MARK provides parameter estimates from marked animals when they are re-encountered at a later time as dead recoveries, or live recaptures or re-sightings. The time intervals between re-encounters do not have to be equal. More than one attribute group of animals can be modelled. The basic input to MARK is the encounter history for each animal. MARK can also estimate the size of closed populations. Parameters can be constrained to be the same across re-encounter occasions, or by age, or group, using the parameter index matrix. A set of common models for initial screening of data are provided. Time effects, group effects, time x group effects and a null model of none of the above, are provided for each parameter. Besides the logit function to link the design matrix to the parameters of the model, other link functions include the log—log, complimentary log—log, sine, log, and identity. The estimates of model parameters are computed via numerical maximum likelihood techniques. The number of parameters that are...

https://www.tandfonline.com/doi/pdf/10.1080/00063659909477239

Program MARK: survival estimation from populations of marked animals

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 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.

/pdf/estimating-site-occupancy-rates-when-detection-probabilities-3listkvns9.pdf

Estimating site occupancy rates when detection probabilities are less than one

Beginning with Emlen (1966) and MacArthur and Pianka (1966) and extending through the last ten years, several authors have sought to predict the foraging behavior of animals by means of mathematical models. These models are very similar,in that they all assume that the fitness of a foraging animal is a function of the efficiency of foraging measured in terms of some "currency" (Schoener, 1971) -usually energy- and that natural selection has resulted in animals that forage so as to maximize this fitness. As a result of these similarities, the models have become known as "optimal foraging models"; and the theory that embodies them, "optimal foraging theory." The situations to which optimal foraging theory has been applied, with the exception of a few recent studies, can be divided into the following four categories: (1) choice by an animal of which food types to eat (i.e., optimal diet); (2) choice of which patch type to feed in (i.e., optimal patch choice); (3) optimal allocation of time to different patches; and (4) optimal patterns and speed of movements. In this review we discuss each of these categories separately, dealing with both the theoretical developments and the data that permit tests of the predictions. The review is selective in the sense that we emphasize studies that either develop testable predictions or that attempt to test predictions in a precise quantitative manner. We also discuss what we see to be some of the future developments in the area of optimal foraging theory and how this theory can be related to other areas of biology. Our general conclusion is that the simple models so far formulated are supported are supported reasonably well by available data and that we are optimistic about the value both now and in the future of optimal foraging theory. We argue, however, that these simple models will requre much modification, espicially to deal with situations that either cannot easily be put into one or another of the above four categories or entail currencies more complicated that just energy.

Citation classic - optimal foraging - a selective review of theory and tests

Information theoretic approaches and model averaging are increasing in popularity, but this approach can be difficult to apply to the realistic, complex models that typify many ecological and evolutionary analyses. This is especially true for those researchers without a formal background in information theory. Here, we highlight a number of practical obstacles to model averaging complex models. Although not meant to be an exhaustive review, we identify several important issues with tentative solutions where they exist (e.g. dealing with collinearity amongst predictors; how to compute model-averaged parameters) and highlight areas for future research where solutions are not clear (e.g. when to use random intercepts or slopes; which information criteria to use when random factors are involved). We also provide a worked example of a mixed model analysis of inbreeding depression in a wild population. By providing an overview of these issues, we hope that this approach will become more accessible to those investigating any process where multiple variables impact an evolutionary or ecological response.

https://www.otago.ac.nz/threatenedbirdgroup/Publications_files/Grueber_et%20al%20JEB_2011_incl%20Corrigendum.pdf

Multimodel inference in ecology and evolution: challenges and solutions

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 occiden- talis caurina) in northern California and tiger salamanders (Ambystoma tigrinum) in Min- nesota, USA.

/pdf/estimating-site-occupancy-colonization-and-local-extinction-8s715obrby.pdf

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

Statistical inference for capture-recapture studies of open animal populations typically relies on the assumption that all emigration from the studied population is per- manent. However, there are many instances in which this assumption is unlikely to be met. We define two general models for the process of temporary emigration: completely random and Markovian. We then consider effects of these two types of temporary emigration on Jolly-Seber estimators and on estimators arising from the full-likelihood approach to robust design data. Capture-recapture data arising from Pollock's robust design provide the basis for ob- taining unbiased estimates of demographic parameters in the presence of temporary emi- gration, and for estimating the probability of temporary emigration. We present a likelihood- based approach to dealing with temporary emigration that permits estimation under different models of temporary emigration and yields tests for completely random and Markovian emigration. In addition, we use the relationship between capture probability estimates based on closed and open models under completely random temporary emigration to derive three ad hoc estimators for the probability of temporary emigration. Two of these should be especially useful in situations where capture probabilities are heterogeneous among indi- vidual animals. Ad hoc and full-likelihood estimators are illustrated for small-mammal capture-recapture data sets. We believe that these models and estimators will be useful for testing hypotheses about the process of temporary emigration, for estimating demographic parameters in the presence of temporary emigration, and for estimating probabilities of temporary emigration. These latter estimates are frequently of ecological interest as indicators of animal movement and, in some sampling situations, as direct estimates of breeding probabilities and proportions.

Estimating temporary emigration using capture-recapture data with pollock's robust design

INTRODUCTION In 1938, Howard M. Wight devoted 9 pages, which was an entire chapter in the first wildlife management techniques manual, to what he termed 'census' methods. As books and chapters such as this attest, the volume of literature on this subject has grown tremendously. Abundance estimation remains an active area of biometrical research, as reflected in the many differences between this chapter and the similar contribution in the previous manual. Our intent in this chapter is to present an overview of the basic and most widely used population estimation techniques and to provide an entree to the relevant literature. Several possible approaches could be taken in writing a chapter dealing with population estimation. For example, we could provide a detailed treatment focusing on statistical models and on derivation of estimators based on these models. Although a chapter using this approach might provide a valuable reference for quantitative biologists and biometricians, it would be of limited use to many field biologists and wildlife managers. Another approach would be to focus on details of actually applying different population estimation techniques. This approach would include both field application (e.g., how to set out a trapping grid or conduct an aerial survey) and detailed instructions on how to use the resulting data with appropriate estimation equations. We are reluctant to attempt such an approach, however, because of the tremendous diversity of real-world field situations defined by factors such as the animal being studied, habitat, available resources, and because of our resultant inability to provide detailed instructions for all possible cases. We believe it is more useful to provide the reader with the conceptual basis underlying estimation methods. Thus, we have tried to provide intuitive explanations for how basic methods work. In doing so, we present relevant estimation equations for many methods and provide citations of more detailed treatments covering both statistical considerations and field applications. We have chosen to present methods that are representative of classes of estimators, rather than address every available method. Our hope is that this chapter will provide the reader with enough background to make an informed decision about what general method(s) will likely perform well in any particular field situation. Readers with a more quantitative background may then be able to consult detailed references and tailor the selected method to suit their particular needs. Less quantitative readers should consult a biometrician, preferably one with experience in wildlife studies, for this 'tailoring,' with the hope they will be able to do so with a basic understanding of the general method, thereby permitting useful interaction and discussion with the biometrician. SUMMARY Estimating the abundance or density of animals in wild populations is not a trivial matter. Virtually all techniques involve the basic problem of estimating the probability of seeing, capturing, or otherwise detecting animals during some type of survey and, in many cases, sampling concerns as well. In the case of indices, the detection probability is assumed to be constant (but unknown). We caution against use of indices unless this assumption can be verified for the comparison(s) of interest. In the case of population estimation, many methods have been developed over the years to estimate the probability of detection associated with various kinds of count statistics. Techniques range from complete counts, where sampling concerns often dominate, to incomplete counts where detection probabilities are also important. Some examples of the latter are multiple observers, removal methods, and capture-recapture. Before embarking on a survey to estimate the size of a population, one must understand clearly what information is needed and for what purpose the information will be used. The key to derivin Additional publication details Publication type Book chapter Publication Subtype Book Chapter Here's how you know Estimating the numb r of animals in wildlife populations Title Estimating the number of animals in wildlife populations Year Published 2005 Language English Publisher Wildlife Society Publisher location Bethesda, Maryland Contributing office(s) Patuxent Wildlife Research Center Description xiv, 974 Larger Work Type Book Larger Work Subtype Other Government Series Larger Work Title Techniques for Wildlife Investigations and Management

Estimating the number of animals in wildlife populations

The Jolly-Seber method has been the traditional approach to the estimation of demographic parameters in long-term capture-recapture studies of wildlife and fish species. This method involves restrictive assumptions about capture probabilities that can lead to biased estimates, especially of population size and recruitment. Pollock (1982, Journal of Wildlife Management 46, 752-757) proposed a sampling scheme in which a series of closely spaced samples were separated by longer intervals such as a year. For this "robust design," Pollock suggested a flexible ad hoc approach that combines the Jolly-Seber estimators with closed population estimators, to reduce bias caused by unequal catchability, and to provide estimates for parameters that are unidentifiable by the Jolly-Seber method alone. In this paper we provide a formal modelling framework for analysis of data obtained using the robust design. We develop likelihood functions for the complete data structure under a variety of models and examine the relationship among the models. We compute maximum likelihood estimates for the parameters by applying a conditional argument, and compare their performance against those of ad hoc and Jolly-Seber approaches using simulation.

A likelihood-based approach to capture-recapture estimation of demographic parameters under the robust design.

Closed-population capture-recapture methods have been used extensively in animal ecology, both by themselves and within the context of Pollock's robust design and multistate models, to estimate various parameters of population and community dynamics. The defining assumption of geographic and demographic closure (i.e., no births, deaths, immigration, or emigration) for the duration of sampling is restrictive and is likely to be violated in many field situations. I evaluated several types of violations of the closure assumption and found that completely random movement in and out of a study area does not introduce bias to estimators from closed-population methods, although it decreases precision. In addition, if capture probabilities vary only with time, the closed-population Lincoln-Petersen estimator is unbiased for the size of the superpopulation when there are only births/immigration or only deaths/emigration. However, for other cases of nonrandom movement, closed-population estimators were biased when movement was Markovian (de- pendent on the presence/absence of the animal in the previous time period), when an animal was allowed one entry to and one exit from the study area, or when there was trap response or heterogeneity among animals in capture probability. In addition, the probability that an animal is present and available for capture (e.g., breeding propensity) can be estimated using Pollock's robust design only when movement occurs at a broader temporal scale than that of sampling.

Robustness of closed capture-recapture methods to violations of the closure assumption

Detecting individuals of amphibian and reptile species can be a daunting task. Detection can be hindered by various factors such as cryptic behavior, color patterns, or observer experience. These factors complicate the estimation of state variables of interest (e.g., abundance, occupancy, species richness) as well as the vital rates that induce changes in these state variables (e.g., survival probabilities for abundance; extinction probabilities for occupancy). Although ad hoc methods (e.g., counts uncorrected for detection, return rates) typically perform poorly in the face of no detection, they continue to be used extensively in various fields, including herpetology. However, formal approaches that estimate and account for the probability of detection, such as capture-mark-recapture (CMR) methods and distance sampling, are available. In this paper, we present classical approaches and recent advances in methods accounting for detectability that are particularly pertinent for herpetological data sets. Through examples, we illustrate the use of several methods, discuss their performance compared to that of ad hoc methods, and we suggest available software to perform these analyses. The methods we discuss control for imperfect detection and reduce bias in estimates of demographic parameters such as population size, survival, or, at other levels of biological organization, species occurrence. Among these methods, recently developed approaches that no longer require marked or resighted individuals should be particularly of interest to field herpetologists. We hope that our effort will encourage practitioners to implement some of the estimation methods presented herein instead of relying on ad hoc methods that make more limiting assumptions.

William L. Kendall

Papers

Estimating temporary emigration using capture-recapture data with pollock's robust design

Estimating the number of animals in wildlife populations

A likelihood-based approach to capture-recapture estimation of demographic parameters under the robust design.

Robustness of closed capture-recapture methods to violations of the closure assumption

Making Great Leaps Forward: Accounting for Detectability in Herpetological Field Studies