Acoustic estimation of wildlife abundance: methodology for vocal mammals in forested habitats

Habitat loss and hunting pressure threaten mammal populations worldwide, generating critical time constraints on trend assessment. This study introduces a new survey method that samples continuously and non‐invasively over long time periods, obtaining estimates of abundance from vocalization rates. We present feasibility assessment methods for acoustic surveys and develop equations for estimating population size. As an illustration, we demonstrate the feasibility of acoustic surveys for African forest elephants (Loxodonta africana cyclotis). Visual surveys and vocalizations from a forest clearing in the Central African Republic were used to establish that low‐frequency elephant calling rate is a useful index of elephant numbers (linear regression P < 0.001, r_adj.² = 0.58). The effective sampling area was 3.22 km² per acoustic sensor, a dramatic increase in coverage over dung survey transects. These results support the use of acoustic surveys for estimating elephant abundance over large remote areas and in diverse habitats, using a distributed network of acoustic sensors. The abundance estimation methods presented can be applied in surveys of any species for which an acoustic abundance index and detection function have been established. This acoustic survey technique provides an opportunity to improve management and conservation of many acoustically‐active taxa whose populations are currently under‐monitored.     

Toward reliable population density estimates of partially marked populations using spatially explicit mark–resight methods

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Optimizing ultrasonic sampling effort for monitoring forest bats

Landscape‐scale monitoring is a key approach for assessing changes in indicators. However, great care needs to be taken to collect rigorous data and avoid wasting resources in long‐term programmes. Insect‐eating bats are diverse, functionally important and are often proposed as indicator species of environmental health. We used acoustic (ultrasonic) data from pilot bat surveys undertaken in forests and woodlands to optimize sampling effort to produce precise estimates of bat activity and occupancy. We also carried out simulations to evaluate the statistical power of different sampling designs to detect changes in activity and occupancy levels of individual bat species. There was little gain in precision for estimates of bat activity by sampling beyond five to six detector nights. To ensure spatial heterogeneity was sampled around a monitoring point, three detectors for two nights or two detectors for three nights would be required. This level of sampling was also sufficient to be 90% certain of recording occupancy for 11 of 12 taxa. Power simulations revealed that a sampling design using two detectors per monitoring point for two nights could detect a 30% decline within 10 years with 90% power for all species, except the white‐striped free tail bat (Tadarida australis), using either changes in activity levels or occupancy. However, fewer years were required when using occupancy. Setting detectors either on‐flyways or off‐flyways contributed only minor differences to the time taken to reach 90% power for both occupancy and activity levels, though sampling both locations has major implications for interpreting trends in bats. We suggest that bat activity levels are more sensitive for detecting change than occupancy because one pass or 1000 passes can be recorded per night by an acoustic detector, and this is not differentiated by occupancy. Bats can be monitored cost‐effectively and should be included in monitoring programmes.     

Large Roads Reduce Bat Activity across Multiple Species

Although the negative impacts of roads on many terrestrial vertebrate and bird populations are well documented, there have been few studies of the road ecology of bats. To examine the effects of large roads on bat populations, we used acoustic recorders to survey bat activity along ten 300 m transects bordering three large highways in northern California, applying a newly developed statistical classifier to identify recorded calls to the species level. Nightly counts of bat passes were analyzed with generalized linear mixed models to determine the relationship between bat activity and distance from a road. Total bat activity recorded at points adjacent to roads was found to be approximately one-half the level observed at 300 m. Statistically significant road effects were also found for the Brazilian free-tailed bat (Tadarida brasiliensis), big brown bat (Eptesicus fuscus), hoary bat (Lasiurus cinereus), and silver-haired bat (Lasionycteris noctivagans). The road effect was found to be temperature dependent, with hot days both increasing total activity at night and reducing the difference between activity levels near and far from roads. These results suggest that the environmental impacts of road construction may include degradation of bat habitat and that mitigation activities for this habitat loss may be necessary to protect bat populations.     

Effects of survey methods on burrowing owl behaviors

Monitoring wildlife populations often involves intensive survey efforts to attain reliable estimates of population size. Such efforts can increase disturbance to animals, alter detection, and bias population estimates. Burrowing owls (Athene cunicularia) are declining across western North America, and information on the relative effects of potential survey methods on owl behaviors is needed. We designed a field experiment to compare burrowing owl flight distances, times displaced, and probabilities of being displaced between 4 potential population survey methods (single walking surveyor, single vehicle stop, single vehicle stop with 2 surveyors, and double vehicle stop with 2 surveyors), and an experimental control in the agricultural matrix of Imperial Valley, California. Between 25 April and 1 May 2008, we randomly applied survey methods to 395 adult male owls during daylight hours (0700 hours through 1900 hours). All survey methods increased odds of displacing owls from perches. Survey methods with observers outside the vehicle were 3 times more likely to displace an owl than a single vehicle stop where observers remained inside the vehicle. Owls were displaced farther distances by all survey methods compared to control trials, but distances and time displaced did not differ among survey methods. We recommend that surveys for counting owls during the breeding season in agroecystems like the Imperial Valley where high densities of owls nest primarily along the borders of fields be conducted using single vehicle stops with or without 2 surveyors, depending on conditions for locating owls from roads. © 2011 The Wildlife Society.     

Habitat associations of bats in a working rangeland landscape

Land‐use change has resulted in rangeland loss and degradation globally. These changes include conversion of native grasslands for row‐crop agriculture as well as degradation of remaining rangeland due to fragmentation and changing disturbance regimes. Understanding how these and other factors influence wildlife use of rangelands is important for conservation and management of wildlife populations. We investigated bat habitat associations in a working rangeland in southeastern North Dakota. We used Petterson d500x acoustic detectors to systematically sample bat activity across the study area on a 1‐km point grid. We identified calls using Sonobat autoclassification software. We detected five species using this working rangeland, which included Lasionycteris noctivagans (2,722 detections), Lasiurus cinereus (2,055 detections), Eptesicus fuscus (749 detections), Lasiurus borealis (62 detections), and Myotis lucifugus (1 detection). We developed generalized linear mixed‐effects models for the four most frequently detected species based on their ecology. The activity of three bat species increased with higher tree cover. While the scale of selection varied between the four species, all three investigated scales were explanatory for at least one bat species. The broad importance of trees to bats in rangelands may put their conservation needs at odds with those of obligate grassland species. Focusing rangeland bat conservation on areas that were treed prior to European settlement, such as riparian forests, can provide important areas for bat conservation while minimizing negative impacts on grassland species.     

Estimating animal population density using passive acoustics

Reliable estimation of the size or density of wild animal populations is very important for effective wildlife management, conservation and ecology. Currently, the most widely used methods for obtaining such estimates involve either sighting animals from transect lines or some form of capture‐recapture on marked or uniquely identifiable individuals. However, many species are difficult to sight, and cannot be easily marked or recaptured. Some of these species produce readily identifiable sounds, providing an opportunity to use passive acoustic data to estimate animal density. In addition, even for species for which other visually based methods are feasible, passive acoustic methods offer the potential for greater detection ranges in some environments (e.g. underwater or in dense forest), and hence potentially better precision. Automated data collection means that surveys can take place at times and in places where it would be too expensive or dangerous to send human observers. Here, we present an overview of animal density estimation using passive acoustic data, a relatively new and fast‐developing field. We review the types of data and methodological approaches currently available to researchers and we provide a framework for acoustics‐based density estimation, illustrated with examples from real‐world case studies. We mention moving sensor platforms (e.g. towed acoustics), but then focus on methods involving sensors at fixed locations, particularly hydrophones to survey marine mammals, as acoustic‐based density estimation research to date has been concentrated in this area. Primary among these are methods based on distance sampling and spatially explicit capture‐recapture. The methods are also applicable to other aquatic and terrestrial sound‐producing taxa. We conclude that, despite being in its infancy, density estimation based on passive acoustic data likely will become an important method for surveying a number of diverse taxa, such as sea mammals, fish, birds, amphibians, and insects, especially in situations where inferences are required over long periods of time. There is considerable work ahead, with several potentially fruitful research areas, including the development of (i) hardware and software for data acquisition, (ii) efficient, calibrated, automated detection and classification systems, and (iii) statistical approaches optimized for this application. Further, survey design will need to be developed, and research is needed on the acoustic behaviour of target species. Fundamental research on vocalization rates and group sizes, and the relation between these and other factors such as season or behaviour state, is critical. Evaluation of the methods under known density scenarios will be important for empirically validating the approaches presented here.     

A comparison of two bat detectors: which is most likely to detect New Zealand’s Chalinolobus tuberculatus?

ABSTRACT

The presence of bat species is commonly determined by placing acoustic bat detectors that record bat echolocation calls in the habitat they are likely to use. Detection rates are affected by variables including type of detection unit used. We compared detection rates of long-tailed bat (Chalinolobus tuberculatus) echolocation calls between two types of automated bat detectors: Wildlife Acoustics SMZC Zero Crossing Bat Recorders (ZC), and Frequency Compression Automated Bat Monitoring units (FC) produced by New Zealand’s Department of Conservation. Units were placed in locations where bats were known to be present, but not all detected bats. The median number of bat passes recorded by FC units over 10 nights was 20 compared with a median of 3 bat passes for ZC units. ZC units also detected bats over significantly fewer nights than FC units. These results suggest FC units are more sensitive and therefore better to use where long-tailed bats are expected to be at low abundance or only present infrequently. Because of inconsistencies in detection rates, we recommend the use of only one model of the detector within a monitoring project. Our data also suggests that surveys should take place over long periods to maximise likelihood of detecting bats, if present.     

Seasonal patterns of habitat use by insectivorous bats in a subtropical African agro‐ecosystem dominated by macadamia orchards

We report on acoustic surveys of insectivorous bats conducted during seven months of the year using ANABAT recordings in two habitats (macadamia orchards and adjacent riparian bush) in a subtropical agro‐ecosystem in northern South Africa. We defined two functional foraging groups of bats based on their echolocation calls: (i) open‐air foragers (family Molossidae) having narrow‐band, low‐frequency, low duty cycle calls; and (ii) clutter‐edge foragers (families Miniopteridae and Vespertilionidae), having broad‐band, higher frequency, low duty cycle calls. Bat activity (number of bat passes) was not significantly influenced by habitat. Total bat activity and activity of both functional groups varied significantly between seasons, being highest in summer and autumn (coinciding with annual peaks in numbers of Twin spotted (Bathycoelia natalicola) and Green (Nezara spp) Stinkbugs, order Heteroptera, family Pentatomidae, and Macadamia Nut Borer moths, Cryptophlebia ombrodelta) and lower in winter and spring. No significant effect of moon phase was detected, either on total activity or activity of the two functional groups. We postulate that the significant pattern of seasonality of commuting and/or foraging activity of bats in macadamia orchards (which is more marked in open‐air foragers) may be driven by the seasonal abundance of pest insects such as stinkbugs and Macadamia Nut Borer moths.     

Habitat-associated and temporal patterns of bat activity in a diverse forest landscape of southern New England, USA

The development and use of acoustic recording technology, surveys have revealed the composition, relative levels of activity, and preliminary habitat use of bat communities of various forest locations. However, detailed examinations of acoustic surveys results to investigate temporal patterns of bat activity are rare. Initial active acoustic surveys of bat activity on the Quabbin Reservoir watershed in central Massachusetts recorded high numbers of call sequences by five bat species. The results showed the importance of aquatic and open habitats for bats in this diverse forest landscape. Because the preliminary surveys were restricted in extent (habitats), number (replicates), and duration (active surveys only, limited number of repeated surveys), the authors strongly recommended more comprehensive and extensive surveys of bat activity on the area. This paper reports the results of active (manual) and passive (automated) acoustic bat activity surveys in three replicates of ten habitats, conducted three to four times per year over the course of the study between 2004 and 2006. Over the full study, 2,800 actively-recorded 37,632 passively-recorded call sequences were examined and identified to species when possible. General patterns in activity did not differ by habitat between the two survey methods. Call sequences were recorded more often in open, especially aquatic habitats than in cluttered habitats. The use of passive surveys allowed for the assessment of temporal patterns of activity and an analysis of the relationship between ambient temperature and activity.     

Acoustic characterization of bats from Malta: setting a baseline for monitoring and conservation of bat populations

Bioacoustic research has made several advancements in developing systems to record extensive acoustic data and classify bat echolocation calls to species level using automated classifiers. These systems are useful as echolocation calls give valuable information on bat behaviour and ecology and hence are widely used for research and conservation of bat populations. Despite the challenges associated with automated classifiers, due to the interspecific differences in call characteristics of bat species found in the Maltese Islands, the use of a quantitative and automated approach is investigated. The sound analysis pipeline involved the use of an algorithm to clean sound files from background noise and measure temporal and spectral parameters of bat echolocation calls. These parameters were then fed to a trained and validated artificial neural network using a bat call library built from reference bat calls from Malta. The automatic classifier achieved an overall correct classification rate of 98%. This high correct classification rate for reliable species identification may have benefitted from the absence of typically problematic species, such as species in the genus Myotis, in the analyses. This study’s results pave the way for efficient and reliable bat acoustic surveys in Malta in aid of necessary monitoring and conservation by providing an updated bat species list and their echolocation characteristics.     

Comparing two fish sampling standards over time: largely congruent results but with caveats

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Survival of Greater Sage‐Grouse broods: survey method affects disturbance and age‐specific detection probability

Investigators rely on brood surveys to estimate annual fecundity of game birds. However, investigators often do not account for factors that influence brood detection probability nor rarely document how much females and their broods are disturbed (flush rates) during surveys, which could lead to biased survival estimates. We used 45 radio‐tagged female Greater Sage‐Grouse (Centrocercus urophasianus) with broods to compare detection probabilities and document disturbance among four survey methods to allow future investigators to select the method that best meets their objectives. These methods included daytime flush, daytime visual, nocturnal spotlight, and fecal surveys at nocturnal roost sites, with the latter being a novel method. We used Cormack–Jolly–Seber (CJS) models to compare detection probability and daily survival estimates for visual and fecal surveys of broods 0–47 d post‐hatch and a double‐survey approach to compare detection probabilities among flush, fecal, and spotlight surveys ~42 d post‐hatch when investigators often determine brood fate. From CJS models, detection probability for visual surveys increased with brood age (0.618–0.881), whereas detection probability for fecal surveys did not (0.748). Daily survival probability estimates increased with brood age and differed annually based on fecal surveys (2016: 0.978–1.000 and 2017: 0.839–0.998). We detected age‐specific daily survival probability with visual surveys (0.956–0.997), but not annual differences. Based on the double‐survey approach, detection probability was high (0.857–1.000) for all methods. We flushed ~310–750% fewer females and broods during fecal and spotlight surveys than during both types of daytime surveys. Our results highlight the need to account for detection probabilities among methods and document disturbance to hens and broods that can help investigators design surveys to minimize impacts to birds. Furthermore, our result suggest that actions to improve brood survival during the first week post‐hatch may improve local recruitment.     

Seasonal patterns of habitat selection in the insectivorous bat‐eared fox

Understanding how animals utilize their habitat provides insights about their ecological needs and is of importance for both theoretical and applied ecology. As changing seasons impact prey habitat selection and vegetation itself, it is important to understand how seasonality impacts microhabitat choice in optimal foragers and their prey. We followed habituated bat‐eared foxes (Otocyon megalotis) in the Kalahari, South Africa, to study their seasonal habitat selection patterns and relate them to the habitat preferences of their main prey, termites (Hodotermes mossambicus). We used Resource Selection Functions (RSFs) to study bat‐eared foxes’ 3rd‐ and 4th‐order habitat selection by comparing used locations to random ones within their home ranges. Third‐order habitat selection for habitat type and composition was weak and varied little between seasons. We found that patterns of fox habitat selection did not mirror habitat selection of Hodotermes (quantified using RSFs), even when feeding on them (4th‐order). Taken together, these results might indicate that bat‐eared foxes’ food resources are homogenously distributed across habitats and that prey other than Hodotermes play an important role in bat‐eared foxes’ space use.     

Improved Campephiline detection: An experiment conducted with the Magellanic Woodpecker

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Heard but not seen: Comparing bat assemblages and study methods in a mosaic landscape in the Western Ghats of India

We used capture (mist‐netting) and acoustic methods to compare the species richness, abundance, and composition of a bat assemblage in different habitats in the Western Ghats of India. In the tropics, catching bats has been more commonly used as a survey method than acoustic recordings. In our study, acoustic methods based on recording echolocation calls detected greater bat activity and more species than mist‐netting. However, some species were detected more frequently or exclusively by capture. Ideally, the two methods should be used together to compensate for the biases in each. Using combined capture and acoustic data, we found that protected forests, forest fragments, and shade coffee plantations hosted similar and diverse species assemblages, although some species were recorded more frequently in protected forests. Tea plantations contained very few species from the overall bat assemblage. In riparian habitats, a strip of forested habitat on the river bank improved the habitat for bats compared to rivers with tea planted up to each bank. Our results show that shade coffee plantations are better bat habitat than tea plantations in biodiversity hotspots. However, if tea is to be the dominant land use, forest fragments and riparian corridors can improve the landscape considerably for bats. We encourage coffee growers to retain traditional plantations with mature native trees, rather than reverting to sun grown coffee or coffee shaded by a few species of timber trees.     

Potential of bat pass duration measures for studies of bat activity

Acoustic detectors have become increasingly used by bat workers to investigate bat ecology and assess the impacts of anthropogenic pressures. Within these studies, the metric used, ‘bat activity’, is based on the number of bat passes, without considering the bat pass duration (i.e. each event of a bat detected within the range of an ultrasonic detector). We expected that bat pass duration may contain information about site quality in terms of foraging potential. Because bats are expected to have a more sinuous trajectory and slower velocity when they exhibit foraging behaviour, as opposed to commuting behaviour, we hypothesize a longer bat pass duration in favourable habitats; during seasons with important energetic demands; or during night peak activity. We used datasets from a large-scale acoustic bat survey (n = 2890 sites), with a total of 24,597 bat pass measures from 6 taxa, and performed GLMM modelling. We detected a significant effect of habitat type on bat pass duration for five taxa. Shorter bat pass durations were detected at the beginning of the night. We detected longer pass durations during the lactation period or just before hibernating, while weather conditions or ageing and wear of the detector rarely influenced bat pass duration. Bat pass duration appears to be a simple and easy measure for position calls on a gradient between commuting vs. foraging behaviour. We suggest that the traditional measure of bat activity may be weighted by bat pass duration by giving more weight to events with potentially stronger links to foraging behaviour.     

Systematics of the Pacific monkey‐faced bats (Chiroptera: Pteropodidae), with a new species of Pteralopex and a new Fijian genus

The fruit‐bat genus Pteralopex comprises the monkey‐faced bats, a group of six endangered species found only in old‐growth forests on certain islands in the south‐west Pacific (the Solomon Islands and Fiji). The taxonomy of the genus is reviewed in detail and updated accordingly. Two ‘cryptic’ biological species are shown to occur in sympatry on both Bougainville and Choiseul in the northern Solomon Islands (corresponding to Pteralopex anceps Andersen, 1909 and a previously undescribed species) and each is accordingly described and reviewed. A new genus (Mirimiri) is erected for the Fijian monkey‐faced bat (formerly Pteralopex acrodonta), which differs greatly both morphologically and genetically from species of Pteralopex in the Solomon Islands. Ecomorphological differences between sympatric Pteralopex species are briefly reviewed, including potential differences in functional morphology and feeding ecology. Geographic patterns of occurrence and future survey priorities for monkey‐faced bats are also discussed.     

The urban matrix and artificial light restricts the nightly ranging behaviour of Gould's long‐eared bat (Nyctophilus gouldi)

Cities are heterogeneous landscapes, with remnant vegetation interspersed amongst areas designed for human use. Native wildlife remaining in urban areas are only likely to thrive and persist if they incorporate human altered areas into what they perceive as habitat. Many sensitive species may be lost if they are restricted to remnant vegetation, and cannot use the urban matrix. In this study, we quantify spatial aspects of the ranging behaviour of Gould's long‐eared bat (Nyctophilus gouldi) using radio‐telemetry and acoustic surveys to determine use of the suburban–bushland interface. This species represents a group prone to extinction due to biological attributes that adapt it to flight within cluttered vegetation, making it more specialized. We radio‐tracked 19 individuals in Cumberland State Forest (CSF), a 40‐ha remnant located in north‐west Sydney, New South Wales, Australia. The nightly range (95% Kernel Density Estimator) was small and localized, and was no greater than 80 ha, where individuals moved on average less than 300 m from roosts each night. All individual N. gouldi tracked used bushland in an almost obligate manner, where up to 100% of used habitat was within local bushland, with little to no use of areas classified as ‘urban’ (residential, commercial and educational land uses). Small open spaces exposed to artificial lighting within the main ranging area of CSF had significantly lower activity (bat passes) of Nyctophilus spp. and significantly higher activity of other species more tolerant of urbanization (P < 0.05). Our results demonstrate that artificial lighting can ‘spill‐over’ into bushland and alter the use of preferred habitat. We conclude that large patches (>40 ha) of protected remnant vegetation must be managed to reduce further degradation, and smaller isolated patches could be restored to provide habitat, particularly in narrow bushland corridors, to assist these species to tolerate urban areas.