James E. Hines

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 Pollocks 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.

ESTIMATING TEMPORARY EMIGRATION USING CAPTURE–RECAPTURE DATA WITH POLLOCK’S ROBUST DESIGN

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 Pollocks 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.

A double-observer approach for estimating detection probability and abundance from point counts

Abstract Although point counts are frequently used in ornithological studies, basic assumptions about detection probabilities often are untested. We apply a double-observer approach developed to estimate detection probabilities for aerial surveys (Cook and Jacobson 1979) to avian point counts. At each point count, a designated “primary” observer indicates to another (“secondary”) observer all birds detected. The secondary observer records all detections of the primary observer as well as any birds not detected by the primary observer. Observers alternate primary and secondary roles during the course of the survey. The approach permits estimation of observer-specific detection probabilities and bird abundance. We developed a set of models that incorporate different assumptions about sources of variation (e.g. observer, bird species) in detection probability. Seventeen field trials were conducted, and models were fit to the resulting data using program SURVIV. Single-observer point counts generally miss varying proportions of the birds actually present, and observer and bird species were found to be relevant sources of variation in detection probabilities. Overall detection probabilities (probability of being detected by at least one of the two observers) estimated using the double-observer approach were very high (>0.95), yielding precise estimates of avian abundance. We consider problems with the approach and recommend possible solutions, including restriction of the approach to fixed-radius counts to reduce the effect of variation in the effective radius of detection among various observers and to provide a basis for using spatial sampling to estimate bird abundance on large areas of interest. We believe that most questions meriting the effort required to carry out point counts also merit serious attempts to estimate detection probabilities associated with the counts. The double-observer approach is a method that can be used for this purpose.

ESTIMATING SPECIES RICHNESS: THE IMPORTANCE OF HETEROGENEITY IN SPECIES DETECTABILITY

Estimating species richness (i.e., the actual number of species present in a given area) is a basic objective of many field studies carried out in community ecology and is also of crucial concern when dealing with the conservation and management of biodiversity. In most studies, the total number of species recorded in an area at a given time is taken as a measure of species richness. Here we use a capture–recapture approach to species richness estimation with North American Breeding Bird Survey (BBS) data in order to estimate species detectability and thus gain insight about its importance. In particular, competing models making different assumptions about species detectability are available. We carried out analyses on all survey routes of four states, Arizona, Maryland, North Dakota, and Wisconsin, in two years, 1970 and 1990. These states were chosen to provide contrasting habitats, bird species composition, and survey quality. We investigated the effect of state, year, and observer ability on the propo...

A REMOVAL MODEL FOR ESTIMATING DETECTION PROBABILITIES FROM POINT-COUNT SURVEYS

Abstract Use of point-count surveys is a popular method for collecting data on abundance and distribution of birds. However, analyses of such data often ignore potential differences in detection probability. We adapted a removal model to directly estimate detection probability during point-count surveys. The model assumes that singing frequency is a major factor influencing probability of detection when birds are surveyed using point counts. This may be appropriate for surveys in which most detections are by sound. The model requires counts to be divided into several time intervals. Point counts are often conducted for 10 min, where the number of birds recorded is divided into those first observed in the first 3 min, the subsequent 2 min, and the last 5 min. We developed a maximum-likelihood estimator for the detectability of birds recorded during counts divided into those intervals. This technique can easily be adapted to point counts divided into intervals of any length. We applied this method to unlimited-radius counts conducted in Great Smoky Mountains National Park. We used model selection criteria to identify whether detection probabilities varied among species, throughout the morning, throughout the season, and among different observers. We found differences in detection probability among species. Species that sing frequently such as Winter Wren (Troglodytes troglodytes) and Acadian Flycatcher (Empidonax virescens) had high detection probabilities (∼90%) and species that call infrequently such as Pileated Woodpecker (Dryocopus pileatus) had low detection probability (36%). We also found detection probabilities varied with the time of day for some species (e.g. thrushes) and between observers for other species. We used the same approach to estimate detection probability and density for a subset of the observations with limited-radius point counts.

Estimating Breeding Proportions and Testing Hypotheses about Costs of Reproduction with Capture-Recapture Data

The proportion of animals in a population that breeds is an important determinant of population growth rate. Usual estimates of this quantity from field sampling data assume that the probability of appearing in the capture or count statistic is the same for animals that do and do not breed. A similar assumption is required by most existing methods used to test ecologically interesting hypotheses about reproductive costs using field sampling data. However, in many field sampling situations breeding and nonbreeding animals are likely to exhibit different probabilities of being seen or caught. In this paper, we propose the use of multistate capture-recapture models for these estimation and testing problems. This methodology permits a formal test of the hypothesis of equal capture/ sighting probabilities for breeding and nonbreeding individuals. Two estimators of breeding proportion (and associated standard errors) are presented, one for the case of equal capture probabilities and one for the case of unequal capture probabilities. The multistate modeling framework also yields formal tests of hypotheses about reproductive costs to future repro- duction or survival or both fitness components. The general methodology is illustrated using capture-recapture data on female meadow voles, Microtus pennsylvanicus. Resulting estimates of the proportion of reproductively active females showed strong seasonal vari- ation, as expected, with low breeding proportions in midwinter. We found no evidence of reproductive costs extracted in subsequent survival or reproduction. We believe that this methodological framework has wide application to problems in animal ecology concerning breeding proportions and phenotypic reproductive costs.

ARE ADULT NONBREEDERS PRUDENT PARENTS? THE KITTIWAKE MODEL

Understanding evolutionary consequences of intermittent breeding (non- breeding in individuals that previously bred) requires investigation of the relationships between adult breeding state and two demographic parameters: survival probability and subsequent breeding probability. One major difficulty raised by comparing the demographic features of breeders and nonbreeders as estimated from capture-recapture data is that breed- ing state is often suspected to influence recapture or resighting probability. We used mul- tistate capture-recapture models to test the hypothesis of equal recapture probabilities for breeding and nonbreeding Kittiwakes and found no evidence of an effect of breeding state on this parameter. The same method was used to test whether reproductive state affects survival probability. Nonbreeding individuals have lower survival rates than breeders. Moreover, nonbreeders have a higher probability of being nonbreeders the following year than do breeders. State-specific survival rates and transition probabilities vary from year to year, but temporal variations of survival and transition probabilities of breeders and nonbreeders are in parallel (on a logit scale). These inferences led us to conclude that nonbreeders tend to be lower quality individuals. The effect of sex was also investigated: males and females do not differ with respect to survival probabilities when reproductive state is taken into account. Similarly, there is no effect of sex on transition probabilities between reproductive states.

Estimating Annual Survival and Movement Rates of Adults within a Metapopulation of Roseate Terns

Several multistratum capture-recapture models were used to test various hypotheses about possible geographic and temporal variation in survival, movement, and recapture/resighting probabilities of 2399 adult Roseate Terns (Sterna dougallii) color-banded from 1988 to 1992 at the sites of the four largest breeding colonies of this species in the northeastern USA. Linear-logistic ultrastructural models also were developed to investigate possible correlates of geographic variation in movement probabilities. Based on goodness-of-fit tests and comparisons of Akaikes Information Criterion (AIC) values, the fully parameterized model (Model A) with time- and location-specific survival, movement, and capture probabilities, was selected as the most appropriate model for this metapopulation structure. With almost all movement accounted for, on average >90% of the surviving adults from each colony site returned to the same site the following year. Variations in movement probabilities were more closely associated with the identity of the destination colony site than with either the identity of the colony site of origin or the distance between colony sites. The average annual survival estimates (0.74-0.84) of terns from all four sites indicate a high rate of annual mortality relative to that of other species of marine birds.

Forest fragmentation and bird community dynamics: inference at regional scales

With increasing fragmentation of natural areas and a dramatic reduction of forest cover in several parts of the world, quantifying the impact of such changes on species richness and community dynamics has been a subject of much concern. Here, we tested whether in more fragmented landscapes there was a lower number of area-sensitive species and higher local extinction and turnover rates, which could explain higher temporal vari- ability in species richness. To investigate such potential landscape effects at a regional scale, we merged two independent, large-scale monitoring efforts: the North American Breeding Bird Survey (BBS) and the Land Use and Land Cover Classification data from the U.S. Geological Survey. We used methods that accounted for heterogeneity in the probability of detecting species to estimate species richness and temporal changes in the bird communities for BBS routes in three mid-Atlantic U.S. states. Forest breeding bird species were grouped prior to the analyses into area-sensitive and non-area-sensitive species according to previous studies. We tested predictions relating measures of forest structure at one point in time (1974) to species richness at that time and to parameters of forest bird community change over the following 22-yr-period (1975-1996). We used the mean size of forest patches to characterize landscape structure, as high correlations among landscape variables did not allow us to disentangle the relative roles of habitat fragmentation per se and habitat loss. As predicted, together with lower species richness for area-sensitive species on routes surrounded by landscapes with lower mean forest-patch size, we found higher mean year- to-year rates of local extinction. Moreover, the mean year-to-year rates of local turnover (proportion of locally new species) for area-sensitive species were also higher in landscapes with lower mean forest-patch size. These associations were not observed for the non-area- sensitive species group. These results suggest that landscape structure may influence forest bird communities at regional scales through its effects on the total number of species but also on the temporal rates of change in community composition. Evidence for higher rates of local extinction and turnover in more fragmented landscapes suggests that bird communities function as metapopulations at a regional scale, and points out the importance of colonizations and recolonizations from surrounding landscapes to local community dynamics. Further, our results illustrate that the methods used to estimate the community parameters can be a powerful statistical tool in addressing questions relative to the dynamics of communities.

SAMPLING DESIGN TRADE‐OFFS IN OCCUPANCY STUDIES WITH IMPERFECT DETECTION: EXAMPLES AND SOFTWARE

Researchers have used occupancy, or probability of occupancy, as a response or state variable in a variety of studies (e.g., habitat modeling), and occupancy is increasingly favored by numerous state, federal, and international agencies engaged in monitoring programs. Recent advances in estimation methods have emphasized that reliable inferences can be made from these types of studies if detection and occupancy probabilities are simultaneously estimated. The need for temporal replication at sampled sites to estimate detection probability creates a trade-off between spatial replication (number of sample sites distributed within the area of interest/inference) and temporal replication (number of repeated surveys at each site). Here, we discuss a suite of questions commonly encountered during the design phase of occupancy studies, and we describe software (program GENPRES) developed to allow investigators to easily explore design trade-offs focused on particularities of their study system and sampling limitations. We illustrate the utility of program GENPRES using an amphibian example from Greater Yellowstone National Park, U.S.A.