A Bayesian approach to line-transect analysis for estimating abundance

Line-transect analysis is a widely used method of estimating plant and animal density and abundance. A Bayesian approach to a basic line-transect analysis is developed for a half-normal detection function. We extend the model of Karunamuni and Quinn [Karunamuni, R.J., Quinn II, T.J., 1995. Bayesian estimation of animal abundance for line-transect sampling. Biometrics 51, 1325–1337] by including a binomial likelihood function for the number of objects detected. The method computes a joint posterior distribution on the effective strip width and the density of objects in the sampled area. Analytical and computational methods for binned and unbinned perpendicular distance data are provided. Existing information about effective strip width and density can be brought into the analysis via prior distributions. The Bayesian approach is compared to a standard line-transect analysis using both real and simulated data. Results of the Bayesian and non-Bayesian analyses are similar when there are no prior data on effective strip width or density, but the Bayesian approach performs better when such data are available from previous or related studies. Practical methods for including prior data on effective strip width and density are suggested. A numerical example shows how the Bayesian approach can provide valid estimates when the sample size is too small for the standard approach to work reliably. The proposed Bayesian approach can form the basis for developing more advanced analyses.     

Bayesian areal wombling via adjacency modeling

Recently, public health professionals and other geostatistical researchers have shown increasing interest in boundary analysis, the detection or testing of zones or boundaries that reveal sharp changes in the values of spatially oriented variables. For areal data (i.e., data which consist only of sums or averages over geopolitical regions), Lu and Carlin (Geogr Anal 37: 265–285, 2005) suggested a fully model-based framework for areal wombling using Bayesian hierarchical models with posterior summaries computed using Markov chain Monte Carlo (MCMC) methods, and showed the approach to have advantages over existing non-stochastic alternatives. In this paper, we develop Bayesian areal boundary analysis methods that estimate the spatial neighborhood structure using the value of the process in each region and other variables that indicate how similar two regions are. Boundaries may then be determined by the posterior distribution of either this estimated neighborhood structure or the regional mean response differences themselves. Our methods do require several assumptions (including an appropriate prior distribution, a normal spatial random effect distribution, and a Bernoulli distribution for a set of spatial weights), but also deliver more in terms of full posterior inference for the boundary segments (e.g., direct probability statements regarding the probability that a particular border segment is part of the boundary). We illustrate three different remedies for the computing difficulties encountered in implementing our method. We use simulation to compare among existing purely algorithmic approaches, the Lu and Carlin (2005) method, and our new adjacency modeling methods. We also illustrate more practical modeling issues (e.g., covariate selection) in the context of a breast cancer late detection data set collected at the county level in the state of Minnesota.     

Inferential permutation tests for maximum entropy models in ecology

Maximum entropy (maxent) models assign probabilities to states that (1) agree with measured macroscopic constraints on attributes of the states and (2) are otherwise maximally uninformative and are thus as close as possible to a specified prior distribution. Such models have recently become popular in ecology, but classical inferential statistical tests require assumptions of independence during the allocation of entities to states that are rarely fulfilled in ecology. This paper describes a new permutation test for such maxent models that is appropriate for very general prior distributions and for cases in which many states have zero abundance and that can be used to test for conditional relevance of subsets of constraints. Simulations show that the test gives correct probability estimates under the null hypothesis. Power under the alternative hypothesis depends primarily on the number and strength of the constraints and on the number of states in the model; the number of empty states has only a small effect on power. The test is illustrated using two empirical data sets to test the community assembly model of B. Shipley, D. Vile, and E. Garnier and the species abundance distribution models of S. Pueyo, F. He, and T. Zillio.     

Poor-data and data-poor species stock assessment using a Bayesian hierarchical approach

Appropriate inference for stocks or species with low-quality data (poor data) or limited data (data poor) is extremely important. Hierarchical Bayesian methods are especially applicable to small-area, small-sample-size estimation problems because they allow poor-data species to borrow strength from species with good-quality data. We used a hammerhead shark complex as an example to investigate the advantages of using hierarchical Bayesian models in assessing the status of poor-data and data-poor exploited species. The hammerhead shark complex (Sphyrna spp.) along the Atlantic and Gulf of Mexico coasts of the United States is composed of three species: the scalloped hammerhead (S. lewini), the great hammerhead (S. mokarran), and the smooth hammerhead (S. zygaena) sharks. The scalloped hammerhead comprises 70-80% of the catch and has catch and relative abundance data of good quality, whereas great and smooth hammerheads have relative abundance indices that are both limited and of low quality presumably because of low stock density and limited sampling. Four hierarchical Bayesian state-space surplus production models were developed to simulate variability in population growth rates, carrying capacity, and catchability of the species. The results from the hierarchical Bayesian models were considerably more robust than those of the nonhierarchical models. The hierarchical Bayesian approach represents an intermediate strategy between traditional models that assume different population parameters for each species and those that assume all species share identical parameters. Use of the hierarchical Bayesian approach is suggested for future hammerhead shark stock assessments and for modeling fish complexes with species-specific data, because the poor-data species can borrow strength from the species with good data, making the estimation more stable and robust.     

The Importance of Being Spatial (and Reserved): Assessing Northern Spotted Owl Habitat Relationships with Hierarchical Bayesian Models

Abstract:  Regional conservation planning increasingly draws on habitat suitability models to support decisions regarding land allocation and management. Nevertheless, statistical techniques commonly used for developing such models may give misleading results because they fail to account for 3 factors common in data sets of species distribution: spatial autocorrelation, the large number of sites where the species is absent (zero inflation), and uneven survey effort. We used spatial autoregressive models fit with Bayesian Markov Chain Monte Carlo techniques to assess the relationship between older coniferous forest and the abundance of Northern Spotted Owl nest and activity sites throughout the species' range. The spatial random-effect term incorporated in the autoregressive models successfully accounted for zero inflation and reduced the effect of survey bias on estimates of species–habitat associations. Our results support the hypothesis that the relationship between owl distribution and older forest varies with latitude. A quadratic relationship between owl abundance and older forest was evident in the southern portion of the range, and a pseudothreshold relationship was evident in the northern portion of the range. Our results suggest that proposed changes to the network of owl habitat reserves would reduce the proportion of the population protected by up to one-third, and that proposed guidelines for forest management within reserves underestimate the proportion of older forest associated with maximum owl abundance and inappropriately generalize threshold relationships among subregions. Bayesian spatial models can greatly enhance the utility of habitat analysis for conservation planning because they add the statistical flexibility necessary for analyzing regional survey data while retaining the interpretability of simpler models.     

Efficacy of extracting indices from large‐scale acoustic recordings to monitor biodiversity

Passive acoustic monitoring could be a powerful way to assess biodiversity across large spatial and temporal scales. However, extracting meaningful information from recordings can be prohibitively time consuming. Acoustic indices (i.e., a mathematical summary of acoustic energy) offer a relatively rapid method for processing acoustic data and are increasingly used to characterize biological communities. We examined the relationship between acoustic indices and the diversity and abundance of biological sounds in recordings. We reviewed the acoustic‐index literature and found that over 60 indices have been applied to a range of objectives with varying success. We used 36 of the most indicative indices to develop a predictive model of the diversity of animal sounds in recordings. Acoustic data were collected at 43 sites in temperate terrestrial and tropical marine habitats across the continental United States. For terrestrial recordings, random‐forest models with a suite of acoustic indices as covariates predicted Shannon diversity, richness, and total number of biological sounds with high accuracy (R² ≥ 0.94, mean squared error [MSE] ≤170.2). Among the indices assessed, roughness, acoustic activity, and acoustic richness contributed most to the predictive ability of models. Performance of index models was negatively affected by insect, weather, and anthropogenic sounds. For marine recordings, random‐forest models poorly predicted Shannon diversity, richness, and total number of biological sounds (R² ≤ 0.40, MSE ≥ 195). Our results suggest that using a combination of relevant acoustic indices in a flexible model can accurately predict the diversity of biological sounds in temperate terrestrial acoustic recordings. Thus, acoustic approaches could be an important contribution to biodiversity monitoring in some habitats.     

New multidimensional functional diversity indices for a multifaceted framework in functional ecology

Functional diversity is increasingly identified as an important driver of ecosystem functioning. Various indices have been proposed to measure the functional diversity of a community, but there is still no consensus on which are most suitable. Indeed, none of the existing indices meets all the criteria required for general use. The main criteria are that they must be designed to deal with several traits, take into account abundances, and measure all the facets of functional diversity. Here we propose three indices to quantify each facet of functional diversity for a community with species distributed in a multidimensional functional space: functional richness (volume of the functional space occupied by the community), functional evenness (regularity of the distribution of abundance in this volume), and functional divergence (divergence in the distribution of abundance in this volume). Functional richness is estimated using the existing convex hull volume index. The new functional evenness index is based on the minimum spanning tree which links all the species in the multidimensional functional space. Then this new index quantifies the regularity with which species abundances are distributed along the spanning tree. Functional divergence is measured using a novel index which quantifies how species diverge in their distances (weighted by their abundance) from the center of gravity in the functional space. We show that none of the indices meets all the criteria required for a functional diversity index, but instead we show that the set of three complementary indices meets these criteria. Through simulations of artificial data sets, we demonstrate that functional divergence and functional evenness are independent of species richness and that the three functional diversity indices are independent of each other. Overall, our study suggests that decomposition of functional diversity into its three primary components provides a meaningful framework for its quantification and for the classification of existing functional diversity indices. This decomposition has the potential to shed light on the role of biodiversity on ecosystem functioning and on the influence of biotic and abiotic filters on the structure of species communities. Finally, we propose a general framework for applying these three functional diversity indices.     

Species abundance distributions result from body size-energetics relationships

Abundance distributions are a central characteristic of ecosystems. Certain distributions have been derived from theoretical models of community organization, and therefore the fit of data to these distributions has been proposed as a test of these theories. However, it is shown here that the geometric sequence distribution can be derived directly from the empirical relationship between population density and body size, with the assumption of random or uniform body size distributions on a log scale (as holds at local scales). The geometric sequence model provides a good to excellent fit to empirical data. The presence of noise in the relationship between population density and body size creates a curve that begins to approximate a lognormal species abundance distribution as the noise term increases. For continental-scale data in which the body size distribution is not flat, the result of sampling tends again toward the lognormal. Repeat sampling over time smooths out species population fluctuations and damps out the noise, giving a more precise geometric sequence abundance distribution. It is argued that the direct derivation of this distribution from empirical relationships gives it priority over distributions derived from complex theoretical community models.     

Spatial statistics for modeling of abundance and distribution of wildlife species in the Masai Mara ecosystem, Kenya

This study illustrates the use of modern statistical procedures for better wildlife management by addressing three key issues: determination of abundance, modeling of animal distributions and variability of diversity in space and time. Prior information in Markov Chain Monte Carlo (MCMC) methods is used to improve estimates of abundance. Measures of autocorrelation are included when modeling distributions of animal counts, and a diversity index to indicate species abundance and richness for large herbivores is developed. Data from the Masai Mara ecosystem in Kenya are used to develop and demonstrate these procedures. The new abundance estimates are up to 35% more accurate than those obtained by existing methods. Significant temporal changes in spatial patterns are found from a space-time analysis of elephant counts over a 20-year period, with strong interactions over 5 km and 6 months space and time separations, respectively. The new diversity index is sensitive to both high abundance and species richness and is also able to capture year to year variation. It indicates an overall marginal decrease in diversity for large herbivores in the Mara ecosystem. The space-time analyses and diversity index can easily be computed thereby providing tools for rapid decision making.     

An integrated approach for predicting fates of reintroductions with demographic data from multiple populations

We devised a novel approach to model reintroduced populations whereby demographic data collected from multiple sites are integrated into a Bayesian hierarchical model. Integrating data from multiple reintroductions allows more precise population-growth projections to be made, especially for populations for which data are sparse, and allows projections that account for random site-to-site variation to be made before new reintroductions are attempted. We used data from reintroductions of the North Island Robin (Petroica longipes), an endemic New Zealand passerine, to 10 sites where non-native mammalian predators are controlled. A comparison of candidate models that we based on deviance information criterion showed that rat-tracking rate (an index of rat density) was a useful predictor of robin fecundity and adult female survival, that landscape connectivity and a binary measure of whether sites were on a peninsula were useful predictors of apparent juvenile survival (probably due to differential dispersal away from reintroduction sites), and that there was unexplained random variation among sites in all demographic rates. We used the two best supported models to estimate the finite rate of increase (λ) for populations at each of the 10 sites, and for a proposed reintroduction site, under different levels of rat control. Only three of the reintroduction sites had λ distributions completely >1 for either model. At two sites, λ was expected to be >1 if rat-tracking rates were <5%. At the other five reintroduction sites, λ was predicted to be close to 1, and it was unclear whether growth was expected. Predictions of λ for the proposed reintroduction site were less precise than for other sites because distributions incorporated the full range of site-to-site random variation in vital rates. Our methods can be applied to any species for which postrelease data on demographic rates are available and potentially can be extended to model multiple species simultaneously.     

Line transect sampling of Karner blue butterflies ( Lycaeides melissa samuelis)

Line transect sampling is an effective survey method for estimating butterfly densities because it provides unbiased estimates of site-density (provided key assumptions are met), and estimates are comparable among sites. For monitoring Karner blue butterflies in Wisconsin, USA, comparable estimates are required because each year a different selection of sites will be monitored. Annual state-wide indices of species abundance can be derived from the site-surveys and compared to previous year's indices to monitor trends. We advocate that line transect sampling is preferable to Pollard-Yates transects as a survey technique for monitoring Karner blue butter- flies. The Pollard-Yates surveys do not adjust for diferences in site detectability. As a consequence, estimates of among-site from Pollard-Yates surveys can be biased. © Rapid Science 1998     

Range-wide effects of breeding- and nonbreeding-season climate on the abundance of a Neotropical migrant songbird

Geographic variation in the population dynamics of a species can result from regional variability in climate and how it affects reproduction and survival. Identifying such effects for migratory birds requires the integration of population models with knowledge of migratory connectivity between breeding and nonbreeding areas. We used Bayesian hierarchical models with 26 years of Breeding Bird Survey data (1982-2007) to investigate the impacts of breeding- and nonbreeding-season climate on abundance of American Redstarts (Setophaga ruticilla) across the species range. We focused on 15 populations defined by Bird Conservation Regions, and we included variation across routes and observers as well as temporal trends and climate effects. American Redstart populations that breed in eastern North America showed increased abundance following winters with higher plant productivity in the Caribbean where they are expected to overwinter. In contrast, western breeding populations showed little response to conditions in their expected wintering areas in west Mexico, perhaps reflecting lower migratory connectivity or differential effects of winter rainfall on individuals across the species range. Unlike the case with winter climate, we found few effects of temperature prior to arrival in spring (March-April) or during the nesting period (May-June) on abundance the following year. Eight populations showed significant changes in abundance, with the steepest declines in the Atlantic Northern Forest (-3.4%/yr) and the greatest increases in the Prairie Hardwood Transition (4%/yr). This study emphasizes how the effects of climate on populations of migratory birds are context dependent and can vary depending on geographic location and the period of the annual cycle. Such knowledge is essential for predicting regional variation in how populations of a species might vary in their response to climate change.     

A Bayesian approach to evaluating site impairment

In the United States, each state is required to list water resources that are declared to be impaired under guidelines set by the Clean Water Act. Measurements are typically collected on a number of chemical constituents and compared with a standard. If there are too many measurements exceeding the standard, then the site is declared impaired. The approach is non-statistical but similar to a Binomial test. The Binomial approach would convert the measurements to binary data then test if the proportion exceeding the standard is excessive. Both methods convert measurements to binary values hence exclude potentially important information in the data. We present a statistical approach using a Bayesian model that uses the raw data instead of the binary transformed data. The population distribution of a family of location-scale parameter models is studied under the model. Posterior distributions from the Bayesian analysis are used in the decision-making process and error probabilities for the Bayesian and the Binomial approaches are compared for a normal population.     

A Bayesian hierarchical framework for calibrating aquatic biogeochemical models

Model practitioners increasingly place emphasis on rigorous quantitative error analysis in aquatic biogeochemical models and the existing initiatives range from the development of alternative metrics for goodness of fit, to data assimilation into operational models, to parameter estimation techniques. However, the treatment of error in many of these efforts is arguably selective and/or ad hoc. A Bayesian hierarchical framework enables the development of robust probabilistic analysis of error and uncertainty in model predictions by explicitly accommodating measurement error, parameter uncertainty, and model structure imperfection. This paper presents a Bayesian hierarchical formulation for simultaneously calibrating aquatic biogeochemical models at multiple systems (or sites of the same system) with differences in their trophic conditions, prior precisions of model parameters, available information, measurement error or inter-annual variability. Our statistical formulation also explicitly considers the uncertainty in model inputs (model parameters, initial conditions), the analytical/sampling error associated with the field data, and the discrepancy between model structure and the natural system dynamics (e.g., missing key ecological processes, erroneous formulations, misspecified forcing functions). The comparison between observations and posterior predictive monthly distributions indicates that the plankton models calibrated under the Bayesian hierarchical scheme provided accurate system representations for all the scenarios examined. Our results also suggest that the Bayesian hierarchical approach allows overcoming problems of insufficient local data by “borrowing strength” from well-studied sites and this feature will be highly relevant to conservation practices of regions with a high number of freshwater resources for which complete data could never be practically collected. Finally, we discuss the prospect of extending this framework to spatially explicit biogeochemical models (e.g., more effectively connect inshore with offshore areas) along with the benefits for environmental management, such as the optimization of the sampling design of monitoring programs and the alignment with the policy practice of adaptive management.     

Studying ecological communities from a neutral standpoint: A review of models’ structure and parameter estimation

Neutral models provide an alternative to niche-based assembly rules of ecological communities by assuming that communities’ properties are shaped by the stochastic interplay between ecological drift, migration and speciation. The recent and ongoing interest about neutral assumptions has produced many developments on the theoretical side, with nevertheless limited echoes in terms of analyses of real-world data. The present review paper aims to help bridge the widening gap between modellers and field ecologists through two objectives. First, to provide a multi-criteria typology of the main neutral models, including those from population genetics that have not yet been transposed to ecology, by considering how the fundamental processes of ecological drift, speciation and migration are modelled and, specifically, how space is taken into account. Second, to review methods recently proposed to estimate models parameters from field data, a point that should be mastered to allow for broader applications.     

Using the negative binomial distribution to model overdispersion in ecological count data

A Poisson process is a commonly used starting point for modeling stochastic variation of ecological count data around a theoretical expectation. However, data typically show more variation than implied by the Poisson distribution. Such overdispersion is often accounted for by using models with different assumptions about how the variance changes with the expectation. The choice of these assumptions can naturally have apparent consequences for statistical inference. We propose a parameterization of the negative binomial distribution, where two overdispersion parameters are introduced to allow for various quadratic mean-variance relationships, including the ones assumed in the most commonly used approaches. Using bird migration as an example, we present hypothetical scenarios on how overdispersion can arise due to sampling, flocking behavior or aggregation, environmental variability, or combinations of these factors. For all considered scenarios, mean-variance relationships can be appropriately described by the negative binomial distribution with two overdispersion parameters. To illustrate, we apply the model to empirical migration data with a high level of overdispersion, gaining clearly different model fits with different assumptions about mean-variance relationships. The proposed framework can be a useful approximation for modeling marginal distributions of independent count data in likelihood-based analyses.     

A Bayesian approach to retransformation bias in transformed regression

Ecological data analysis often involves fitting linear or nonlinear equations to data after transforming either the response variable, the right side of the equation, or both, so that the standard suite of regression assumptions are more closely met. However, inference is usually done in the natural metric and it is well known that retransforming back to the original metric provides a biased estimator for the mean of the response variable. For the normal linear model, fit under a log-transformation, correction factors are available to reduce this bias, but these factors may not be generally applicable to all model forms or other transformations. We demonstrate that this problem is handled in a straightforward manner using a Bayesian approach, which is general for linear and nonlinear models and other transformations and model error structures. The Bayesian framework provides a predictive distribution for the response variable so that inference can be made at the mean, or over the entire distribution to incorporate the predictive uncertainty.     

The distribution of a generalized diversity index due to Good

Good (1953, 1982) proposed a generalized diversity index which includes as special cases both Shannon's and Simpson's indices. This index can be further generalized as described in Baczkowski et al. (1997, 1998). In this paper the first four moments of this generalized index are derived for both a general species abundance distribution and the case with all species abundances equal, the equiprobable case. This allows the skewness and kurtosis of the index to be determined and thus gives information about the distribution of the index.     

The apportionment of quadratic entropy: a useful alternative for partitioning diversity in ecological data

Many methods that study the diversity within hierarchically structured populations have been developed in genetics. Among them, the analysis of molecular variance (AMOVA) (Excoffier et al., 1992) has the advantage of including evolutionary distances between individuals. AMOVA is a special case of a far more general statistical scheme produced by Rao (1982a; 1986) and called the apportionment of quadratic entropy (APQE). It links diversity and dissimilarity and allows the decomposition of diversity according to a given hierarchy. We apply this framework to ecological data showing that APQE may be very useful for studying diversity at various spatial scales. Moreover, the quadratic entropy has a critical advantage over usual diversity indices because it takes into account differences between species. Finally, the differences that can be incorporated in APQE may be either taxonomic or functional (biological traits), which may be of critical interest for ecologists.     

Modelling environmental stochasticity in adult survival for a long-lived species

Stochastic matrix population models are often used to help guide the management of animal populations. For a long-lived species, environmental stochasticity in adult survival will play an important role in determining outcomes from the model. One of the most common methods for modelling such stochasticity is to randomly select the value of adult survival for each year from a distribution with a specified mean and standard deviation. We consider four distributions that can provide realistic models for stochasticity in adult survival. For values of the mean and standard deviation that cover the range we would expect for long-lived species, all four distributions have similar shapes, with small differences in their skewness and kurtosis. This suggests that many of the outcomes from a population model will be insensitive to the choice of distribution, assuming that distribution provides a realistic model for environmental stochasticity in adult survival. For a generic age-structured model, the estimate of the long-run stochastic growth rate is almost identical for the four distributions, across this range of values for the mean and standard deviation. Model outcomes based on short-term projections, such as the probability of a decline over a 20-year period, are more sensitive to the choice of distribution.