Double-observer line transect surveys with Markov-modulated Poisson process models for animal availability.

We develop maximum likelihood methods for line transect surveys in which animals go undetected at distance zero, either because they are stochastically unavailable while within view or because they are missed when they are available. These incorporate a Markov-modulated Poisson process model for animal availability, allowing more clustered availability events than is possible with Poisson availability models. They include a mark-recapture component arising from the independent-observer survey, leading to more accurate estimation of detection probability given availability. We develop models for situations in which (a) multiple detections of the same individual are possible and (b) some or all of the availability process parameters are estimated from the line transect survey itself, rather than from independent data. We investigate estimator performance by simulation, and compare the multiple-detection estimators with estimators that use only initial detections of individuals, and with a single-observer estimator. Simultaneous estimation of detection function parameters and availability model parameters is shown to be feasible from the line transect survey alone with multiple detections and double-observer data but not with single-observer data. Recording multiple detections of individuals improves estimator precision substantially when estimating the availability model parameters from survey data, and we recommend that these data be gathered. We apply the methods to estimate detection probability from a double-observer survey of North Atlantic minke whales, and find that double-observer data greatly improve estimator precision here too.

A Unifying Model for Capture-Recapture and Distance Sampling Surveys of Wildlife Populations.

A fundamental problem in wildlife ecology and management is estimation of population size or density. The two dominant methods in this area are capture-recapture (CR) and distance sampling (DS), each with its own largely separate literature. We develop a class of models that synthesizes them. It accommodates a spectrum of models ranging from nonspatial CR models (with no information on animal locations) through to DS and mark-recapture distance sampling (MRDS) models, in which animal locations are observed without error. Between these lie spatially explicit capture-recapture (SECR) models that include only capture locations, and a variety of models with less location data than are typical of DS surveys but more than are normally used on SECR surveys. In addition to unifying CR and DS models, the class provides a means of improving inference from SECR models by adding supplementary location data, and a means of incorporating measurement error into DS and MRDS models. We illustrate their utility by comparing inference on acoustic surveys of gibbons and frogs using only capture locations, using estimated angles (gibbons) and combinations of received signal strength and time-of-arrival data (frogs), and on a visual MRDS survey of whales, comparing estimates with exact and estimated distances. Supplementary materials for this article are available online.

Using hidden Markov models to deal with availability bias on line transect surveys.

We develop estimators for line transect surveys of animals that are stochastically unavailable for detection while within detection range. The detection process is formulated as a hidden Markov model with a binary state-dependent observation model that depends on both perpendicular and forward distances. This provides a parametric method of dealing with availability bias when estimates of availability process parameters are available even if series of availability events themselves are not. We apply the estimators to an aerial and a shipboard survey of whales, and investigate their properties by simulation. They are shown to be more general and more flexible than existing estimators based on parametric models of the availability process. We also find that methods using availability correction factors can be very biased when surveys are not close to being instantaneous, as can estimators that assume temporal independence in availability when there is temporal dependence.

Point transect sampling along linear features.

Distance sampling is a widely used methodology for assessing animal abundance. A key requirement of distance sampling is that samplers (lines or points) are placed according to a randomized design, which ensures that samplers are positioned independently of animals. Often samplers are placed along linear features such as roads, so that bias is expected if animals are not uniformly distributed with respect to distance from the linear feature. We present an approach for analyzing distance data from a survey when the samplers are points placed along a linear feature. Based on results from a simulation study and from a survey of Irish hares in Northern Ireland conducted from roads, we conclude that large bias may result if the position of samplers is not randomized, and analysis methods fail to account for nonuniformity.

Spatially explicit maximum likelihood methods for capture-recapture studies.

Live-trapping capture-recapture studies of animal populations with fixed trap locations inevitably have a spatial component: animals close to traps are more likely to be caught than those far away. This is not addressed in conventional closed-population estimates of abundance and without the spatial component, rigorous estimates of density cannot be obtained. We propose new, flexible capture-recapture models that use the capture locations to estimate animal locations and spatially referenced capture probability. The models are likelihood-based and hence allow use of Akaike's information criterion or other likelihood-based methods of model selection. Density is an explicit parameter, and the evaluation of its dependence on spatial or temporal covariates is therefore straightforward. Additional (nonspatial) variation in capture probability may be modeled as in conventional capture-recapture. The method is tested by simulation, using a model in which capture probability depends only on location relative to traps. Point estimators are found to be unbiased and standard error estimators almost unbiased. The method is used to estimate the density of Red-eyed Vireos (Vireo olivaceus) from mist-netting data from the Patuxent Research Refuge, Maryland, U.S.A. Estimates agree well with those from an existing spatially explicit method based on inverse prediction. A variety of additional spatially explicit models are fitted; these include models with temporal stratification, behavioral response, and heterogeneous animal home ranges.

Line transect methods for plant surveys.

Interest in surveys for monitoring plant abundance is increasing, due in part to the need to quantify the rate of loss of biodiversity. Line transect sampling offers an efficient way to monitor many species. However, the method does not work well in some circumstances, for example on small survey plots, when the plant species has a strongly aggregated distribution, or when plants that are on the line are not easily detected. We develop a crossed design, together with methods that exploit the additional information from such a design, to address these problems. The methods are illustrated using data on a colony of cowslips.

Accommodating unmodeled heterogeneity in double-observer distance sampling surveys.

Mark-recapture models applied to double-observer distance sampling data neglect the information on relative detectability of objects contained in the distribution of observed distances. A difference between the observed distribution and that predicted by the mark-recapture model is symptomatic of a failure of the assumption of zero correlation between detection probabilities implicit in the mark-recapture model. We develop a mark-recapture-based model that uses the observed distribution to relax this assumption to zero correlation at only one distance. We demonstrate its usefulness in coping with unmodeled heterogeneity using data from an aerial survey of crabeater seals in the Antarctic.

Wildlife population assessment: past developments and future directions.

We review the major developments in wildlife population assessment in the past century. Three major areas are considered: mark-recapture, distance sampling, and harvest models. We speculate on how these fields will develop in the next century. Topics for which we expect to see methodological advances include integration of modeling with Geographic Information Systems, automated survey design algorithms, advances in model-based inference from sample survey data, a common inferential framework for wildlife population assessment methods, improved methods for estimating population trends, the embedding of biological process models into inference, substantially improved models for conservation management, advanced spatiotemporal models of ecosystems, and greater emphasis on incorporating model selection uncertainty into inference. We discuss the kind of developments that might be anticipated in these topics.