Estimation of abundance from presence–absence maps using cluster models

A presence–absence map consists of indicators of the occurrence or nonoccurrence of a given species in each cell over a grid, without counting the number of individuals in a cell once it is known it is occupied. They are commonly used to estimate the distribution of a species, but our interest is in using these data to estimate the abundance of the species. In practice, certain types of species (in particular flora types) may be spatially clustered. For example, some plant communities will naturally group together according to similar environmental characteristics within a given area. To estimate abundance, we develop an approach based on clustered negative binomial models with unknown cluster sizes. Our approach uses working clusters of cells to construct an estimator which we show is consistent. We also introduce a new concept called super-clustering used to estimate components of the standard errors and interval estimators. A simulation study is conducted to examine the performance of the estimators and they are applied to real data.     

Improving occupancy estimation when two types of observational error occur: non-detection and species misidentification

Efforts to draw inferences about species occurrence frequently account for false negatives, the common situation when individuals of a species are not detected even when a site is occupied. However, recent studies suggest the need to also deal with false positives, which occur when species are misidentified so that a species is recorded as detected when a site is unoccupied. Bias in estimators of occupancy, colonization, and extinction can be severe when false positives occur. Accordingly, we propose models that simultaneously account for both types of error. Our approach can be used to improve estimates of occupancy for study designs where a subset of detections is of a type or method for which false positives can be assumed to not occur. We illustrate properties of the estimators with simulations and data for three species of frogs. We show that models that account for possible misidentification have greater support (lower AIC for two species) and can yield substantially different occupancy estimates than those that do not. When the potential for misidentification exists, researchers should consider analytical techniques that can account for this source of error, such as those presented here.     

Spatiotemporal hierarchical modelling of extreme precipitation in Western Australia using anisotropic Gaussian random fields

We discuss an approach for the statistical modelling of extreme precipitation events in South-West Australia over space and time, using a latent spatiotemporal process where precipitation maxima follow a generalised extreme value distribution. Temporal features are captured by modelling trends on the location and scale parameters. Spatial features are captured using anisotropic Gaussian random fields. Site specific explanatory variables are also incorporated. We fit several models using Bayesian inferential methods to precipitation extremes recorded at 36 weather stations around the Western Australian state capital city of Perth over the period 1907–2009. Model choice is performed using the DIC criterion. The best fitting model shows significant non-stationarity over time, with extreme precipitation events becoming less frequent. Extreme precipitation events are stronger at coastal locations, with the intensity decreasing as we head to the higher and drier areas to the North-East.     

Analyzing Variability and the Rate of Decline of Migratory Shorebirds in Moreton Bay,Australia

Abstract: Estimating the abundance of migratory species is difficult because sources of variability differ substantially among species and populations. Recently developed state‐space models address this variability issue by directly modeling both environmental and measurement error, although their efficacy in detecting declines is relatively untested for empirical data. We applied state‐space modeling, generalized least squares (with autoregression error structure), and standard linear regression to data on abundance of wetland birds (shorebirds and terns) at Moreton Bay in southeast Queensland, Australia. There are internationally significant numbers of 8 species of waterbirds in the bay, and it is a major terminus of the large East Asian‐Australasian Flyway. In our analyses, we considered 22 migrant and 8 resident species. State‐space models identified abundances of 7 species of migrants as significantly declining and abundance of one species as significantly increasing. Declines in migrant abundance over 15 years were 43–79%. Generalized least squares with an autoregressive error structure showed abundance changes in 11 species, and standard linear regression showed abundance changes in 15 species. The higher power of the regression models meant they detected more declines, but they also were associated with a higher rate of false detections. If the declines in Moreton Bay are consistent with trends from other sites across the flyway as a whole, then a large number of species are in significant decline.     

Detecting and Modeling Spatial and Temporal Dependence in Conservation Biology

Abstract: Due to the structuring forces and large-scale physical processes that shape our biosphere, we often find that environmental and ecological data are either spatially or temporally—or both spatially and temporally—dependent. When these data are analyzed, statistical techniques and models are frequently applied that were developed for independent data. We describe some of the detrimental consequences, such as inefficient parameter estimators, biased hypothesis test results, and inaccurate predictions, of ignoring spatial and temporal data dependencies, and we cite an example of adverse statistical results occurring when spatial dependencies were disregarded. We also discuss and recommend available techniques used to detect and model spatial and temporal dependence, including variograms, covariograms, autocorrelation and partial autocorrelation plots, geostatistical techniques, Gaussian autoregressive models, K functions, and ARIMA models, in environmental and ecological research to avoid the aforementioned difficulties.     

Combining multistate capture-recapture data with tag recoveries to estimate demographic parameters

Matrix population models that allow an animal to occupy more than one state over time are important tools for population and evolutionary ecologists. Definition of state can vary, including location for metapopulation models and breeding state for life history models. For populations whose members can be marked and subsequently reencountered, multistate mark-recapture models are available to estimate the survival and transition probabilities needed to construct population models. Multistate models have proved extremely useful in this context, but they often require a substantial amount of data and restrict estimation of transition probabilities to those areas or states subjected to formal sampling effort. At the same time, for many species, there are considerable tag recovery data provided by the public that could be modeled in order to increase precision and to extend inference to a greater number of areas or states. Here we present a statistical model for combining multistate capture-recapture data (e.g., from a breeding ground study) with multistate tag recovery data (e.g., from wintering grounds). We use this method to analyze data from a study of Canada Geese (Branta canadensis) in the Atlantic Flyway of North America. Our analysis produced marginal improvement in precision, due to relatively few recoveries, but we demonstrate how precision could be further improved with increases in the probability that a retrieved tag is reported.     

Estimation bias under model selection for distance sampling detection functions

Many simulation studies have examined the properties of distance sampling estimators of wildlife population size. When assumptions hold, if distances are generated from a detection model and fitted using the same model, they are known to perform well. However, in practice, the true model is unknown. Therefore, standard practice includes model selection, typically using model comparison tools like Akaike Information Criterion. Here we examine the performance of standard distance sampling estimators under model selection. We compare line and point transect estimators with distances simulated from two detection functions, hazard-rate and exponential power series (EPS), over a range of sample sizes. To mimic the real-world context where the true model may not be part of the candidate set, EPS models were not included as candidates, except for the half-normal parameterization. We found median bias depended on sample size (being asymptotically unbiased) and on the form of the true detection function: negative bias (up to 15% for line transects and 30% for point transects) when the shoulder of maximum detectability was narrow, and positive bias (up to 10% for line transects and 15% for point transects) when it was wide. Generating unbiased simulations requires careful choice of detection function or very large datasets. Practitioners should collect data that result in detection functions with a shoulder similar to a half-normal and use the monotonicity constraint. Narrow-shouldered detection functions can be avoided through good field procedures and those with wide shoulder are unlikely to occur, due to heterogeneity in detectability.     

Comparison of Two Types of Metapopulation Models in Real and Artificial Landscapes

Abstract: Application of metapopulation models is becoming increasingly widespread in the conservation of species in fragmented landscapes. We provide one of the first detailed comparisons of two of the most common modeling techniques, incidence function models and stage-based matrix models, and test their accuracy in predicting patch occupancy for a real metapopulation. We measured patch occupancies and demographic rates for regional populations of the Florida scrub lizard (   Sceloporus woodi ) and compared the observed occupancies with those predicted by each model. Both modeling strategies predicted patch occupancies with good accuracy ( 77–80%) and gave similar results when we compared hypothetical management scenarios involving removal of key habitat patches and degradation of habitat quality. To compare the two modeling approaches over a broader set of conditions, we simulated metapopulation dynamics for 150 artificial landscapes composed of equal-sized patches (2–1024 ha) spaced at equal distances (50–750 m). Differences in predicted patch occupancy were small to moderate (<20%) for about 74% of all simulations, but 22% of the landscapes had differences openface> 50%. Incidence function models and stage-based matrix models differ in their approaches, assumptions, and requirements for empirical data, and our findings provide evidence that the two models can produce different results. We encourage researchers to use both techniques and further examine potential differences in model output. The feasibility of obtaining data for population modeling varies widely among species and limits the modeling approaches appropriate for each species. Understanding different modeling approaches will become increasingly important as conservation programs undertake the challenge of managing for multiple species in a landscape context.     

Variation partitioning of species data matrices: estimation and comparison of fractions

Establishing relationships between species distributions and environmental characteristics is a major goal in the search for forces driving species distributions. Canonical ordinations such as redundancy analysis and canonical correspondence analysis are invaluable tools for modeling communities through environmental predictors. They provide the means for conducting direct explanatory analysis in which the association among species can be studied according to their common and unique relationships with the environmental variables and other sets of predictors of interest, such as spatial variables. Variation partitioning can then be used to test and determine the likelihood of these sets of predictors in explaining patterns in community structure. Although variation partitioning in canonical analysis is routinely used in ecological analysis, no effort has been reported in the literature to consider appropriate estimators so that comparisons between fractions or, eventually, between different canonical models are meaningful. In this paper, we show that variation partitioning as currently applied in canonical analysis is biased. We present appropriate unbiased estimators. In addition, we outline a statistical test to compare fractions in canonical analysis. The question addressed by the test is whether two fractions of variation are significantly different from each other. Such assessment provides an important step toward attaining an understanding of the factors patterning community structure. The test is shown to have correct Type I. error rates and good power for both redundancy analysis and canonical correspondence analysis.     

Life history and viability of a long-lived marine invertebrate: the octocoral Paramuricea clavata

The red gorgonian Paramuricea clavata is a long-lived, slow-growing sessile invertebrate of ecological and conservation importance in the northwestern Mediterranean Sea. We develop a series of size-based matrix models for two Paramuricea clavata populations. These models were used to estimate basic life history traits for this species and to evaluate the viability of the red gorgonian populations we studied. As for many other slow-growing species, sensitivity and elasticity analysis demonstrate that gorgonian population growth is far more sensitive to changes in survival rates than to growth, shrinkage, or reproductive rates. The slow growth and low mortality of red gorgonians results in low damping ratios, indicating slow convergence to stable size structures (at least 50 years). The stable distributions predicted by the model did not differ from the observed ones. However, our simulations point out the fragility of this species, showing both populations in decline and high risk of extinction over moderate time horizons. These declines appear to be related to a recent increase in anthropogenic disturbances. Relative to their life span, the values of recruitment elasticity for Paramuricea clavata are lower than those reported for other marine organisms but are similar to those reported for some long-lived plants. These values and the delayed age of sexual maturity, in combination with the longevity of the species, show a clear fecundity/mortality trade-off. Full demographic studies of sessile marine species are quite scarce but can provide insight into population dynamics and life history patterns for these difficult and under-studied species. While our work shows clear results for the red gorgonian, the variability in some of our estimates suggest that future work should include data collection over longer temporal and spatial scales to better understand the long-term effects of natural and anthropogenic disturbances on red gorgonian populations.     

Efficient Markov chain Monte Carlo sampling for hierarchical hidden Markov models

Traditional Markov chain Monte Carlo (MCMC) sampling of hidden Markov models (HMMs) involves latent states underlying an imperfect observation process, and generates posterior samples for top-level parameters concurrently with nuisance latent variables. When potentially many HMMs are embedded within a hierarchical model, this can result in prohibitively long MCMC runtimes. We study combinations of existing methods, which are shown to vastly improve computational efficiency for these hierarchical models while maintaining the modeling flexibility provided by embedded HMMs. The methods include discrete filtering of the HMM likelihood to remove latent states, reduced data representations, and a novel procedure for dynamic block sampling of posterior dimensions. The first two methods have been used in isolation in existing application-specific software, but are not generally available for incorporation in arbitrary model structures. Using the NIMBLE package for R, we develop and test combined computational approaches using three examples from ecological capture–recapture, although our methods are generally applicable to any embedded discrete HMMs. These combinations provide several orders of magnitude improvement in MCMC sampling efficiency, defined as the rate of generating effectively independent posterior samples. In addition to being computationally significant for this class of hierarchical models, this result underscores the potential for vast improvements to MCMC sampling efficiency which can result from combinations of known algorithms.     

Estimating parameters of hidden Markov models based on marked individuals: use of robust design data

Development and use of multistate mark-recapture models, which provide estimates of parameters of Markov processes in the face of imperfect detection, have become common over the last 20 years. Recently, estimating parameters of hidden Markov models, where the state of an individual can be uncertain even when it is detected, has received attention. Previous work has shown that ignoring state uncertainty biases estimates of survival and state transition probabilities, thereby reducing the power to detect effects. Efforts to adjust for state uncertainty have included special cases and a general framework for a single sample per period of interest. We provide a flexible framework for adjusting for state uncertainty in multistate models, while utilizing multiple sampling occasions per period of interest to increase precision and remove parameter redundancy. These models also produce direct estimates of state structure for each primary period, even for the case where there is just one sampling occasion. We apply our model to expected-value data, and to data from a study of Florida manatees, to provide examples of the improvement in precision due to secondary capture occasions. We have also implemented these models in program MARK. This general framework could also be used by practitioners to consider constrained models of particular interest, or to model the relationship between within-primary-period parameters (e.g., state structure) and between-primary-period parameters (e.g., state transition probabilities).     

Nonparametric MLE incorporation of heterogeneity and model testing into premarked cohort studies

We utilize mixture models and nonparametric maximum likelihood estimation to both develop a likelihood ratio test (lrt) for a common simplifying assumption and to allow heterogeneity within premarked cohort studies. Our methods allow estimation of the entire probability model and thus one can not only estimate many parameters of interest but one can also bootstrap from the estimated model to predict many things, including the standard deviations of estimators. Simulations suggest that our lrt has the appropriate protection for Type I error and often has good power. In practice, our lrt is important for determining the appropriateness of estimators and in examining if a simple design with only one capture period could be utilized for a future similar study.     

An incidence-based richness estimator for quadrats sampled without replacement

Most richness estimators currently in use are derived from models that consider sampling with replacement or from the assumption of infinite populations. Neither of the assumptions is suitable for sampling sessile organisms such as plants where quadrats are often sampled without replacement and the area of study is always limited. In this paper, we propose an incidence-based parametric richness estimator that considers quadrat sampling without replacement in a fixed area. The estimator is derived from a zero-truncated binomial distribution for the number of quadrats containing a given species (e.g., species i) and a modified beta distribution for the probability of presence-absence of a species in a quadrat. The maximum likelihood estimate of richness is explicitly given and can be easily solved. The variance of the estimate is also obtained. The performance of the estimator is tested against nine other existing incidence-based estimators using two tree data sets where the true numbers of species are known. Results show that the new estimator is insensitive to sample size and outperforms the other methods as judged by the root mean squared errors. The superiority of the new method is particularly noticeable when large quadrat size is used, suggesting that a few large quadrats are preferred over many small ones when sampling diversity.     

Flexible hierarchical mark-recapture modeling for open populations using WinBUGS

Hierarchical mark-recapture models offer three advantages over classical mark-recapture models: (i) they allow expression of complicated models in terms of simple components; (ii) they provide a convenient way of modeling missing data and latent variables in a way that allows expression of relationships involving latent variables in the model; (iii) they provide a convenient way of introducing parsimony into models involving many nuisance parameters. Expressing models using the complete data likelihood we show how many of the standard mark-recapture models for open populations can be readily fitted using the software WinBUGS. We include examples that illustrate fitting the Cormack–Jolly–Seber model, multi-state and multi-event models, models including auxiliary data, and models including density dependence.

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.     

Lotka-Volterra competition models for sessile organisms

Markov models are widely used to describe the dynamics of communities of sessile organisms, because they are easily fitted to field data and provide a rich set of analytical tools. In typical ecological applications, at any point in time, each point in space is in one of a finite set of states (e.g., species, empty space). The models aim to describe the probabilities of transitions between states. In most Markov models for communities, these transition probabilities are assumed to be independent of state abundances. This assumption is often suspected to be false and is rarely justified explicitly. Here, we start with simple assumptions about the interactions among sessile organisms and derive a model in which transition probabilities depend on the abundance of destination states. This model is formulated in continuous time and is equivalent to a Lotka-Volterra competition model. We fit this model and a variety of alternatives in which transition probabilities do not depend on state abundances to a long-term coral reef data set. The Lotka-Volterra model describes the data much better than all models we consider other than a saturated model (a model with a separate parameter for each transition at each time interval, which by definition fits the data perfectly). Our approach provides a basis for further development of stochastic models of sessile communities, and many of the methods we use are relevant to other types of community. We discuss possible extensions to spatially explicit models.     

Optimal model averaging by data perturbation for spatial noisy data

Geostatistical models play an important role in spatial data analysis, in which model selection is inevitable. Model selection methods, such as AIC and BIC, are popular for selecting appropriate models. In recent years, some model averaging methods, such as smoothed AIC and smoothed BIC, are also applied to spatial data models. However, the corresponding averaging estimators are outperformed by optimal model averaging estimators (Hansen in Econometrica 75:1175–1189, 2007) for the ordinary linear models. Therefore, this paper focuses on the optimal model averaging method for geostatistical models. We propose a weight choice criterion for the model averaging estimator on the basis of the generalized degrees of freedom and data perturbation technique. We further theoretically prove the resultant estimator is asymptotically optimal in terms of the mean squared error, and numerically demonstrate its satisfactory performance. Finally, the proposed method is applied to a mercury data set.     

Habitat Suitability Models and the Shortfall in Conservation Planning for African Vertebrates

Abstract:  Ongoing loss of biodiversity requires identifying large-scale conservation priorities, but the detailed information on the distribution of species required for this purpose is often missing. We present a systematic reserve selection for 1223 African mammals and amphibians in which habitat suitability models are used as estimates of the area occupied by species. In the framework of the World Conservation Union (IUCN) Global Amphibian Assessment and IUCN Global Mammal Assessment, we collected the geographic range (extent of occurrence) and habitat preferences for each species. We used the latter to build species-specific habitat suitability models inside geographic ranges, and for 181 species we verified the models by comparing suitability levels to presence-absence data collected in the field. We then used the suitable areas as estimators of the area of occupancy and compared the results of systematic reserve selection based on geographic ranges to those based on estimated areas of occupancy. Our results showed that the reserve system would need a 30-100% expansion to achieve minimal conservation targets, concentrated in the tropics, where species richness reaches a maximum. Comparative analyses revealed that using geographic ranges, which overestimate the area occupied by species, underestimates the total amount of area that needs to be conserved. The area selected for conservation doubled when we used the estimated area of occupancy in place of the geographic ranges. This happened because the suitable areas potentially occupied by each species overlapped less than their geographic ranges. As a result, any given protected area contained fewer species than predicted by the analysis of ranges. Because species are more specialized than our estimates of distribution based on extent of occurrence suggest, we propose that this is a general effect in systematic conservation planning.     

Nonparametric mean estimation using partially ordered sets

In ranked-set sampling (RSS), the ranker must give a complete ranking of the units in each set. In this paper, we consider a modification of RSS that allows the ranker to declare ties. Our sampling method is simply to break the ties at random so that we obtain a standard ranked-set sample, but also to record the tie structure for use in estimation. We propose several different nonparametric mean estimators that incorporate the tie information, and we show that the best of these estimators is substantially more efficient than estimators that ignore the ties. As part of our comparison of estimators, we develop new results about models for ties in rankings. We also show that there are settings where, to achieve more efficient estimation, ties should be declared not just when the ranker is actually unsure about how units rank, but also when the ranker is sure about the ranking, but believes that the units are close.