Conditional and unconditional estimators of population size

Estimators for the population sizes of animal species are similar to Horvitz-Thompson estimators — they involve dividing counts of detected animals by the probabilities of detection. Knowing detection probabilities for different subpopulations allows one to estimate each sub-population size with such an estimator, and to add the results for an estimator of the total population. In the case where the proportions of animals belonging to the different sub-populations are also known, this paper shows that using those proportions to arrive at a common average detection probability will, when used in conjunction with the total number of animals detected, result in a better estimator. We provide two examples where the inferior estimator may seem sensible.     

Estimating Population Size in a Continuous-time Removal Experiment with a Known Sub-population Size Ratio

We study a continuous-time removal model for estimating the population size for a population in which a sub-population size ratio is known. The maximum likelihood estimate and the optimal martingale estimate of the population size are obtained; these are shown to be equivalent. A comparison between the proposed estimator and the maximum likelihood estimate which ignores the information on the known size ratio is made, using a simulation study. The asymptotic variances of these two estimators are also obtained, and a comparison between them is made. The sensitivity of mis-specification of the known size ratio is examined. We also apply the corresponding discrete-time model to the proposed continuous-time setting, and study the efficiency of the corresponding discrete-time type estimator relative to the proposed estimator.     

Multiple data sources improve DNA-based mark-recapture population estimates of grizzly bears.

A fundamental challenge to estimating population size with mark-recapture methods is heterogeneous capture probabilities and subsequent bias of population estimates. Confronting this problem usually requires substantial sampling effort that can be difficult to achieve for some species, such as carnivores. We developed a methodology that uses two data sources to deal with heterogeneity and applied this to DNA mark-recapture data from grizzly bears (Ursus arctos). We improved population estimates by incorporating additional DNA "captures" of grizzly bears obtained by collecting hair from unbaited bear rub trees concurrently with baited, grid-based, hair snag sampling. We consider a Lincoln-Petersen estimator with hair snag captures as the initial session and rub tree captures as the recapture session and develop an estimator in program MARK that treats hair snag and rub tree samples as successive sessions. Using empirical data from a large-scale project in the greater Glacier National Park, Montana, USA, area and simulation modeling we evaluate these methods and compare the results to hair-snag-only estimates. Empirical results indicate that, compared with hair-snag-only data, the joint hair-snag-rub-tree methods produce similar but more precise estimates if capture and recapture rates are reasonably high for both methods. Simulation results suggest that estimators are potentially affected by correlation of capture probabilities between sample types in the presence of heterogeneity. Overall, closed population Huggins-Pledger estimators showed the highest precision and were most robust to sparse data, heterogeneity, and capture probability correlation among sampling types. Results also indicate that these estimators can be used when a segment of the population has zero capture probability for one of the methods. We propose that this general methodology may be useful for other species in which mark-recapture data are available from multiple sources.     

Modeling misidentification errors that result from use of genetic tags in capture–recapture studies

Misidentification of animals is potentially important when naturally existing features (natural tags) such as DNA fingerprints (genetic tags) are used to identify individual animals. For example, when misidentification leads to multiple identities being assigned to an animal, traditional estimators tend to overestimate population size. Accounting for misidentification in capture–recapture models requires detailed understanding of the mechanism. Using genetic tags as an example, we outline a framework for modeling the effect of misidentification in closed population studies when individual identification is based on natural tags that are consistent over time (non-evolving natural tags). We first assume a single sample is obtained per animal for each capture event, and then generalize to the case where multiple samples (such as hair or scat samples) are collected per animal per capture occasion. We introduce methods for estimating population size and, using a simulation study, we show that our new estimators perform well for cases with moderately high capture probabilities or high misidentification rates. In contrast, conventional estimators can seriously overestimate population size when errors due to misidentification are ignored.     

Comparison of distribution function estimators for forestry applications following variable probability sampling

Practical considerations often motivate employing variable probability sampling designs when estimating characteristics of forest populations. Three distribution function estimators, the Horvitz-Thompson estimator, a difference estimator, and a ratio estimator, are compared following variable probability sampling in which the inclusion probabilities are proportional to an auxiliary variable, X. Relative performance of the estimators is affected by several factors, including the distribution of the inclusion probabilities, the correlation () between X and the response Y, and the position along the distribution function being estimated. Both the ratio and difference estimators are superior to the Horvitz-Thompson estimator. The difference estimator gains better precision than the ratio estimator toward the upper portion of the distribution function, but the ratio estimator is superior toward the lower end of the distribution function. The point along the distribution function at which the difference estimator becomes more precise than the ratio estimator depends on the sampling design, as well as the coefficient of variation of X and . A simple confidence interval procedure provides close to nominal coverage for intervals constructed from both the difference and ratio estimators, with the exception that coverage may be poor for the lower tail of the distribution function when using the ratio estimator.     

Design based estimation for ranked set sampling in finite populations

In this paper, we consider design-based estimation using ranked set sampling (RSS) in finite populations. We first derive the first and second-order inclusion probabilities for an RSS design and present two Horvitz–Thompson type estimators using these inclusion probabilities. We also develop an alternate Hansen–Hurwitz type estimator and investigate its properties. In particular, we show that this alternate estimator always outperforms the usual Hansen–Hurwitz type estimator in the simple random sampling with replacement design with comparable sample size. We also develop formulae for ratio estimator for all three developed estimators. The theoretical results are augmented by numerical and simulation studies as well as a case study using a well known data set. These show that RSS design can yield a substantial improvement in efficiency over the usual simple random sampling design in finite populations.     

Comparison between Hansen-Hurwitz and Horvitz-Thompson estimators for adaptive cluster sampling

Thompson (1990) introduced the adaptive cluster sampling design. This sampling design has been shown to be a useful sampling method for parameter estimation of a clustered and scattered population (Roesch, 1993; Smith et al., 1995; Thompson and Seber, 1996). Two estimators, the modified Hansen-Hurwitz (HH) and Horvitz-Thompson (HT) estimators, are available to estimate the mean or total of a population. Empirical results from previous researches indicate that the modified HT estimator has smaller variance than the modified HH estimator. We analytically compare the properties of these two estimators. Some results are obtained in favor of the modified HT estimator so that practitioners are strongly recommended to use the HT estimator despite easiness of computations for the HH estimator.     

A combination line transect and capture-recapture sampling model for multiple observers in aerial surveys

We present a robust sampling methodology to estimate population size using line transect and capture-recapture procedures for aerial surveys. Aerial surveys usually underestimate population density due to animals being missed. A combination of capture-recapture and line transect sampling methods with multiple observers allows violation of the assumption that all animals on the centreline are sighted from the air. We illustrate our method with an example of inanimate objects which shows evidence of failure of the assumption that all objects on the centreline have probability 1 of being detected. A simulation study is implemented to evaluate the performance of three variations of the Lincoln-Petersen estimator: the overall estimator, the stratified estimator, and the general stratified estimator based on the combined likelihood proposed in this paper. The stratified Lincoln-Petersen estimator based on the combined likelihood is found to be generally superior to the other estimators.     

Rao-Blackwell versions of the Horvitz-Thompson and Hansen-Hurwitz in adaptive cluster sampling

Thompson (1990) introduced the adaptive cluster sampling design and developed two unbiased estimators, the modified Horvitz-Thompson (HT) and Hansen-Hurwitz (HH) estimators, for this sampling design and noticed that these estimators are not a function of the minimal sufficient statistics. He applied the Rao-Blackwell theorem to improve them. Despite having smaller variances, these latter estimators have not received attention because a suitable method or algorithm for computing them was not available. In this paper we obtain closed forms of the Rao-Blackwell versions which can easily be computed. We also show that the variance reduction for the HH estimator is greater than that for the HT estimator using Rao-Blackwell versions. When the condition for extra samples is 0$$ " align="middle" border="0"> , one can expect some Rao-Blackwell improvement in the HH estimator but not in the HT estimator. Two examples are given.     

Randomized graph sampling

Randomized graph sampling (RGS) is an approach for sampling populations associated with or describable as graphs, when the structure of the graph is known and the parameter of interest is the total weight of the graph. RGS is related to, but distinct from, other graph-based approaches such as snowball and network sampling. Graph elements are clustered into walks that reflect the structure of the graph, as well as operational constraints on sampling. The basic estimator in RGS can be constructed as a Horvitz-Thompson estimator. I prove it to be design-unbiased, and also show design-unbiasedness of an estimator of the sample variance when walks are sampled with replacement. Covariates can be employed for variance reduction either through improved assignment of selection probabilities to walks in the design step, or through the use of alternative estimators during analysis. The approach is illustrated with a trail maintenance example, which demonstrates that complicated approaches to assignment of selection probabilities can be counterproductive. I describe conditions under which RGS may be efficient in practice, and suggest possible applications.     

Estimation of the CDF of a finite population in the presence of acalibration sample

We compare the performance of a number of estimators of the cumulative distribution function (CDF) for the following scenario: imperfect measurements are taken on an initial sample from afinite population and perfect measurements are obtained on a small calibration subset of the initial sample. The estimators we considered include two naive estimators using perfect and imperfect measurements; the ratio, difference and regression estimators for a two-phasesample; a minimum MSE estimator; Stefanski and Bay's SIMEX estimator (1996); and two proposed estimators. The proposed estimators take the form of a weighted average of perfect and imperfect measurements. They are constructed by minimizing variance among the class of weighted averages subject to an unbiasedness constraint. They differ in the manner of estimating the weight parameters. The first one uses direct sample estimates. The second one tunes the unknown parameters to an underlying normal distribution. We compare the root mean square error (RMSE) of the proposed estimator against other potential competitors through computer simulations. Our simulations show that our second estimator has the smallest RMSE among thenine compared and that the reduction in RMSE is substantial when the calibration sample is small and the error is medium or large.     

An EM algorithm for capture-recapture estimation

Analysis of capture—recapture data often involves maximizing a complex likelihood function with many unknown parameters. Statistical inference based on selection of a proper model depends on successful attainment of this maximum. An EM algorithm is developed for obtaining maximum likelihood estimates of capture and survival probabilities conditional on first capture from standard capture—recapture data. The algorithm does not require the use of numerical derivatives which may improve precision and stability relative to other estimation schemes. The asymptotic covariance matrix of the estimated parameters can be obtained using the supplemented EM algorithm. The EM algorithm is compared to a more traditional Newton-Raphson algorithm with both a simulated and a real dataset. The two algorithms result in the same parameter estimates, but Newton-Raphson variance estimates depend on a numerically estimated Hessian matrix that is sensitive to step size choice.     

Estimating Sonoran pronghorn abundance and survival with fecal DNA and capture–recapture methods

Population abundance estimates are important for management but can be challenging to determine in low‐density, wide‐ranging, and endangered species, such as Sonoran pronghorn (Antilocapra americana sonoriensis). The Sonoran pronghorn population has been increasing; however, population estimates are currently derived from a biennial aerial count that does not provide survival or recruitment estimates. We identified individuals through noninvasively collected fecal DNA and used robust‐design capture–recapture to estimate abundance and survival for Sonoran pronghorn in the United States from 2013 to 2014. In 2014 we generated separate population estimates for pronghorn gathered near 13 different artificial water holes and for pronghorn not near water holes. The population using artificial water holes had 116 (95% CI 102–131) and 121 individuals (95% CI 112–132) in 2013 and 2014, respectively. For all locations, we estimated there were 144 individuals (95% CI 132–157). Adults had higher annual survival probabilities (0.83, 95% CI 0.69–0.92) than fawns (0.41, 95% CI 0.21–0.65). Our use of targeted noninvasive genetic sampling and capture–recapture with Sonoran pronghorn fecal DNA was an effective method for monitoring a large proportion of the population. Our results provided the first survival estimates for this population in over 2 decades and precise estimates of the population using artificial water holes. Our method could be used for targeted sampling of broadly distributed species in other systems, such as in African savanna ecosystems, where many species congregate at watering sites.     

Estimator bias and efficiency for adaptive cluster sampling with order statistics and a stopping rule

Practical problems facing adaptive cluster sampling with order statistics (acsord) are explored using Monte Carlo simulation for three simulated fish populations and two known waterfowl populations. First, properties of an unbiased Hansen-Hurwitz (HH) estimator and a biased alternative Horvitz-Thompson (HT) estimator are evaluated. An increase in the level of population aggregation or the initial sample size increases the efficiencies of the two acsord estimators. For less aggregated fish populations, the efficiencies decrease as the order statistic parameter r (the number of units about which adaptive sampling is carried out) increases; for the highly aggregated fish and waterfowl populations, they increase with r. Acsord is almost always more efficient than simple random sampling for the highly aggregated populations. Positive bias is observed for the HT estimator, with the maximum bias usually occurring at small values of r. Secondly, a stopping rule at the Sth iteration of adaptive sampling beyond the initial sampling unit was applied to the acsord design to limit the otherwise open-ended sampling effort. The stopping rule induces relatively high positive bias to the HH estimator if the level of the population aggregation is high, the stopping level S is small, and r is large. The bias of HT is not very sensitive to the stopping rule and its bias is often reduced by the stopping rule at smaller values of r. For more aggregated populations, the stopping rule often reduces the efficiencies of the estimators compared to the non-stopping-rule scheme, but acsord still remains more efficient than simple random sampling. Despite its bias and lack of theoretical grounding, the HT estimator is usually more efficient than the HH estimator. In the stopping rule case, the HT estimator is preferable, because its bias is less sensitive to the stopping level.     

Variances and variance estimators of the improved ratio estimators under adaptive cluster sampling

Although not design-unbiased, the ratio estimator is recognized as more efficient when a certain degree of correlation exists between the variable of primary interest and the auxiliary variable. Meanwhile, the Rao–Blackwell method is another commonly used procedure to improve estimation efficiency. Various improved ratio estimators under adaptive cluster sampling (ACS) that make use of the auxiliary information together with the Rao–Blackwellized univariate estimators have been proposed in past research studies. In this article, the variances and the associated variance estimators of these improved ratio estimators are proposed for a thorough framework of statistical inference under ACS. Performance of the proposed variance estimators is evaluated in terms of the absolute relative percentage bias and the empirical mean-squared error. As expected, results show that both the absolute relative percentage bias and the empirical mean-squared error decrease as the initial sample size increases for all the variance estimators. To evaluate the confidence intervals based on these variance estimators and the finite-population Central Limit Theorem, the coverage rate and the interval width are used. These confidence intervals suffer a disadvantage similar to that of the conventional ratio estimator. Hence, alternative confidence intervals based on a certain type of adjusted variance estimators are constructed and assessed in this article.     

A statistical evaluation of non-ergodic variogram estimators

Geostatistics is a set of statistical techniques that is increasingly used to characterize spatial dependence in spatially referenced ecological data. A common feature of geostatistics is predicting values at unsampled locations from nearby samples using the kriging algorithm. Modeling spatial dependence in sampled data is necessary before kriging and is usually accomplished with the variogram and its traditional estimator. Other types of estimators, known as non-ergodic estimators, have been used in ecological applications. Non-ergodic estimators were originally suggested as a method of choice when sampled data are preferentially located and exhibit a skewed frequency distribution. Preferentially located samples can occur, for example, when areas with high values are sampled more intensely than other areas. In earlier studies the visual appearance of variograms from traditional and non-ergodic estimators were compared. Here we evaluate the estimators' relative performance in prediction. We also show algebraically that a non-ergodic version of the variogram is equivalent to the traditional variogram estimator. Simulations, designed to investigate the effects of data skewness and preferential sampling on variogram estimation and kriging, showed the traditional variogram estimator outperforms the non-ergodic estimators under these conditions. We also analyzed data on carabid beetle abundance, which exhibited large-scale spatial variability (trend) and a skewed frequency distribution. Detrending data followed by robust estimation of the residual variogram is demonstrated to be a successful alternative to the non-ergodic approach.     

Robust optimal observation of a metapopulation

We consider a two-parameter continuous-time Markovian model for patch occupancy in metapopulation, and address the question of when to observe the population in order to obtain the most accurate and precise estimator of the parameters. Using the likelihood obtained from a diffusion approximation we derive a robust procedure for determining the optimal observation schedule, which is considerably simpler than would otherwise be possible. We investigate the performance of two optimality criteria, ED-optimality and a particular form of maximin-optimality. Both allow one to incorporate prior belief about the parameter values before any data is collected. Kernel density estimates of the maximum likelihood estimator are compared under the various combinations of optimality criteria and prior distributions. Our methods are illustrated with reference to a model for the spread of crown of thorns starfish (Acanthaster planci) among the 55 islands comprising the Ryukyu group in Japan.     

A robust design capture-recapture model with multiple age classes augmented with population assignment data

The relative contribution of in situ reproduction versus immigration to the recruitment process is important to ecologists. Here we consider a robust design superpopulation capture-recapture model for a population with two age classes augmented with population assignment data. We first use age information to estimate the entry probabilities of new animals originating via in situ reproduction and immigration separately for all except the first period. Then we combine age and population assignment information with the capture-recapture model, which enables us to estimate the entry probability of in situ births and the entry probability of immigrants separately for all sampling periods. Further, this augmentation of age specific capture-recapture data with population assignment data greatly improves the estimators’ precision. We apply our new model to a capture-recapture data set with genetic information for banner-tailed kangaroo rats in Southern Arizona. We find that many more individuals are born in situ than are immigrants for all time periods. Young animals have lower survival probabilities than adults born in situ. Adult animals born in situ have higher survival probabilities than adults that were immigrants.     

An estimating function approach to the inference of catch-effort models

A class of catch-effort models, which allows for heterogeneous removal probabilities, is proposed for closed populations. The model includes three types of removal probabilities: multiplicative, Poisson and logistic. The usual removal and generalized removal models then become special cases. The equivalence of the proposed model and a special type of capture-recapture model is discussed. A unified estimating function approach is used to estimate the initial population size. For the homogeneous model, the resulting population size estimator based on optimal estimating functions is asymptotically equivalent to the maximum likelihood estimator. One advantage for our approach is that it can be extended to handle the heterogeneous populations in which the maximum likelihood estimators do not exist. The bootstrap method is applied to construct variance estimators and confidence intervals. We illustrate the method by two real data examples. Results of a simulation study investigating the performance of the proposed estimation procedure are presented.     

Regression-type estimators for adaptive two-stage sequential sampling

Adaptive two-stage sequential sampling (ATSSS) design was developed to observe more rare units and gain higher efficiency, in the sense of having a smaller variance estimator, than conventional sampling designs with equal effort for rare and spatially cluster populations. For certain rare populations, incorporating auxiliary variables into a sampling design can further improve the observation of rare units and increase efficiency. In this article, we develop regression-type estimators for ATSSS so that auxiliary variables can be incorporated into the ATSSS design when warranted. Simulation studies on two populations show that the regression-type estimators can significantly increase the efficiency of ATSSS and the detection of more rare units as compared to conventional sampling counterparts. Simulation of sampling of desert shrubs in Inner Mongolia (one of the two populations studied) showed that by incorporating a GIS auxiliary variable into ATSSS with the regression estimators resulted in a gain in efficiency over ATSSS without the auxiliary variable. Further, we found that the use of the GIS auxiliary variable in a conventional two-stage design with a regression estimator did not show a gain in efficiency.