost) defined the architecture of the network because it had the most links. That the network showed scale-free properties has profound implications for the management of the habitat trees of this roosting group. Scale-free networks provide high tolerance against stochastic events such as random roost removals but are susceptible to the selective removal of hub nodes. Network analysis is a useful tool for understanding the structural organization of habitat tree usage and allows the informed judgment of the relative importance of individual trees and hence the derivation of appropriate management decisions, Conservation planners and managers should emphasize the differential importance of habitat trees and think of them as being analogous to vital service centers in human societies.

Roost tree characteristics determine use by the white-striped freetail bat (Tadarida australis, Chiroptera : Molossidae) in suburban subtropical Brisbane, Australia

We examined factors affecting roost tree selection by the white-striped freetail bat Tadarida australis (Chiroptera: Molossidae), a large insectivorous bat in suburban Brisbane, Australia. We compared biophysical characteristics associated with 34 roost trees and 170 control trees of similar diameter, height and tree senescence characters. Roost trees used by the white-striped freetail bat had significantly higher numbers of hollows in the trunk and branches (P < 0.003) and were more likely to contain a large trunk cavity with an internal diameter of > 30 cm (P < 0.001) than control trees. These trees also accommodated more species of hollow-using fauna (P = 0.005). When comparing roost trees with control trees of similar diameters and heights, roost trees were on average at a later stage of tree senescence (P < 0.001). None of the roost trees were found in the large forest reserves fringing the Brisbane metropolitan area despite these areas being used for foraging by the white-striped freetail bat. Although all tree locations in this study were in modified landscapes, roost trees tended to be surrounded by groups of trees and undergrowth. Roost trees provide important habitat requirements for hollow-using fauna in suburban, rural and forested environments.

Palaeoecology of the Mount Etna bat fauna, coastal Eastern Queensland

Global warming is already threatening many animal and plant communities worldwide, however, the effect of climate change on bat populations is poorly known. Understanding the factors influencing the survival of bats is crucial to their conservation, and this cannot be achieved solely by modern ecological studies. Palaeoecological investigations provide a perspective over a much longer temporal scale, allowing the understanding of the dynamic patterns that shaped the distribution of modern taxa. In this study twelve microchiropteran fossil assemblages from Mount Etna, central-eastern Queensland, ranging in age from more than 500,000 years to the present day, were investigated. The aim was to assess the responses of insectivorous bats to Quaternary environmental changes, including climatic fluctuations and recent anthropogenic impacts. In particular, this investigation focussed on the effects of increasing late Pleistocene aridity, the subsequent retraction of rainforest habitat, and the impact of cave mining following European settlement at Mount Etna. A thorough examination of the dental morphology of all available extant Australian bat taxa was conducted in order to identify the fossil taxa prior to their analysis in term of species richness and composition. This detailed odontological work provided new diagnostic dental characters for eighteen species and one genus. It also provided additional useful dental characters for three species and seven genera. This odontological analysis allowed the identification of fifteen fossil bat taxa from the Mount Etna deposits, all being representatives of extant bats, and included ten taxa identified to the species level (i.e., Macroderma gigas, Hipposideros semoni, Rhinolophus megaphyllus, Miniopterus schreibersii, Miniopterus australis, Scoteanax rueppellii, Chalinolobus gouldii, Chalinolobus dwyeri, Chalinolobus nigrogriseus and Vespadelus troughtoni) and five taxa identified to the generic level (i.e., Mormopterus, Taphozous, Nyctophilus, Scotorepens and Vespadelus). Palaeoecological analysis of the fossil taxa revealed that, unlike the non-volant mammal taxa, bats have remained essentially stable in terms of species diversity and community membership between the mid-Pleistocene rainforest habitat and the mesic habitat that occurs today in the region. The single major exception is Hipposideros semoni, which went locally extinct at Mount Etna. Additionally, while intensive mining operations resulted in the abandonment of at least one cave that served as a maternity roost in the recent past, the diversity of the Mount Etna bat fauna has not declined since European colonisation. The overall resilience through time of the bat species discussed herein is perhaps due to their unique ecological, behavioural, and physiological characteristics as well as their ability to fly, which have allowed them to successfully adapt to their changing environment. This study highlights the importance of palaeoecological analyses as a tool to gain an understanding of how bats have responded to environmental change in the past and provides valuable information for the conservation of threatened modern species, such as H. semoni.