Design and field methods for sighting surveys of cetaceans in coastal and riverine habitats

• 1 Dolphins and porpoises in coastal and/or riverine habitats face serious conservation threats, yet surveys of their abundance are often especially difficult due to the challenges imposed by the habitats. Because many of these species occur in developing countries, lack of resources imposes a further set of challenges.
• 2 We offer advice on designing and conducting line‐transect surveys with a focus on sound, practical, design rather than analytical sophistication, and we attempt, where possible, to offer simple, inexpensive solutions.
• 3 We guide the reader through the questions of what kind of survey should be done, whether by boat or aircraft, and we discuss ways to avoid bias and increase precision.
• 4 Our treatment of field methods focuses especially on robust, but low‐cost, approaches. We provide two case studies to illustrate the implementation of these ideas.

     

Accounting for imperfect detection of groups and individuals when estimating abundance

If animals are independently detected during surveys, many methods exist for estimating animal abundance despite detection probabilities <1. Common estimators include double‐observer models, distance sampling models and combined double‐observer and distance sampling models (known as mark‐recapture‐distance‐sampling models; MRDS). When animals reside in groups, however, the assumption of independent detection is violated. In this case, the standard approach is to account for imperfect detection of groups, while assuming that individuals within groups are detected perfectly. However, this assumption is often unsupported. We introduce an abundance estimator for grouped animals when detection of groups is imperfect and group size may be under‐counted, but not over‐counted. The estimator combines an MRDS model with an N‐mixture model to account for imperfect detection of individuals. The new MRDS‐Nmix model requires the same data as an MRDS model (independent detection histories, an estimate of distance to transect, and an estimate of group size), plus a second estimate of group size provided by the second observer. We extend the model to situations in which detection of individuals within groups declines with distance. We simulated 12 data sets and used Bayesian methods to compare the performance of the new MRDS‐Nmix model to an MRDS model. Abundance estimates generated by the MRDS‐Nmix model exhibited minimal bias and nominal coverage levels. In contrast, MRDS abundance estimates were biased low and exhibited poor coverage. Many species of conservation interest reside in groups and could benefit from an estimator that better accounts for imperfect detection. Furthermore, the ability to relax the assumption of perfect detection of individuals within detected groups may allow surveyors to re‐allocate resources toward detection of new groups instead of extensive surveys of known groups. We believe the proposed estimator is feasible because the only additional field data required are a second estimate of group size.     

Line Transect and Triangulation Surveys Provide Reliable Estimates of the Density of Kloss’ Gibbons (Hylobates klossii) on Siberut Island, Indonesia

Estimating population densities of key species is crucial for many conservation programs. Density estimates provide baseline data and enable monitoring of population size. Several different survey methods are available, and the choice of method depends on the species and study aims. Few studies have compared the accuracy and efficiency of different survey methods for large mammals, particularly for primates. Here we compare estimates of density and abundance of Kloss’ gibbons (Hylobates klossii) using two of the most common survey methods: line transect distance sampling and triangulation. Line transect surveys (survey effort: 155.5 km) produced a total of 101 auditory and visual encounters and a density estimate of 5.5 gibbon clusters (groups or subgroups of primate social units)/km². Triangulation conducted from 12 listening posts during the same period revealed a similar density estimate of 5.0 clusters/km². Coefficients of variation of cluster density estimates were slightly higher from triangulation (0.24) than from line transects (0.17), resulting in a lack of precision in detecting changes in cluster densities of <66 % for triangulation and <47 % for line transect surveys at the 5 % significance level with a statistical power of 50 %. This case study shows that both methods may provide estimates with similar accuracy but that line transects can result in more precise estimates and allow assessment of other primate species. For a rapid assessment of gibbon density under time and financial constraints, the triangulation method also may be appropriate.     

Total Counts Versus Line Transects for Estimating Abundance of Small Gopher Tortoise Populations

Abstract: Status and trends of gopher tortoise (Gopherus polyphemus) populations are a critical information need for natural resource managers, researchers, and policy makers. Many tortoise populations are small and isolated, which can present challenges for deriving population estimates. Our objective was to compare abundance and density estimates for a small tortoise population derived using a total burrow count versus estimates obtained with line transect distance sampling (LTDS) using repeated surveys. We also compared results of the 2 survey methods using standard burrow-to-tortoise correction factors versus assessing occupancy of all burrows with a camera scope. In addition, we compared LTDS data obtained using a compass and measuring tape to define transects to those obtained using a Global Positioning System (GPS) and Personal Data Assistant (PDA) field computer to navigate transects. Line transect distance sampling with repeated surveys (both with a measuring tape and compass and with a GPS—PDA) yielded sufficient observations of tortoises to calculate population estimates. From 18% to 31% of burrows were occupied by tortoises as determined with the burrow camera. We found 25 burrows during the LTDS survey that we did not find in the total count survey, which demonstrated that the assumption of 100% detection for the total count was not met; hence, density or abundance measurements derived with this method were underestimates. We recommend using GPS—PDA technology, scoping all burrows detected, and using LTDS with repeated surveys to estimate abundance and density for small gopher tortoise populations.     

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.     

Improving Aquatic Warbler Population Assessments by Accounting for Imperfect Detection

Monitoring programs designed to assess changes in population size over time need to account for imperfect detection and provide estimates of precision around annual abundance estimates. Especially for species dependent on conservation management, robust monitoring is essential to evaluate the effectiveness of management. Many bird species of temperate grasslands depend on specific conservation management to maintain suitable breeding habitat. One such species is the Aquatic Warbler (Acrocephalus paludicola), which breeds in open fen mires in Central Europe. Aquatic Warbler populations have so far been assessed using a complete survey that aims to enumerate all singing males over a large area. Because this approach provides no estimate of precision and does not account for observation error, detecting moderate population changes is challenging. From 2011 to 2013 we trialled a new line transect sampling monitoring design in the Biebrza valley, Poland, to estimate abundance of singing male Aquatic Warblers. We surveyed Aquatic Warblers repeatedly along 50 randomly placed 1-km transects, and used binomial mixture models to estimate abundances per transect. The repeated line transect sampling required 150 observer days, and thus less effort than the traditional ‘full count’ approach (175 observer days). Aquatic Warbler abundance was highest at intermediate water levels, and detection probability varied between years and was influenced by vegetation height. A power analysis indicated that our line transect sampling design had a power of 68% to detect a 20% population change over 10 years, whereas raw count data had a 9% power to detect the same trend. Thus, by accounting for imperfect detection we increased the power to detect population changes. We recommend to adopt the repeated line transect sampling approach for monitoring Aquatic Warblers in Poland and in other important breeding areas to monitor changes in population size and the effects of habitat management.     

Evaluating abundance estimates and evidence of breeding for Bobolinks from transect and point-count surveys

Estimating the abundance and breeding success of territorial songbirds is challenging. Various types of surveys and analyses are available, but all receive some criticism in the literature, and most methods are rarely compared with results obtained using intensive monitoring efforts. We assessed the efficacy of transect and point-count surveys to estimate the abundance of male Bobolinks (Dolichonyx oryzivorus) and detect evidence of nesting and fledging by comparing the results of those surveys to results from more intensive monitoring (i.e., spot mapping and nest monitoring). We monitored 36 fields (254 ha) of late-harvest hay, restored grassland, and fallow fields in the Luther Marsh Wildlife Management Area and on four farms in southern Ontario, Canada, in 2018. Compared to the number of territories identified based on spot mapping (197), distance sampling analysis of transect survey data provided a more accurate estimate of the abundance of male Bobolinks (230, 95% CI: 187, 282) than N-mixture models of transect (668, 95% CI: 332, 1342) and point-count (337, 95% CI: 203, 559) data. Three visits to survey transects and five to point counts did not effectively detect evidence of Bobolink breeding (i.e., nesting or fledging) in fields compared to spot mapping and nest monitoring. Distance sampling analysis of transect data appears promising for estimating the number of Bobolink territories in an area, e.g., those impacted by conservation programs. If estimates of the number of nesting Bobolinks and frequency of fledging are of interest, spot mapping and nest monitoring could be implemented at a subset of sampled fields. Our results suggest that additional studies to evaluate model-based estimates of abundance with the best available information (e.g., from spot mapping of marked or unmarked populations and nest monitoring) would be useful to ensure that robust estimates are provided to support population estimates and conservation actions.     

Optimising sampling methods for small mammal communities in Neotropical rainforests

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Design and Analysis of Line Transect Surveys for Primates

Line transect surveys are widely used for estimating abundance of primate populations. The method relies on a small number of key assumptions, and if these are not met, substantial bias may occur. For a variety of reasons, primate surveys often do not follow what is generally considered to be best practice, either in survey design or in analysis. The design often comprises too few lines (sometimes just 1), subjectively placed or placed along trails, so lacks both randomization and adequate replication. Analysis often involves flawed or inefficient models, and often uses biased estimates of the locations of primate groups relative to the line. We outline the standard method, emphasizing the assumptions underlying the approach. We then consider options for when it is difficult or impossible to meet key assumptions. We explore the performance of these options by simulation, focusing particularly on the analysis of primate group sizes, where many of the variations in survey methods have been developed. We also discuss design issues, field methods, analysis, and potential alternative methodologies for when standard line transect sampling cannot deliver reliable abundance estimates.     

鸟类调查的样线法和样点法比较:以崇明东滩春季盐沼鸟类调查为例

为了探讨样线法和样点法对盐沼湿地鸟类调查的有效性及适用性, 我们于2008年3–5月采用固定距离样线法和固定半径样点法对崇明东滩盐沼鸟类进行了调查。5次调查共记录到鸟类24种, 其中样线法记录到19种, 样点法记录到21种, 两种调查方法平均每次记录到的鸟类种数没有显著差异, 表明两种方法对盐沼鸟类种类的发现能力相似。两种方法得到的鸟类密度之间有显著的相关性,说明两种方法在反映鸟类群落中每种鸟类的相对数量方面具有可比性。但除了仅在一种调查方法中记录到的种类, 样点法调查得到的每种鸟类密度和鸟类总密度均高于样线法的调查结果。因此, 在对鸟类种群或群落的时空特征进行比较时, 需要考虑所采用的调查方法对调查结果的影响。     

Aerial surveys suggest long‐term stability in the seasonally ice‐free Foxe Basin (Nunavut) polar bear population

Significant information gaps exist regarding the status of polar bears, especially with respect to the impacts of climate change, across large portions of the Arctic. To obtain an updated abundance estimate for the Foxe Basin population, we conducted comprehensive aerial surveys during the 2009 and 2010 ice‐free seasons, when bears are confined to land. We sampled with mark‐recapture distance sampling protocols on inland and coastal transects and surveyed small islands and remnant ice floes. We observed 816 and 1,003 bears in 2009 and 2010, respectively. Although detection functions differed substantially between years, estimates were consistent between analytical methods and years. Averaging four estimates (two from each year) yielded 2,585 (2,096–3,189) bears, which is similar to an estimate from the 1990s. This result, along with robust cub production, suggests a stable and healthy population despite deteriorating sea ice conditions. Collectively, this and other recent on‐land surveys provide a framework for implementing aerial surveys elsewhere. Although aerial surveys do not yield estimates of vital rates or population growth, they enable more rapid and frequent monitoring than mark‐recapture. Integrating them in long‐term monitoring programs will require consideration of ancillary data to infer status and facilitate setting harvest levels.     

Comparative performance of four survey methods for assessing Lilian’s Lovebird abundance in Liwonde National Park,Malawi

Monitoring abundance of threatened species is important for conservation planning. Lilian’s Lovebird Agapornis lilianae is a near-threatened small parrot found in mopane Colophospermum mopane woodland. Its population has not been investigated in any part of its range. We investigated the abundance and density of the Lilian’s Lovebird in Liwonde National Park, Malawi. Both distance sampling (line and point transects) methods and total counts (waterhole and flyway counts) were applied. The point count method gave very low numbers and was discontinued after the first year. Line transects conducted during the wet season had the highest density estimates of 17 ± 4.8 lovebirds km?2 of mopane woodland. However, number of observations per transect in each year were low. Waterhole counts had the lowest density estimates (10 ± 3.5 lovebirds km?2). Flyway counts gave an intermediate estimate (13 ± 3.0 lovebirds km?2). The total population of Lilian’s Lovebirds in Liwonde National Park is therefore estimated to be about 4 000 individuals. The use of line transect counts at the end of the rainy season is recommended for continued monitoring of Lilian’s Lovebirds abundance in Liwonde National Park.     

An Evaluation of Survey Methods for Estimating Northern Bobwhite Abundance in Southern Texas

Abstract: Distance sampling has been identified as a reliable and well-suited method for estimating northern bobwhite (Colinus virginianus) density. However, distance sampling using walked transects requires intense sampling to obtain precise estimates, thus making the technique impractical for large acreages. Researchers have addressed this limitation by either resorting to the use of indices (e.g., morning covey-call surveys) or incorporating the use of aerial surveys with distance sampling. Both approaches remain relatively untested. Our objectives were to 1) compare density estimates among morning covey-call surveys, helicopter transects, and walked transects; 2) test a critical assumption of distance sampling pertinent to helicopter surveys (i.e., all objects on line are detected); and 3) evaluate the underlying premise of morning covey-call surveys (i.e., that the no. of calling coveys correlates with bobwhite density). Our study was conducted on 3 study sites in Brooks County, Texas, USA, during October to December, 2001 to 2005. Comparisons between walked transects and morning covey-call surveys involved the entire 5-year data set, whereas helicopter transects involved only the latter 2 years. Density estimates obtained from helicopter transects were similar to walked transect estimates for both years. We documented a detection probability on the helicopter transect line of 70 ± 10.2% (% ± SE; n = 20 coveys). Morning covey-call surveys yielded similar density estimates to walked transect estimates during only 2 of 5 years, when walked transect estimates were the least accurate and precise. We detected a positive relationship (R² = 0.51; 95% CI for slope: 29.5–53.1; n = 63 observations) between covey density and number of coveys heard calling. We conclude that helicopter transects appear to be a viable alternative to walked transects for estimating density of bobwhites. Morning covey-call surveys appear to be a poor method to estimate absolute abundance and to depict general population trajectories.     

Abundance estimation of long-diving animals using line transect methods

Line transect sampling is one of the most widely used methods for estimating the size of wild animal populations. An assumption in standard line transect sampling is that all the animals on the trackline are detected without fail. This assumption tends to be violated for marine mammals with surfacing/diving behaviors. The detection probability on the trackline is estimated using duplicate sightings from double-platform line transect methods. The double-platform methods, however, are insufficient to estimate the abundance of long-diving animals because these animals can be completely missed while the observers pass. We developed a more flexible hazard probability model that incorporates information on surfacing/diving patterns obtained from telemetry data. The model is based on a stochastic point process and is statistically tractable. A simulation study showed that the new model provides near-unbiased abundance estimates, whereas the traditional hazard rate and hazard probability models produce considerably biased estimates. As an illustration, we applied the model to data on the Baird's beaked whale (Berardius bairdii) in the western North Pacific.     

Modelling heterogeneity in detection probabilities in land and aerial abundance surveys in humpback whales (<Emphasis Type="Italic">Megaptera novaeangliae</Emphasis>)

The effective management and conservation of animal populations relies on statistically-sound and replicable surveys to obtain estimates of abundance and assess trends. Surveys of cetaceans, such as humpback whales Megaptera novaeangliae, are difficult to conduct and are particularly affected by bias in detection probability. For example, the probability of detection of whales from land decreases substantially with increased distance from the platform. This distance effect is also true for aerial surveys, combined with the problem that animals are unavailable for detection (underwater) whilst in the field of view. We present a novel approach that combines corrected double-platform land surveys with corrected aerial surveys to obtain a robust estimate of g(0), the probability of detection on the survey line, for aerial surveys of migrating humpback whales. Several sources of heterogeneity in detection probabilities were identified within the land and aerial surveys (including group composition, bearing of first sighting, number of groups being tracked simultaneously and cloud cover). After including these into our estimate of ĝ(0), we found that only 29% of available whales are being detected on the survey line (ĝ(0) = 0.288), which is a considerably smaller estimate than many available for humpback whales using other methods. Incorporating heterogeneity into the population surveys shows that we are likely to be underestimating the population size of whales on the east coast of Australia. The implications of this result for their conservation and management in light of increased whale-human conflict is discussed.     

Inferring trackline detection probabilities,g(0), for cetaceans from apparent densities in different survey conditions

Visual line‐transect surveys are commonly used to estimate cetacean abundance. A key parameter in such studies is g(0), the probability of detecting an animal that is directly on the transect line. This is typically considered to be constant for a species across survey conditions. A method is developed to estimate the relative values of g(0) in different survey conditions (Beaufort state) by comparing Beaufort‐specific density estimates. The approach is based on fitting generalized additive models, with the presence of a sighting on a survey segment as the dependent variable, Beaufort state as the key explanatory variable, and year, latitude, and longitude as nuisance variables to control for real differences in density over time and space. Values of relative g(0) are estimated for 20 cetacean taxa using 175,000 km of line‐transect survey data from the eastern and central Pacific Ocean from 1986 to 2010. Results show that g(0) decreases as Beaufort state increases, even for visually conspicuous species. This effect is greatest for the least conspicuous species (rough‐toothed dolphins, beaked whales, minke whales, and dwarf and pygmy sperm whales). Ignoring these large effects results in a nontrivial bias in cetacean abundance estimates.     

Distance models as a tool for modelling detection probability and density of native bumblebees

Effective monitoring of native bee populations requires accurate estimates of population size and relative abundance among habitats. Current bee survey methods, such as netting or pan trapping, may be adequate for a variety of study objectives but are limited by a failure to account for imperfect detection. Biases due to imperfect detection could result in inaccurate abundance estimates or erroneous insights about the response of bees to different environments. To gauge the potential biases of currently employed survey methods, we compared abundance estimates of bumblebees (Bombus spp.) derived from hierarchical distance sampling models (HDS) to bumblebee counts collected from fixed‐area net surveys (“net counts”) and fixed‐width transect counts (“transect counts”) at 47 early‐successional forest patches in Pennsylvania. Our HDS models indicated that detection probabilities of Bombus spp. were imperfect and varied with survey‐ and site‐covariates. Despite being conspicuous, Bombus spp. were not reliably detected beyond 5 m. Habitat associations of Bombus spp. density were similar across methods, but the strength of association with shrub cover differed between HDS and net counts. Additionally, net counts suggested sites with more grass hosted higher Bombus spp. densities whereas HDS suggested that grass cover was associated with higher detection probability but not Bombus spp. density. Density estimates generated from net counts and transect counts were 80%–89% lower than estimates generated from distance sampling. Our findings suggest that distance modelling provides a reliable method to assess Bombus spp. density and habitat associations, while accounting for imperfect detection caused by distance from observer, vegetation structure, and survey covariates. However, detection/non‐detection data collected via point‐counts, line‐transects and distance sampling for Bombus spp. are unlikely to yield species‐specific density estimates unless individuals can be identified by sight, without capture. Our results will be useful for informing the design of monitoring programs for Bombus spp. and other pollinators.     

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.

     

To See or Not to See: Investigating Detectability of Ganges River Dolphins Using a Combined Visual-Acoustic Survey

Detection of animals during visual surveys is rarely perfect or constant, and failure to account for imperfect detectability affects the accuracy of abundance estimates. Freshwater cetaceans are among the most threatened group of mammals, and visual surveys are a commonly employed method for estimating population size despite concerns over imperfect and unquantified detectability. We used a combined visual-acoustic survey to estimate detectability of Ganges River dolphins (Platanista gangetica gangetica) in four waterways of southern Bangladesh. The combined visual-acoustic survey resulted in consistently higher detectability than a single observer-team visual survey, thereby improving power to detect trends. Visual detectability was particularly low for dolphins close to meanders where these habitat features temporarily block the view of the preceding river surface. This systematic bias in detectability during visual-only surveys may lead researchers to underestimate the importance of heavily meandering river reaches. Although the benefits of acoustic surveys are increasingly recognised for marine cetaceans, they have not been widely used for monitoring abundance of freshwater cetaceans due to perceived costs and technical skill requirements. We show that acoustic surveys are in fact a relatively cost-effective approach for surveying freshwater cetaceans, once it is acknowledged that methods that do not account for imperfect detectability are of limited value for monitoring.     

Line Transect Sampling in Small Regions

This article develops an approach to estimating population abundance from line transect surveys that uses a calibration survey to estimate the detection function, which is then employed as a weight function in constructing the abundance estimate. Nonparametric methods of estimating the detection function via local regression and via a kernel density estimator are considered. The proposed methods are evaluated using a set of Western Australian plant data and weed enumeration data.