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D.I. MacKenzie

Publications and source records attributed to D.I. MacKenzie.

11 recordsLinked to original sources

Occupancy estimation and modeling with multiple states and state uncertainty

The distribution of a species over space is of central interest in ecology, but species occurrence does not provide all of the information needed to characterize either the well-being of a population or the suitability of occupied habitat. Recent methodological development has focused on drawing inferences about species occurrence in the face of imperfect detection. Here we extend those methods by characterizing occupied locations by some additional state variable ( e. g., as producing young or not). Our modeling approach deals with both detection probabilities,1 and uncertainty in state classification. We then use the approach with occupancy and reproductive rate data from California Spotted Owls ( Strix occidentalis occidentalis ) collected in the central Sierra Nevada during the breeding season of 2004 to illustrate the utility of the modeling approach. Estimates of owl reproductive rate were larger than naive estimates, indicating the importance of appropriately accounting for uncertainty in detection and state classification.

Ecology

The use of photographic rates to estimate densities of tigers and other cryptic mammals: a comment on misleading conclusions

The search for easy-to-use indices that substitute for direct estimation of animal density is a common theme in wildlife and conservation science, but one fraught with well-known perils (Nichols & Conroy, 1996; Yoccoz, Nichols & Boulinier, 2001; Pollock et al., 2002). To establish the utility of an index as a substitute for an estimate of density, one must: (1) demonstrate a functional relationship between the index and density that is invariant over the desired scope of inference; (2) calibrate the functional relationship by obtaining independent measures of the index and the animal density; (3) evaluate the precision of the calibration (Diefenbach et al., 1994). Carbone et al. (2001) argue that the number of camera-days per photograph is a useful index of density for large, cryptic, forest-dwelling animals, and proceed to calibrate this index for tigers (Panthera tigris). We agree that a properly calibrated index may be useful for rapid assessments in conservation planning. However, Carbone et al. (2001), who desire to use their index as a substitute for density, do not adequately address the three elements noted above. Thus, we are concerned that others may view their methods as justification for not attempting directly to estimate animal densities, without due regard for the shortcomings of their approach.

Animal Conservation

Occupancy Estimation and Modeling : Inferring Patterns and Dynamics of Species Occurrence

This is the first book to examine the latest methods in analyzing presence/absence data surveys. Using four classes of models (single-species, single-season; single-species, multiple season; multiple-species, single-season; and multiple-species, multiple-season), the authors discuss the practical sampling situation, present a likelihood-based model enabling direct estimation of the occupancy-related parameters while allowing for imperfect detectability, and make recommendations for designing studies using these models. It provides authoritative insights into the latest in estimation modeling; discusses multiple models which lay the groundwork for future study designs; addresses critical issues of imperfect detectibility and its effects on estimation; and explores the role of probability in estimating in detail.

Book

Investigating species co-occurrence patterns when species are detected imperfectly

1. Over the last 30 years there has been a great deal of interest in investigating patterns of species co-occurrence across a number of locations, which has led to the development of numerous methods to determine whether there is evidence that a particular pattern may not have occurred by random chance. 2. A key aspect that seems to have been largely overlooked is the possibility that species may not always be detected at a location when present, which leads to 'false absences' in a species presence/absence matrix that may cause incorrect inferences to be made about co-occurrence patterns. Furthermore, many of the published methods for investigating patterns of species co-occurrence do not account for potential differences in the site characteristics that may partially (at least) explain non-random patterns (e.g. due to species having similar/different habitat preferences). 3. Here we present a statistical method for modelling co-occurrence patterns between species while accounting for imperfect detection and site characteristics. This method requires that multiple presence/absence surveys for the species be conducted over a reasonably short period of time at most sites. The method yields unbiased estimates of probabilities of occurrence, and is practical when the number of species is small (< 4). 4. To illustrate the method we consider data collected on two terrestrial salamander species, Plethodon jordani and members of the Plethodon glutinosus complex, collected in the Great Smoky Mountains National Park, USA. We find no evidence that the species do not occur independently at sites once site elevation has been allowed for, although we find some evidence of a statistical interaction between species in terms of detectability that we suggest may be due to changes in relative abundances.

North Carolina, Tennessee

Assessing the fit of site-occupancy models

Few species are likely to be so evident that they will always be detected at a site when present. Recently a model has been developed that enables estimation of the proportion of area occupied, when the target species is not detected with certainty. Here we apply this modeling approach to data collected on terrestrial salamanders in the Plethodon glutinosus complex in the Great Smoky Mountains National Park, USA, and wish to address the question 'how accurately does the fitted model represent the data?' The goodness-of-fit of the model needs to be assessed in order to make accurate inferences. This article presents a method where a simple Pearson chi-square statistic is calculated and a parametric bootstrap procedure is used to determine whether the observed statistic is unusually large. We found evidence that the most global model considered provides a poor fit to the data, hence estimated an overdispersion factor to adjust model selection procedures and inflate standard errors. Two hypothetical datasets with known assumption violations are also analyzed, illustrating that the method may be used to guide researchers to making appropriate inferences. The results of a simulation study are presented to provide a broader view of the methods properties.

North Carolina, Tennessee

Abundance estimation and conservation biology

Abundance is the state variable of interest in most population–level ecological research and in most programs involving management and conservation of animal populations. Abundance is the single parameter of interest in capture–recapture models for closed populations (e.g., Darroch, 1958; Otis et al., 1978; Chao, 2001). The initial capture–recapture models developed for partially (Darroch, 1959) and completely (Jolly, 1965; Seber, 1965) open populations represented efforts to relax the restrictive assumption of population closure for the purpose of estimating abundance. Subsequent emphases in capture–recapture work were on survival rate estimation in the 1970’s and 1980’s (e.g., Burnham et al., 1987; Lebreton et al.,1992), and on movement estimation in the 1990’s (Brownie et al., 1993; Schwarz et al., 1993). However, from the mid–1990’s until the present time, capture–recapture investigators have expressed a renewed interest in abundance and related parameters (Pradel, 1996; Schwarz & Arnason, 1996; Schwarz, 2001). The focus of this session was abundance, and presentations covered topics ranging from estimation of abundance and rate of change in abundance, to inferences about the demographic processes underlying changes in abundance, to occupancy as a surrogate of abundance. The plenary paper by Link & Barker (2004) is provocative and very interesting, and it contains a number of important messages and suggestions. Link & Barker (2004) emphasize that the increasing complexity of capture–recapture models has resulted in large numbers of parameters and that a challenge to ecologists is to extract ecological signals from this complexity. They offer hierarchical models as a natural approach to inference in which traditional parameters are viewed as realizations of stochastic processes. These processes are governed by hyperparameters, and the inferential approach focuses on these hyperparameters. Link & Barker (2004) also suggest that our attention should be focused on relationships between demographic processes such as survival and recruitment, the two quantities responsible for changes in abundance, rather than simply on the magnitudes of these quantities. They describe a type of Jolly–Seber capture–recapture model that permits inference about the underlying relationship between per capita recruitment rates and survival rates (Link & Barker, this volume). Implementation used Bayesian Markov Chain Monte Carlo methods and appeared to work well, yielding inferences about the relationship between recruitment and survival that were robust to selection of prior distribution. We believe that readers will find their arguments compelling, and we expect to see increased use of hierarchical modeling approaches in capture–recapture and related fields. Otto (presentation without paper) also recommended use of hierarchical models in analysis of multiple data sources dealing with population dynamics of North American mallards. He integrated survival inferences from ringing data, abundance information from aerial survey data, and recruitment information based on age ratios from a harvest survey. He used a Leslie matrix population projection model as an integrating framework and obtained estimates of breeding population size using all data.Otto’s approach also permitted inference about biases in estimated quantities. As with the work of Link & Barker (2004), we find Otto’s recommendation to use hierarchical models to integrate data from multiple sources to be very compelling. Alisauskas et al. (2004) report results of an analysis of capture–recapture data for a askatchewan population of white–winged scoters. They used the approach of Pradel (1996) to estimate population growth rate (See the PDF) directly. Estimates for 1975–1985 were quite low, but estimates for the recent period, 2000–2003,increased to values > 1. Parameter estimates for seniority, survival and per capita recruitment (Pradel, 1996) led to the inference that increased recruitment was largely responsible for the improvements in population status and growth. However, various data sources also indicated that this increase in recruitment was likely a result of increased immigration rather than improved reproduction on the area. This latter inference is important from a conservation perspective in indicating the importance of birds in other locations to growth and health of the study population. Lukacs and Burnham presented material to be published elsewhere that dealt with the use of genetic markers in capture–recapture studies. The data sources for such studies are samples of hair or feces, which are then analyzed using molecular genetic techniques in order to determine individual genotypes with respect to a usually small number of loci. Two types of classification error can arise in such analyses. First, if only a small number of loci is examined, then there may be nonnegligible probabilities that multiple individual animals will have the same genotypes. The second type of error arises during the polymerase chain reaction (PCR) process and can result from failure of alleles to amplify (allelic dropout) or from PCR inhibitors in hair and feces that produce the appearance of false alleles or misprinting (Creel et al., 2003). Lukacs and Burnham developed models that formally incorporate possible misclassification of samples resulting from these errors. These models permit estimation of parameters such as abundance and survival in a manner that properly incorporates this uncertainty of individual identity. We anticipate that noninvasive sampling based on molecular genetic analyses of hair or feces will become extremely important for some species, and that the models of Lukacs and Burnham will become very popular for such analyses. MacKenzie & Nichols (2004) discuss the use of occupancy (proportion of patches or habitat area that is occupied) as a surrogate for abundance. In cases of territorial species and where birds occur at low densities, the number of occupied patches may provide a reasonable estimate of abundance. In other cases, occupancy can be viewed as providing information about one tail of the abundance distribution, P (N = 0). The motivation for considering occupancy as a surrogate for abundance is that occupancy is based on so–called presence–absence surveys that are frequently less expensive of time and effort than methods that estimate abundance directly. We describe one set of models that can be used to estimate occupancy for a single season and another that can be used to estimate parameters such as local probabilities of extinction and colonization that are associated with occupancy dynamics. We outline a possible hybrid approach that combines occupancy data with data on marked individuals in order to betterexplore the mechanisms underlying occupancy dynamics. These five presentations made for an interesting session containing useful information and recommendations for future work. A number of themes connecting these presentations could be emphasized. For example, two of the presentations considered alternatives to standard capture–recapture sampling that can be used to draw inferences about abundance, or a portion of the abundance distribution, with field methods that should be less expensive than usual capture–recapture approaches of handling animals. We believe that the most important theme of the session was the emphasis on the processes responsible for changes in abundance. In particular, we are excited by the potential for using hierarchical models as a means of investigating relationships among vital rates and as a means of combining multiple sources of data relevant to system dynamics. Indeed, we expect the importance of this session theme to be reflected in the content and presentations of the next EURING meeting.

Animal Biodiversity and Conservation

Occupancy as a surrogate for abundance estimation

In many monitoring programmes it may be prohibitively expensive to estimate the actual abundance of a bird species in a defined area, particularly at large spatial scales, or where birds occur at very low densities. Often it may be appropriate to consider the proportion of area occupied by the species as an alternative state variable. However, as with abundance estimation, issues of detectability must be taken into account in order to make accurate inferences: the non?detection of the species does not imply the species is genuinely absent. Here we review some recent modelling developments that permit unbiased estimation of the proportion of area occupied, colonization and local extinction probabilities. These methods allow for unequal sampling effort and enable covariate information on sampling locations to be incorporated. We also describe how these models could be extended to incorporate information from marked individuals, which would enable finer questions of population dynamics (such as turnover rate of nest sites by specific breeding pairs) to be addressed. We believe these models may be applicable to a wide range of bird species and may be useful for investigating various questions of ecological interest. For example, with respect to habitat quality, we might predict that a species is more likely to have higher local extinction probabilities, or higher turnover rates of specific breeding pairs, in poor quality habitats.

Animal Biodiversity and Conservation

Estimating site occupancy, colonization, and local extinction when a species is detected imperfectly

Few species are likely to be so evident that they will always be detected when present. Failing to allow for the possibility that a target species was present, but undetected, at a site will lead to biased estimates of site occupancy, colonization, and local extinction probabilities. These population vital rates are often of interest in long-term monitoring programs and metapopulation studies. We present a model that enables direct estimation of these parameters when the probability of detecting the species is less than 1. The model does not require any assumptions of process stationarity, as do some previous methods, but does require detection/nondetection data to be collected in a manner similar to Pollock's robust design as used in mark?recapture studies. Via simulation, we show that the model provides good estimates of parameters for most scenarios considered. We illustrate the method with data from monitoring programs of Northern Spotted Owls (Strix occidentalis caurina) in northern California and tiger salamanders (Ambystoma tigrinum) in Minnesota, USA.

Ecology

Estimating site occupancy rates when detection probabilities are less than one

Nondetection of a species at a site does not imply that the species is absent unless the probability of detection is 1. We propose a model and likelihood-based method for estimating site occupancy rates when detection probabilities are < 1. The model provides a flexible framework enabling covariate information to be included and allowing for missing observations. Via computer simulation, we found that the model provides good estimates of the occupancy rates, generally unbiased for moderate detection probabilities (>0.3). We estimated site occupancy rates for two anuran species at 32 wetland sites in Maryland, USA, from data collected during 2000 as part of an amphibian monitoring program, Frogwatch USA. Site occupancy rates were estimated as 0.49 for American toads ( Bufo americanus ), a 44% increase over the proportion of sites at which they were actually observed, and as 0.85 for spring peepers ( Pseudacris crucifer ), slightly above the observed proportion of 0.83.

Maryland

How should detection probability be incorporated into estimates of relative abundance?

Determination of the relative abundance of two populations, separated by time or space, is of interest in many ecological situations. We focus on two estimators of relative abundance, which assume that the probability that an individual is detected at least once in the survey is either equal or unequal for the two populations. We present three methods for incorporating the collected information into our inference. The first method, proposed previously, is a traditional hypothesis test for evidence that detection probabilities are unequal. However, we feel that, a priori, it is more likely that detection probabilities are actually different; hence, the burden of proof should be shifted, requiring evidence that detection probabilities are practically equivalent. The second method we present, equivalence testing, is one approach to doing so. Third, we suggest that model averaging could be used by combining the two estimators according to derived model weights. These differing approaches are applied to a mark-recapture experiment on Nuttail's cottontail rabbit ( Sylvilagus nuttallii ) conducted in central Oregon during 1974 and 1975, which has been previously analyzed by other authors.

Oregon