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A temporally stratified extension of space‐for‐time Cormack–Jolly–Seber for migratory animals

Understanding drivers of temporal variation in demographic parameters is a central goal of mark‐recapture analysis. To estimate the survival of migrating animal populations in migration corridors, space‐for‐time mark–recapture models employ discrete sampling locations in space to monitor marked populations as they move past monitoring sites, rather than the standard practice of using fixed sampling points in time. Because these models focus on estimating survival over discrete spatial segments, model parameters are implicitly integrated over the temporal dimension. Furthermore, modeling the effect of time‐varying covariates on model parameters is complicated by unknown passage times for individuals that are not detected at monitoring sites. To overcome these limitations, we extended the Cormack–Jolly–Seber (CJS) framework to estimate temporally stratified survival and capture probabilities by including a discretized arrival time process in a Bayesian framework. We allow for flexibility in the model form by including temporally stratified covariates and hierarchical structures. In addition, we provide tools for assessing model fit and comparing among alternative structural models for the parameters. We demonstrate our framework by fitting three competing models to estimate daily survival, capture, and arrival probabilities at four hydroelectric dams for over 200 000 individually tagged migratory juvenile salmon released into the Snake River, USA.

Biometrics

Multivariate Bayesian clustering using covariate-informed components with application to boreal vegetation sensitivity

Climate change is impacting both the distribution and abundance of vegetation, especially in far northern latitudes. The effects of climate change are different for every plant assemblage and vary heterogeneously in both space and time. Small changes in climate could result in large vegetation responses in sensitive assemblages but weak responses in robust assemblages. But, patterns and mechanisms of sensitivity and robustness are not yet well understood, largely due to a lack of long-term measurements of climate and vegetation. Fortunately, observations are sometimes available across a broad spatial extent. We develop a novel statistical model for a multivariate response based on unknown cluster-specific effects and covariances, where cluster labels correspond to sensitivity and robustness. Our approach utilizes a prototype model for cluster membership that offers flexibility while enforcing smoothness in cluster probabilities across sites with similar characteristics. We demonstrate our approach with an application to vegetation abundance in Alaska, USA, in which we leverage the broad spatial extent of the study area as a proxy for unrecorded historical observations. In the context of the application, our approach yields interpretable site-level cluster labels associated with assemblage-level sensitivity and robustness without requiring strong a priori assumptions about the drivers of climate sensitivity.

Alaska

Combining band recovery data and Pollock's robust design to model temporary and permanent emigration

Capture-recapture models are widely used to estimate demographic parameters of marked populations. Recently, this statistical theory has been extended to modeling dispersal of open populations. Multistate models can be used to estimate movement probabilities among subdivided populations if multiple sites are sampled. Frequently, however, sampling is limited to a single site, Models described by Burnham (1993, in Marked Individuals in the Study of Bird Populations , 199–213), which combined open population capture-recapture and band-recovery models, can be used to estimate permanent emigration when sampling is limited to a single population. Similarly, Kendall, Nichols, and Hines (1997, Ecology 51 , 563–578) developed models to estimate temporary emigration under Pollock's (1982, Journal of Wildlife Management 46 , 757–760) robust design. We describe a likelihood-based approach to simultaneously estimate temporary and permanent emigration when sampling is limited to a single population. We use a sampling design that combines the robust design and recoveries of individuals obtained immediately following each sampling period. We present a general form for our model where temporary emigration is a first-order Markov process, and we discuss more restrictive models. We illustrate these models with analysis of data on marked Canvasback ducks. Our analysis indicates that probability of permanent emigration for adult female Canvasbacks was 0.193 ( ) and that birds that were present at the study area in year i — 1 had a higher probability of presence in year i than birds that were not present in year i — 1.

Biometrics

Using open robust design models to estimate temporary emigration from capture-recapture data

Capture-recapture studies are crucial in many circumstances for estimating demographic parameters for wildlife and fish populations. Pollock's robust design, involving multiple sampling occasions per period of interest, provides several advantages over classical approaches. This includes the ability to estimate the probability of being present and available for detection, which in some situations is equivalent to breeding probability. We present a model for estimating availability for detection that relaxes two assumptions required in previous approaches. The first is that the sampled population is closed to additions and deletions across samples within a period of interest. The second is that each member of the population has the same probability of being available for detection in a given period. We apply our model to estimate survival and breeding probability in a study of hawksbill sea turtles ( Eretmochelys imbricata ), where previous approaches are not appropriate.

Biometrics

Demographic analysis from summaries of an age-structured population

Demographic analyses of age-structured populations typically rely on life history data for individuals, or when individual animals are not identified, on information about the numbers of individuals in each age class through time. While it is usually difficult to determine the age class of a randomly encountered individual, it is often the case that the individual can be readily and reliably assigned to one of a set of age classes. For example, it is often possible to distinguish first-year from older birds. In such cases, the population age structure can be regarded as a latent variable governed by a process prior, and the data as summaries of this latent structure. In this article, we consider the problem of uncovering the latent structure and estimating process parameters from summaries of age class information. We present a demographic analysis for the critically endangered migratory population of whooping cranes (Grus americana), based only on counts of first-year birds and of older birds. We estimate age and year-specific survival rates. We address the controversial issue of whether management action on the breeding grounds has influenced recruitment, relating recruitment rates to the number of seventh-year and older birds, and examining the pattern of variation through time in this rate.

Biometrics

Nonidentifiability of population size from capture-recapture data with heterogeneous detection probabilities

Heterogeneity in detection probabilities has long been recognized as problematic in mark-recapture studies, and numerous models developed to accommodate its effects. Individual heterogeneity is especially problematic, in that reasonable alternative models may predict essentially identical observations from populations of substantially different sizes. Thus even with very large samples, the analyst will not be able to distinguish among reasonable models of heterogeneity, even though these yield quite distinct inferences about population size. The problem is illustrated with models for closed and open populations.

Biometrics

N-mixture models for estimating population size from spatially replicated counts

Spatial replication is a common theme in count surveys of animals. Such surveys often generate sparse count data from which it is difficult to estimate population size while formally accounting for detection probability. In this article, I describe a class of models (n-mixture models) which allow for estimation of population size from such data. The key idea is to view site-specific population sizes, n, as independent random variables distributed according to some mixing distribution (e.g., Poisson). Prior parameters are estimated from the marginal likelihood of the data, having integrated over the prior distribution for n. Carroll and Lombard (1985, Journal of American Statistical Association 80, 423-426) proposed a class of estimators based on mixing over a prior distribution for detection probability. Their estimator can be applied in limited settings, but is sensitive to prior parameter values that are fixed a priori. Spatial replication provides additional information regarding the parameters of the prior distribution on n that is exploited by the n-mixture models and which leads to reasonable estimates of abundance from sparse data. A simulation study demonstrates superior operating characteristics (bias, confidence interval coverage) of the n-mixture estimator compared to the Caroll and Lombard estimator. Both estimators are applied to point count data on six species of birds illustrating the sensitivity to choice of prior on p and substantially different estimates of abundance as a consequence.

Biometrics

A general model for the analysis of mark-resight, mark-recapture, and band-recovery data under tag loss

Estimates of waterfowl demographic parameters often come from resighting studies where birds fit with individually identifiable neck collars are resighted at a distance. Concerns have been raised about the effects of collar loss on parameter estimates, and the reliability of extrapolating from collared individuals to the population. Models previously proposed to account for collar loss do not allow survival or harvest parameters to depend on neck collar presence or absence. Also, few models have incorporated recent advances in mark‐recapture theory that allow for multiple states or auxiliary encounters such as band recoveries. We propose a multistate model for tag loss in which the presence or absence of a collar is considered as a state variable. In this framework, demographic parameters are corrected for tag loss and questions related to collar effects on survival and recovery rates can be addressed. Encounters of individuals between closed sampling periods also can be incorporated in the analysis. We discuss data requirements for answering questions related to tag loss and sampling designs that lend themselves to this purpose. We illustrate the application of our model using a study of lesser snow geese ( Chen caerulescens caerulescens ).

Biometrics

Modeling association among demographic parameters in analysis of open population capture-recapture data

We present a hierarchical extension of the Cormack–Jolly–Seber (CJS) model for open population capture–recapture data. In addition to recaptures of marked animals, we model first captures of animals and losses on capture. The parameter set includes capture probabilities, survival rates, and birth rates. The survival rates and birth rates are treated as a random sample from a bivariate distribution, thus the model explicitly incorporates correlation in these demographic rates. A key feature of the model is that the likelihood function, which includes a CJS model factor, is expressed entirely in terms of identifiable parameters; losses on capture can be factored out of the model. Since the computational complexity of classical likelihood methods is prohibitive, we use Markov chain Monte Carlo in a Bayesian analysis. We describe an efficient candidate-generation scheme for Metropolis–Hastings sampling of CJS models and extensions. The procedure is illustrated using mark-recapture data for the moth Gonodontis bidentata .

Biometrics

Improving removal-based estimates of abundance by sampling a population of spatially distinct subpopulations

A statistical modeling framework is described for estimating the abundances of spatially distinct subpopulations of animals surveyed using removal sampling. To illustrate this framework, hierarchical models are developed using the Poisson and negative-binomial distributions to model variation in abundance among subpopulations and using the beta distribution to model variation in capture probabilities. These models are fitted to the removal counts observed in a survey of a federally endangered fish species. The resulting estimates of abundance have similar or better precision than those computed using the conventional approach of analyzing the removal counts of each subpopulation separately. Extension of the hierarchical models to include spatial covariates of abundance is straightforward and may be used to identify important features of an animal's habitat or to predict the abundance of animals at unsampled locations.

Biometrics

Analysis of capture–recapture models with individual covariates using data augmentation

I consider the analysis of capture–recapture models with individual covariates that influence detection probability. Bayesian analysis of the joint likelihood is carried out using a flexible data augmentation scheme that facilitates analysis by Markov chain Monte Carlo methods, and a simple and straightforward implementation in freely available software. This approach is applied to a study of meadow voles ( Microtus pennsylvanicus ) in which auxiliary data on a continuous covariate (body mass) are recorded, and it is thought that detection probability is related to body mass. In a second example, the model is applied to an aerial waterfowl survey in which a double‐observer protocol is used. The fundamental unit of observation is the cluster of individual birds, and the size of the cluster (a discrete covariate) is used as a covariate on detection probability.

Biometrics

Predicting the geographic distribution of a species from presence-only data subject to detection errors

Several models have been developed to predict the geographic distribution of a species by combining measurements of covariates of occurrence at locations where the species is known to be present with measurements of the same covariates at other locations where species occurrence status (presence or absence) is unknown. In the absence of species detection errors, spatial point-process models and binary-regression models for case-augmented surveys provide consistent estimators of a species’ geographic distribution without prior knowledge of species prevalence. In addition, these regression models can be modified to produce estimators of species abundance that are asymptotically equivalent to those of the spatial point-process models. However, if species presence locations are subject to detection errors, neither class of models provides a consistent estimator of covariate effects unless the covariates of species abundance are distinct and independently distributed from the covariates of species detection probability. These analytical results are illustrated using simulation studies of data sets that contain a wide range of presence-only sample sizes. Analyses of presence-only data of three avian species observed in a survey of landbirds in western Montana and northern Idaho are compared with site-occupancy analyses of detections and nondetections of these species.

Biometrics

A parametric generalization of the Hayne estimator for line transect sampling

The Hayne model for line transect sampling is generalized by using an elliptical (rather than circular) flushing model for animal detection. By assuming the ration of major and minor axes lengths is constant for all animals, a model results which allows estimation of population density based directly upon sighting distances and sighting angles. The derived estimator of animal density is a generalization of the Hayne estimator for line transect sampling.

Biometrics

Nonparametric estimation of plant density by the distance method

A relation between the plant density and the probability density function of the nearest neighbor distance (squared) from a random point is established under fairly broad conditions. Based upon this relationship, a nonparametric estimator for the plant density is developed and presented in terms of order statistics. Consistency and asymptotic normality of the estimator are discussed. An interval estimator for the density is obtained. The modifications of this estimator and its variance are given when the distribution is truncated. Simulation results are presented for regular, random and aggregated populations to illustrate the nonparametric estimator and its variance. A numerical example from field data is given. Merits and deficiencies of the estimator are discussed with regard to its robustness and variance.

Biometrics

Goodness-of-fit tests for open capture-recapture models

General goodness-of-fit tests for the Jolly-Seber model are proposed. These tests are based on conditional arguments using minimal sufficient statistics. The tests are shown to be of simple hypergeometric form so that a series of independent contingency table chi-square tests can be performed. The relationship of these tests to other proposed tests is discussed. This is followed by a simulation study of the power of the tests to detect departures from the assumptions of the Jolly-Seber model. Some meadow vole capture-recapture data are used to illustrate the testing procedure which has been implemented in a computer program available from the authors.

Biometrics

A general methodology for maximum likelihood inference from band-recovery data

A numerical procedure is described for obtaining maximum likelihood estimates and associated maximum likelihood inference from band- recovery data. The method is used to illustrate previously developed one-age-class band-recovery models, and is extended to new models, including the analysis with a covariate for survival rates and variable-time-period recovery models. Extensions to R-age-class band- recovery, mark-recapture models, and twice-yearly marking are discussed. A FORTRAN program provides computations for these models.

Biometrics

The use of auxiliary variables in capture-recapture and removal experiments

The dependence of animal capture probabilities on auxiliary variables is an important practical problem which has not been considered in the development of estimation procedures for capture-recapture and removal experiments. In this paper the linear logistic binary regression model is used to relate the probability of capture to continuous auxiliary variables. The auxiliary variables could be environmental quantities such as air or water temperature, or characteristics of individual animals, such as body length or weight. Maximum likelihood estimators of the population parameters are considered for a variety of models which all assume a closed population. Testing between models is also considered. The models can also be used when one auxiliary variable is a measure of the effort expended in obtaining the sample.

Biometrics