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.     

Small-scale dung survey reveals high forest elephant density and preference for mixed species forest in an intact protected area

Understanding the ecological factors influencing African forest elephant (Loxodonta cyclotis) abundance and distribution is crucial for their conservation management in Central Africa. Dung surveys have been conducted at the landscape scale and confirmed the overwhelming impact of anthropogenic activities on forest elephants. We present results from a small-scale survey in a pristine protected area without anthropogenic activities to elucidate the ecological factors influencing forest elephant density. We conducted a line transect dung survey in a small study area (110 km²) around Mbeli Bai, a natural forest clearing in the Nouabalé-Ndoki National Park, Republic of Congo, and compare results with a landscape survey conducted during the same period. We used habitat specific dung decay data collected on site to estimate elephant density using distance sampling. We fitted Generalized Additive Models to elephant dung encounter rate using explanatory variables collected during the transect survey and from geospatial data. The small-scale survey revealed a precise estimate of forest elephant density that was twice as high as the result from a landscape survey with higher density in mixed species forest for the small-scale survey. We could not find an impact of the proximity of forest clearings and proximity to rivers at the small scale. Fine-scale habitat features, e.g. degree of canopy and understorey closure, had little explanatory power for elephant dung encounter rate. Small-scale dung surveys are a useful method to reveal spatio-temporal variation in forest elephant density and distribution which can inform conservation practitioners in a timely manner. Combining monitoring methods at various spatial scales improves our knowledge and conservation efforts of forest elephants. Nouabalé-Ndoki National Park is a stronghold for forest elephants and of global importance for their conservation.

     

Elephants caught in the middle: impacts of war, fences and people on elephant distribution and abundance in the Caprivi Strip, Namibia

We conducted wet [26 March–4 April 2003 (Apr03)] and dry [1–8 November 2005 (Nov05)] season aerial surveys of African elephants ( Loxodonta africana Blumenbach) in the Caprivi Strip, Namibia to provide an updated status report on elephant numbers and distribution and assist with a historical analysis of elephant distribution and abundance in the Caprivi Strip. During the wet season when water was available in seasonal pans, elephants were widely distributed throughout the survey area. In contrast, during the dry season, a majority of elephant herds occurred within 30 km of the perennial Kwando, Linyanti and Okavango rivers and few herds occurred within the West Caprivi Game Reserve where water in the seasonal pans was limited. We estimated 5318 elephants for the 7731-km² survey area (0.71 elephants km⁻²) for the Apr03 wet season survey and 6474 elephants for the 8597-km² survey area (0.75 elephants km⁻²) for the Nov05 dry season survey. Based on our aerial surveys and reports of elephant numbers and distribution from historical aerial surveys and telemetry studies, civil war, veterinary fences and human activities appear to have effected changes in African elephant abundance, distribution and movements in the Caprivi Strip, Namibia since 1988 when the first comprehensive aerial surveys were conducted.     

Dung survey bias and elephant population estimates in southern Mozambique

We used dung surveys to estimate population size and extracted an age structure from boli diameters for the elephants living in the Maputo Elephant Reserve. Our estimate was based on published defecation rates, dung decay rates, distance-sampling techniques and 1,672 dung piles encountered on 204 line-transects. The reserve had at least 311 (95% CI: 198–490) elephants at a density of 0.60 (95% CI: 0.38–0.94) per km². However, observer bias reduced effective strip widths and inflated estimates and their confidence limits. The age structure extrapolated from dung measurements indicated few newborn calves compared with other populations. To detect population changes of 2–5% at 80% power, dung surveys should be carried out every second year for the next 20 years using 100 transects of at least 500 m each. Comparison with a 1995 dung survey suggests that the population is stable and that previous fears of a major population decline during the civil war have no foundation.     

Estimating population sizes for elusive animals: the forest elephants of Kakum National Park,Ghana

African forest elephants are difficult to observe in the dense vegetation, and previous studies have relied upon indirect methods to estimate population sizes. Using multilocus genotyping of noninvasively collected samples, we performed a genetic survey of the forest elephant population at Kakum National Park, Ghana. We estimated population size, sex ratio and genetic variability from our data, then combined this information with field observations to divide the population into age groups. Our population size estimate was very close to that obtained using dung counts, the most commonly used indirect method of estimating the population sizes of forest elephant populations. As their habitat is fragmented by expanding human populations, management will be increasingly important to the persistence of forest elephant populations. The data that can be obtained from noninvasively collected samples will help managers plan for the conservation of this keystone species.     

Trends in populations of elephant and other large herbivores in Gonarezhou National Park,Zimbabwe, as revealed by sample aerial surveys

For 30 years, regular aerial surveys in Zimbabwean protected areas were funded, designed and executed primarily to estimate elephant numbers. Other large herbivores were recorded, even though some species were not easily seen from the air in savannah woodlands. Population estimates for species other than elephant provided indices of abundance that could be used to determine temporal trends in population size. This study tests for significant trends in the abundance of large herbivores in Gonarezhou National Park, assuming that data from aerial sample surveys designed for elephant also provide accurate estimates of real trends in the populations of other herbivores. For each species, the exponential rate of population change was calculated using weighted regression, with the variance of this rate based on the sampling variances of the population estimates. Significant population trends were detected for eight species. Before the 1992 drought, elephant number was held approximately constant by frequent culls, but afterwards, it increased at a mean annual rate of 6.2% (confidence limits 4.0% and 8.6%). Elephants in cow herds increased at 7.3%, significantly faster than elephants in bull herds (?0.5%). Buffalo, eland, kudu, nyala, waterbuck, wildebeest and zebra all increased in number, after population declines during the drought.     

Application of photographic capture–recapture modelling to estimate demographic parameters for male Asian elephants

In addition to the threats of habitat loss and degradation, adult males of the Asian elephant Elephas maximus also face greater threats from ivory poaching and conflict with humans. To understand the impact of these threats, conservationists need robust estimates of abundance and vital rates specifically for the adult male segment of elephant populations. By integrating the identification of individual male elephants in a population from distinct morphology and natural markings, with modern capture–recapture (CR) sampling designs, it is possible to estimate various demographic parameters that are otherwise difficult to obtain from this long-lived and wide-ranging megaherbivore. In this study, we developed systematic individual identification protocols and integrated them into CR sampling designs to obtain capture histories and thereby estimate the abundance of adult bull elephants in a globally important population in southern India. We validated these estimates against those obtained from an independent method combining line-transect density estimates with age–sex composition data for elephants. The sampled population was open to gains and losses between sampling occasions. The abundance of adult males in the 176 km² study area was (SÊ ) = 134(14.20) and they comprised 14% (±1%) of the total elephant population. Time-specific abundance estimates for each sampling occasion showed a distinct increase in adult male numbers over the sampling period, explained by seasonal patterns of local migration. CR-based estimates for adult male abundance closely matched estimates from distance-based methods. Thus, while providing abundance data of comparable rigour and precision, photographic CR methods permit estimation of demographic parameters for the Asian elephant that are both urgently needed and difficult to obtain.     

Estimating forest elephant density in Sapo National Park (Liberia) with a rainfall model

The number of elephant dung‐piles lying on the forest floor is a function of the number of elephants present and the rainfall in the 2 preceding months. We present the results of a stochastic model that describes this relationship and we show how it can be used to estimate elephant numbers. The data from a survey in Sapo NP (Liberia) in 1989 are used as an example. The dung‐pile density was estimated at 152 km^?2 with confidence interval from 72 to 322, and the number of elephants was estimated to be 313 with confidence interval from 172 to 617.     

How does a re-colonizing population of Asian elephants affect the forest habitat?

The Asian elephant Elephas maximus is currently re-colonizing the Bardia National Park in lowland Nepal. We studied their impact on woody vegetation in the nutrient-rich floodplain and in the relatively nutrient-poor sal forest. The types and extent of tree impact were recorded along fixed-width transects (335 km). Species composition, density and size classes ≥8 cm diameter breast height (dbh) were recorded in 15-m radius random plots ( n =95). Impact was higher in the floodplain complex than in the sal-dominated forest. Our hypothesis that elephants were more selective on species in the nutrient-poor sal forest was only partly supported; the niche breadth of impacted trees was slightly higher in the floodplain complex. Pushed-over trees accounted for the highest proportion of impact (55%), followed by killed trees (39%). Of the pushed trees, 10% were not used for food. Among food trees, elephants selectively impacted size class 12–16 cm dbh, whereas non-food trees were impacted independently of size. A large proportion of the freshly browsed trees had been felled previously, indicating that most felled trees survived, enabling elephants to feed on them again. This may reflect an evolutionary adaptation among long-lived species with high site fidelity. Owing to preferential use but low abundance, two species in sal forest, Grewia spp. and Desmodium oojeinense , were found to be particularly vulnerable to local extinction due to elephants. Although the elephants had impacted a large number of species (62, 73% of all), 56.4% of the impacted trees consisted of Mallotus phillippinensis . A recently observed increase in the density of M. phillippinensis and the concurrent reduction of the hardly utilized Shorea robusta indicates that the rapidly growing elephant population may modify the composition of the forest by increasing its preferred food species.     

Estimating abundance from detection–nondetection data for randomly distributed or aggregated elusive populations

Estimating abundance is important in many ecological studies in order to understand the spatial and temporal dynamics of a population, which can assist in management and conservation. However, direct estimates of abundance can be difficult and expensive to obtain, particularly for wide-ranging, rare or elusive species. An alternative – estimating from detection-nondetection data – is a challenging but alluring concept to ecologists since the cost and effort of a study can be greatly reduced. This paper describes a method for estimating the abundance of randomly distributed or aggregated populations by using binary data where the probability of detection is less than one. The performances of the models were evaluated by computer simulations comprising 1620 cases. The results show that the accuracy of the abundance estimates increases as the sampling rate, efficiency of survey method, and the number of repeated surveys increase, whereas the accuracy declines as individuals become more aggregated. For a randomly distributed population, using a sampling rate of 0.05 in a survey method with a detection probability of 0.5, and repeating surveys three times provides sufficient accuracy of abundance. For an aggregated population, to achieve reasonably accurate abundance estimates the sampling rate should be doubled and each cell should be repeatedly surveyed on 4 to 6 occasions.     

Abundance and distribution of the endangered loggerhead turtle in Spanish Mediterranean waters and the conservation implications

During 2 years (2001–2003), we performed seasonal aerial surveys in the central Spanish Mediterranean following the transect line methodology in order to determine the abundance and distribution patterns of loggerhead turtles Caretta caretta . We surveyed a total of 16 700 km, accounting for 770 turtle sightings. Loggerhead turtles were present with high abundance all year round. No seasonal differences in abundance were found, except in spring 2001, where the density of turtles was higher than in the other seasons. Our results show that the Western Mediterranean is not a 'summer' feeding area as proposed previously, as a high number of turtles are present throughout the year. The average surface density of turtles in the whole study area was 0.21 turtles km⁻² [95% confidence interval (CI): 0.17–0.25], and the mean abundance was 6653 turtles (95% CI: 5514–8027). The data relate to the number of turtles on the surface only, as diving turtles escape observation. Correcting our estimations of diving behaviour data in the area, the absolute abundance was 18 954 turtles (95% CI: 6679–53 786). Bearing in mind that around 25 000 loggerheads are caught per year in the Spanish Mediterranean, our results indicate that accidental captures seem to be a significant threat for this species, and conservation measures have to be implemented to avoid a non-sustainable situation.     

Effect of count duration on abundance estimates of Black-capped Vireos

ABSTRACT.   Distance sampling applied to point count surveys (point transects) has become a common method for estimating the absolute abundance of birds. When conducting point transects, detections of focal species are typically recorded during a fixed time interval. However, count duration has varied among studies and the effect of such variation on the resulting abundance estimates is unclear. My objective was to examine the effect of count duration on abundance estimates of male Black-capped Vireos ( Vireo atricapilla ). The abundance of these vireos in a 349-ha area in central Texas was estimated using 3-, 5-, and 6-min point transects and results were then compared to actual number present as determined by banding and territory mapping. The 3-min counts provided an estimate that was 26% greater than the actual number of male Vireos present ( N = 201), but this number was within the corresponding 95% confidence interval ( N = 157–413). Confidence intervals for the 5- and 6-min counts did not include the actual number of vireos present. The shortest count duration may have provided the most accurate abundance estimate because male Black-capped Vireos are typically active, sing intermittently, and sometimes move tens of meters between songs. Thus, shorter-duration counts may also yield the most accurate abundance estimates for other species that exhibit similar behavior. However, because behavior varies among species, I recommend that investigators collect preliminary data to establish an appropriate count duration when accurate estimates of absolute, rather than relative, abundance are important.     

How reliable are dung counts for estimating elephant numbers?

Dung counts are the most commonly used techniques for estimating elephant numbers in forests, yet there is considerable scepticism concerning their accuracy. Published accounts of dung counts show that they give estimates similar to those from other methods for vertebrates ranging in size from lizards to elephants. For ungulates, macropods and elephants there are strong correlations between estimates from dung counts and other methods. Thus, dung counts are as accurate or inaccurate as other methods for estimating vertebrate numbers, including elephants. Dung counts for elephants give estimates that are as precise as, and sometimes more precise than, those from aerial surveys of elephants. This is because the variance in dung density is usually low and results in a lower than expected variance for the final elephant estimate when combined with the variances of defecation and decay rates. Dung counts may be more appropriate than aerial surveys for monitoring small or declining elephant populations.     

Estimating the population size of the Sanje mangabey (Cercocebus sanjei) using acoustic distance sampling

The Sanje mangabey (Cercocebus sanjei) is endemic to the Udzungwa Mountains, Tanzania, and is classified as Endangered due to its putatively declining population size, habitat degradation and fragmentation. Previous population size estimates have ranged from 1,350 to 3,500 individuals, with the last direct survey being conducted 15 years before the present study. Previous estimates are now thought to have underestimated the population due to a limited knowledge of group and habitat size, nonsystematic approaches and the use of visual methods that are not suitable for surveying the Sanje mangabey with its semi-terrestrial and elusive behaviors. We used an acoustic survey method with observers recording the distinctive “whoop-gobble” vocalization produced by mangabeys and point transect distance sampling to model a detection function and estimate abundance. Twenty-eight surveys were conducted throughout the two forests where Sanje mangabeys are found: Mwanihana forest in the Udzungwa Mountains National Park (n = 13), and the Uzungwa Scarp Nature Reserve (n = 15). Group density was found to be significantly lower in the relatively unprotected Uzungwa Scarp forest (0.15 groups/km²; 95% CI: 0.08–0.27) compared to the well-protected Mwanihana forest (0.29 groups/km²; 95% CI: 0.19–0.43; p = .03). We estimate that there are 1,712 (95% CI: 1,141–2,567) individuals in Mwanihana and 1,455 (95% CI: 783–2,702) in the Uzungwa Scarp, resulting in a total population size of 3,167 (95% CI: 2,181–4,596) individuals. The difference in group density between sites is likely a result of the differing protection status and levels of enforcement between the forests, suggesting that protection of the Uzungwa Scarp should be increased to encourage recovery of the population, and reduce the threat of degradation and hunting. Our results contribute to the reassessment of the species' IUCN Red List status and informing management and conservation action planning.     

Forest elephant crisis in the Congo Basin

Debate over repealing the ivory trade ban dominates conferences of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Resolving this controversy requires accurate estimates of elephant population trends and rates of illegal killing. Most African savannah elephant populations are well known; however, the status of forest elephants, perhaps a distinct species, in the vast Congo Basin is unclear. We assessed population status and incidence of poaching from line-transect and reconnaissance surveys conducted on foot in sites throughout the Congo Basin. Results indicate that the abundance and range of forest elephants are threatened from poaching that is most intense close to roads. The probability of elephant presence increased with distance to roads, whereas that of human signs declined. At all distances from roads, the probability of elephant occurrence was always higher inside, compared to outside, protected areas, whereas that of humans was always lower. Inside protected areas, forest elephant density was correlated with the size of remote forest core, but not with size of protected area. Forest elephants must be prioritised in elephant management planning at the continental scale.     

Estimating the defaecation rate of the African forest elephant (Loxodonta cyclotis) in Banyang-Mbo Wildlife Sanctuary, south-western Cameroon

Defaecation rate is an important parameter in the conversion of dung counts to elephant numbers, yet estimates of elephant defaecation rates are available for a few sites, especially rainforest sites. The Banyang‐Mbo Wildlife Sanctuary in south‐western Cameroon is no exception in this regard. We tracked a radio‐collared herd of four forest elephants, counted their fresh droppings in two 24‐h periods each month in 36 consecutive months. Our results show a mean defaecation rate of 15.9 [95% CI (12.2–19.7)] dung piles per elephant per day. We found significant monthly and inter‐annual variation in defaecation rates with implications for their predictability and forest elephant surveys. In particular, our results show that mean defaecation rates are higher for the wettest period of the year than the relatively dry months.     

Estimating abundance for a savanna elephant population using mark–resight methods: a case study for the Tembe Elephant Park, South Africa

Elephants living in dense woodlands are difficult to count. Many elephant populations in Africa occur in such conditions. Estimates of these populations based on total counts, aerial counts and dung counts often lack information on precision and accuracy. We use standard mark–recapture field methods to obtain estimates of population size with associated confidence limits. We apply this approach to a closed elephant population in the Tembe Elephant Park (300 km²), South Africa. A registration count completed in 4 months gives a known population size. We evaluate mark–recapture models against the known population size. Individual identification profiles obtained for elephants during the registration count and mark–recapture events indicate that at least 167 elephants live in the park. We consider this value as an estimate of the minimum number alive. We include 189 sightings of bulls and 37 sightings of breeding herds in the mark–recapture modelling. Of the models we test (Petersen, Schnabel, Schumacher, Jolly–Seber, Bowden's, Poisson and negative binomial), Bowden's gives an estimate closest to the registration count. Assumptions of the model are not violated. For all models except one (negative binomial), our estimates improve with increased sampling intensity. Confidence intervals do not improve with increased effort except for the Schnabel model. Mark–recapture methods should be considered as reliable estimators of population size for elephants occurring in dense woodlands and forests when other methods cannot be relied on.     

Changes in Elephant Abundance Affect Forest Composition or Regeneration?

While overall numbers of African elephant have declined dramatically in recent times, some populations are now confined to protected areas and are locally overabundant—an undesirable situation for both biodiversity conservation and elephants. In forested protected areas, options to manage elephants are limited because it is difficult to safely approach animals, yet it is vital that these populations are managed because browsing by elephants can dramatically alter forest ecosystems. Using data collected over 50 yr in Kibale National Park, Uganda, we examine the prediction that increasing elephant numbers and associated changes in their foraging behavior have caused a shift in tree community composition. Although the relative abundance of elephants increased significantly between 1996 and 2010, the population structure of their preferred tree food species did not change, nor did tree community composition change in favor of species able to re‐sprout after elephant damage. Furthermore, over the last 50 yr Kibale elephants have not become more selective foragers, as would be expected if more nutritious tree species were declining. However, elephants are more abundant in disturbed areas dominated by shrubs and grasses and appear to have arrested forest succession in these areas. At their current abundance, elephants have not selectively altered the composition of intact old growth forest, but they do inhibit the regeneration of disturbed areas.     

Estimating the Barents Sea polar bear subpopulation size

A large-scale survey was conducted in August 2004 to estimate the size of the Barents Sea polar bear subpopulation. We combined helicopter line transect distance sampling (DS) surveys in most of the survey area with total counts in small areas not suitable for DS. Due to weather constraints we failed to survey some of the areas originally planned to be covered by DS. For those, abundance was estimated using a ratio estimator, in which the auxiliary variable was the number of satellite telemetry fixes (in previous years). We estimated that the Barents Sea subpopulation had approximately 2,650 (95% CI approximately 1,900–3,600) bears. Given current intense interest in polar bear management due to the potentially disastrous effects of climate change, it is surprising that many subpopulation sizes are still unknown. We show here that line transect sampling is a promising method for addressing the need for abundance estimates.     

Quantity and Configuration of Available Elephant Habitat and Related Conservation Concerns in the Lower Kinabatangan Floodplain of Sabah,Malaysia

The approximately 300 (298, 95% CI: 152–581) elephants in the Lower Kinabatangan Managed Elephant Range in Sabah, Malaysian Borneo are a priority sub-population for Borneo''s total elephant population (2,040, 95% CI: 1,184–3,652). Habitat loss and human-elephant conflict are recognized as the major threats to Bornean elephant survival. In the Kinabatangan region, human settlements and agricultural development for oil palm drive an intense fragmentation process. Electric fences guard against elephant crop raiding but also remove access to suitable habitat patches. We conducted expert opinion-based least-cost analyses, to model the quantity and configuration of available suitable elephant habitat in the Lower Kinabatangan, and called this the Elephant Habitat Linkage. At 184 km², our estimate of available habitat is 54% smaller than the estimate used in the State''s Elephant Action Plan for the Lower Kinabatangan Managed Elephant Range (400 km²). During high flood levels, available habitat is reduced to only 61 km². As a consequence, short-term elephant densities are likely to surge during floods to 4.83 km⁻² (95% CI: 2.46–9.41), among the highest estimated for forest-dwelling elephants in Asia or Africa. During severe floods, the configuration of remaining elephant habitat and the surge in elephant density may put two villages at elevated risk of human-elephant conflict. Lower Kinabatangan elephants are vulnerable to the natural disturbance regime of the river due to their limited dispersal options. Twenty bottlenecks less than one km wide throughout the Elephant Habitat Linkage, have the potential to further reduce access to suitable habitat. Rebuilding landscape connectivity to isolated habitat patches and to the North Kinabatangan Managed Elephant Range (less than 35 km inland) are conservation priorities that would increase the quantity of available habitat, and may work as a mechanism to allow population release, lower elephant density, reduce human-elephant conflict, and enable genetic mixing.