Robert M. Dorazio

ESTIMATING SPECIES RICHNESS AND ACCUMULATION BY MODELING SPECIES OCCURRENCE AND DETECTABILITY

A statistical model is developed for estimating species richness and accumulation by formulating these community-level attributes as functions of model-based estimators of species occurrence while accounting for imperfect detection of individual species. The model requires a sampling protocol wherein repeated observations are made at a collection of sample locations selected to be representative of the community. This temporal replication provides the data needed to resolve the ambiguity between species absence and nondetection when species are unobserved at sample locations. Estimates of species richness and accumulation are computed for two communities, an avian community and a butterfly community. Our model-based estimates suggest that detection failures in many bird species were attributed to low rates of occurrence, as opposed to simply low rates of detection. We estimate that the avian community contains a substantial number of uncommon species and that species richness greatly exceeds the number of species actually observed in the sample. In fact, predictions of species accumulation suggest that even doubling the number of sample locations would not have revealed all of the species in the community. In contrast, our analysis of the butterfly community suggests that many species are relatively common and that the estimated richness of species in the community is nearly equal to the number of species actually detected in the sample. Our predictions of species accumulation suggest that the number of sample locations actually used in the butterfly survey could have been cut in half and the asymptotic richness of species still would have been attained. Our approach of developing occurrence-based summaries of communities while allowing for imperfect detection of species is broadly applicable and should prove useful in the design and analysis of surveys of biodiversity.

Estimating Size and Composition of Biological Communities by Modeling the Occurrence of Species

We develop a model that uses repeated observations of a biological community to estimate the number and composition of species in the community. Estimators of community-level attributes are constructed from model-based estimators of occurrence of individual species that incorporate imperfect detection of individuals. Data from the North American Breeding Bird Survey are analyzed to illustrate the variety of ecologically important quantities that are easily constructed and estimated using our model-based estimators of species occurrence. In particular, we compute site-specific estimates of species richness that honor classical notions of species-area relationships. We suggest extensions of our model to estimate maps of occurrence of individual species and to compute inferences related to the temporal and spatial dynamics of biological communities.

Mixture Models for Estimating the Size of a Closed Population When Capture Rates Vary among Individuals

We develop a parameterization of the beta-binomial mixture that provides sensible inferences about the size of a closed population when probabilities of capture or detection vary among individuals. Three classes of mixture models (beta-binomial, logistic-normal, and latent-class) are fitted to recaptures of snowshoe hares for estimating abundance and to counts of bird species for estimating species richness. In both sets of data, rates of detection appear to vary more among individuals (animals or species) than among sampling occasions or locations. The estimates of population size and species richness are sensitive to model-specific assumptions about the latent distribution of individual rates of detection. We demonstrate using simulation experiments that conventional diagnostics for assessing model adequacy, such as deviance, cannot be relied on for selecting classes of mixture models that produce valid inferences about population size. Prior knowledge about sources of individual heterogeneity in detection rates, if available, should be used to help select among classes of mixture models that are to be used for inference.

Analysis of Multinomial Models With Unknown Index Using Data Augmentation

Multinomial models with unknown index (“sample size”) arise in many practical settings. In practice, Bayesian analysis of such models has proved difficult because the dimension of the parameter space is not fixed, being in some cases a function of the unknown index. We describe a data augmentation approach to the analysis of this class of models that provides for a generic and efficient Bayesian implementation. Under this approach, the data are augmented with all-zero detection histories. The resulting augmented dataset is modeled as a zero-inflated version of the complete-data model where an estimable zero-inflation parameter takes the place of the unknown multinomial index. Interestingly, data augmentation can be justified as being equivalent to imposing a discrete uniform prior on the multinomial index. We provide three examples involving estimating the size of an animal population, estimating the number of diabetes cases in a population using the Rasch model, and the motivating example of estimating the number of species in an animal community with latent probabilities of species occurrence and detection.

Hierarchical models of animal abundance and occurrence

Much of animal ecology is devoted to studies of abundance and occurrence of species, based on surveys of spatially referenced sample units. These surveys frequently yield sparse counts that are contaminated by imperfect detection, making direct inference about abundance or occurrence based on observational data infeasible. This article describes a flexible hierarchical modeling framework for estimation and inference about animal abundance and occurrence from survey data that are subject to imperfect detection. Within this framework, we specify models of abundance and detectability of animals at the level of the local populations defined by the sample units. Information at the level of the local population is aggregated by specifying models that describe variation in abundance and detection among sites. We describe likelihood-based and Bayesian methods for estimation and inference under the resulting hierarchical model. We provide two examples of the application of hierarchical models to animal survey data, the first based on removal counts of stream fish and the second based on avian quadrat counts. For both examples, we provide a Bayesian analysis of the models using the software WinBUGS.

Bayesian inference and decision theory - A framework for decision making in natural resource management

Bayesian inference and decision theory may be used in the solution of relatively complex problems of natural resource management, owing to recent advances in statistical theory and computing. In particular, Markov chain Monte Carlo algorithms provide a computational framework for fitting models of adequate complexity and for evaluating the expected consequences of alternative management actions. We illustrate these features using an example based on management of waterfowl habitat. Corresponding Editor: D. B. Lindenmayer.

Parameter-expanded data augmentation for Bayesian analysis of capture-recapture models

Data augmentation (DA) is a flexible tool for analyzing closed and open population models of capture–recapture data, especially models which include sources of hetereogeneity among individuals. The essential concept underlying DA, as we use the term, is based on adding “observations” to create a dataset composed of a known number of individuals. This new (augmented) dataset, which includes the unknown number of individuals N in the population, is then analyzed using a new model that includes a reformulation of the parameter N in the conventional model of the observed (unaugmented) data. In the context of capture–recapture models, we add a set of “all zero” encounter histories which are not, in practice, observable. The model of the augmented dataset is a zero-inflated version of either a binomial or a multinomial base model. Thus, our use of DA provides a general approach for analyzing both closed and open population models of all types. In doing so, this approach provides a unified framework for the analysis of a huge range of models that are treated as unrelated “black boxes” and named procedures in the classical literature. As a practical matter, analysis of the augmented dataset by MCMC is greatly simplified compared to other methods that require specialized algorithms. For example, complex capture–recapture models of an augmented dataset can be fitted with popular MCMC software packages (WinBUGS or JAGS) by providing a concise statement of the model’s assumptions that usually involves only a few lines of pseudocode. In this paper, we review the basic technical concepts of data augmentation, and we provide examples of analyses of closed-population models (M0, Mh, distance sampling, and spatial capture–recapture models) and open-population models (Jolly–Seber) with individual effects.

Models for inference in dynamic metacommunity systems

A variety of processes are thought to be involved in the formation and dynamics of species assemblages. For example, various metacommunity theories are based on differences in the relative contributions of dispersal of species among local communities and interactions of species within local communities. Interestingly, metacommunity theories continue to be advanced without much empirical validation. Part of the problem is that statistical models used to analyze typical survey data either fail to specify ecological processes with sufficient, complexity or they fail to account for errors in detection of species during sampling. In this paper, we describe a statistical modeling framework for the analysis of metacommunity dynamics that is based on the idea of adopting a unified approach, multispecies occupancy modeling, for computing inferences about individual species, local communities of species, or the entire metacommunity of species. This approach accounts for errors in detection of species during sampling and also allows different metacommunity paradigms to be specified in terms of species- and location-specific probabilities of occurrence, extinction, and colonization: all of which are estimable. In addition, this approach can be used to address inference problems that arise in conservation ecology, such as predicting temporal and spatial changes in biodiversity for use in making conservation decisions. To illustrate, we estimate changes in species composition associated with the species-specific phenologies of flight patterns of butterflies in Switzerland for the purpose of estimating regional differences in biodiversity.

A new parameterization for estimating co‐occurrence of interacting species

Models currently used to estimate patterns of species co-occurrence while accounting for errors in detection of species can be difficult to fit when the effects of covariates on species occurrence probabilities are included. The source of the estimation problems is the particular parameterization used to specify species co-occurrence probability. We develop a new parameterization for estimating patterns of co-occurrence of interacting species that allows the effects of covariates to be specified quite naturally without estimation problems. In our model, the occurrence of one species is assumed to depend on the occurrence of another, but the occurrence of the second species is not assumed to depend on the presence of the first species. This pattern of co-occurrence, wherein one species is dominant and the other is subordinate, can be produced by several types of ecological interactions (predator-prey, parasitism, and so on). A simulation study demonstrated that estimates of species occurrence probabilities were unbiased in samples of 50-100 locations and three surveys per location, provided species are easily detected (probability of detection > or = 0.5). Higher sample sizes (>200 locations) are needed to achieve unbiasedness when species are more difficult to detect. An analysis of data from treefrog surveys in southern Florida indicated that the occurrence of Cuban treefrogs, an invasive predator species, was highest near the point of its introduction and declined with distance from that location. Sites occupied by Cuban treefrogs were 9.0 times less likely to contain green treefrogs and 15.7 times less likely to contain squirrel treefrogs compared to sites without Cuban treefrogs. The detection probabilities of native treefrog species did not depend on the presence of Cuban treefrogs, suggesting that the native treefrog species are naive to the introduced species.

ON THE CHOICE OF STATISTICAL MODELS FOR ESTIMATING OCCURRENCE AND EXTINCTION FROM ANIMAL SURVEYS

In surveys of natural animal populations the number of animals that are present and available to be detected at a sample location is often low, resulting in few or no detections. Low detection frequencies are especially common in surveys of imperiled species; however, the choice of sampling method and protocol also may influence the size of the population that is vulnerable to detection. In these circumstances, probabilities of animal occurrence and extinction will generally be estimated more accurately if the models used in data analysis account for differences in abundance among sample locations and for the dependence between site-specific abundance and detection. Simulation experiments are used to illustrate conditions wherein these types of models can be expected to outperform alternative estimators of population site occupancy and extinction.