On the use of correlated beta random variables with animal population modelling

We give reasons why demographic parameters such as survival and reproduction rates are often modelled well in stochastic population simulation using beta distributions. In practice, it is frequently expected that these parameters will be correlated, for example with survival rates for all age classes tending to be high or low in the same year. We therefore discuss a method for producing correlated beta random variables by transforming correlated normal random variables, and show how it can be applied in practice by means of a simple example. We also note how the same approach can be used to produce correlated uniform, triangular, and exponential random variables.     

A general random walk model for the leptokurtic distribution of organism movement: Theory and application

The movement of organisms is usually leptokurtic in which some individuals move long distances while the majority remains at or near the area they are released. There has been extensive research into the origin of such leptokurtic movement, but one important aspect that has been overlooked is that the foraging behaviour of most organisms is not Brownian as assumed in most existing models. In this paper we show that such non-Brownian foraging indeed gives rise to leptokurtic distribution. We first present a general random walk model to describe the organism movement by breaking the foraging of each individual into events of active movement and inactive stationary period; its foraging behaviour is therefore fully characterized by a joint probability of how far the individual can move in each active movement and the duration it remains stationary between two consecutive movements. The spatio-temporal distribution of the organism can be described by a generalized partial differential equation, and the leptokurtic distribution is a special case when the stationary period is not exponentially distributed. Empirical observations of some organisms living in different habitats indicated that their rest time shows a power-law distribution, and we speculate that this is general for other organisms. This leads to a fractional diffusion equation with three parameters to characterize the distributions of stationary period and movement distance. A method to estimate the parameters from empirical data is given, and we apply the model to simulate the movement of two organisms living in different habitats: a stream fish (Cyprinidae: Nocomis leptocephalus) in water, and a root-feeding weevil, Sitona lepidus in the soil. Comparison of the simulations with the measured data shows close agreement. This has an important implication in ecology that the leptokurtic distribution observed at population level does not necessarily mean population heterogeneity as most existing models suggested, in which the population consists of different phenotypes; instead, a homogeneous population moving in homogeneous habitat can also lead to leptokurtic distribution.     

Estimating age-specific hazards from wildlife telemetry data

We illustrate 2 techniques for estimating age-specific hazards with wildlife telemetry data: Siler’s (Ecology 60:750–757, 1979) competing risk model fit using maximum likelihood and a penalized likelihood estimate that only assumes the hazard varies smoothly with age. In most telemetry studies, animals enter at different points in time (and at different ages), leading to data that are left-truncated. In addition, death times may only be known to occur within an interval of time (interval-censoring). Observations may also be right-censored (e.g., due to the end of the study, radio-collar failure, or emigration from the study area). It is important to consider the observation process, since the contribution of each individual’s data to the likelihood will depend on whether data are left-truncated or censored. We estimate age-specific hazards using telemetry data collected in two Phases during a 13-year study of white-tailed deer (Odocoileus virginianus) in northern Minnesota. The hazards estimated from the two methods were similar for the full data set that included 302 adults and 76 neonates (followed since or shortly after birth). However, estimated hazards for early-aged individuals differed considerably for subsets of the data that did not include neonates. We discuss the advantages and disadvantages of these two modeling approaches and also compare the estimators using a short simulation study.     

Simulating correlated count data

In this study we compare two techniques for simulating count-valued random n-vectors Y with specified mean and correlation structure. The first technique is to use a lognormal-Poisson hierarchy (L-P method). A vector of correlated normals Z is generated and transformed to a vector of lognormals X. Then, Y is generated as conditionally independent Poissons with means X _i . The L-P method is simple, fast, and familiar to many researchers. However, the method requires each Y _i to be overdispersed (i.e., σ² > μ), and only low correlations are possible with this method when the variables have small means. We develop a second technique to generate the elements of Y as overlapping sums (OS) of independent X _j ’s (OS method). For example, suppose X, X ₁, and X ₂ are independent. If Y ₁ = X + X ₁ and Y ₂ = X + X ₂, then Y ₁ and Y ₂ are correlated because they share the common component X. A generalized version of the OS method for simulating n-vectors of two-parameter count-valued distributions is presented. The OS method is shown to address some of the shortcomings of the L-P method. In particular, underdispersed random variables can be simulated, and high correlations are feasible even when the means are small. However, negative correlations cannot be simulated with the OS method, and when n > 3, the OS method is more complicated to implement than the L-P method.     

Habitat selection models to account for seasonal persistence in radio telemetry data

Models for the analysis of habitat selection data incorporate covariates in an independent multinomial selections model (McCracken et al. 1998) Ramsey and Usner 2003 and an extension of that model to include a persistence parameter (2003). In both cases, all parameters are assumed to be fixed through time. Radio telemetry data collected for habitat selection studies typically consist of animal relocations through time, suggesting the need for an extension to these models. We use a Bayesian approach that allows for the habitat selection probabilities, persistence parameter, or both, to change with season. These extensions are particularly important when movement patterns are expected to differ seasonally and/or when availabilities of habitats change throughout the study period due to weather or migration. We implement and compare the models using radio telemetry data for westslope cutthroat trout in two streams in eastern Oregon.     

Diffusion about the mean drift location in a biased random walk

Random walks are used to model movement in a wide variety of contexts: from the movement of cells undergoing chemotaxis to the migration of animals. In a two-dimensional biased random walk, the diffusion about the mean drift position is entirely dependent on the moments of the angular distribution used to determine the movement direction at each step. Here we consider biased random walks using several different angular distributions and derive expressions for the diffusion coefficients in each direction based on either a fixed or variable movement speed, and we use these to generate a probability density function for the long-time spatial distribution. We demonstrate how diffusion is typically anisotropic around the mean drift position and illustrate these theoretical results using computer simulations. We relate these results to earlier studies of swimming microorganisms and explain how the results can be generalized to other types of animal movement.     

Fitting animal survival models with temporal random effects

Estimating temporal variance in animal demographic parameters is of particular importance in population biology. We implement the Schall’s algorithm for incorporating temporal random effects in survival models using recovery data. Our frequentist approach is based on a formulation of band-recovery models with random effects as generalized linear mixed models and a linearization of the link function conditional on the random effects. A simulation study shows that our procedure provides unbiased and precise estimates. The method is then implemented on two case studies using recovery data on fish and birds.     

A nonparametric procedure for analyzing repeated measures of spatially correlated data

Many agricultural, biological, and environmental studies involve detecting temporal changes of a response variable, based on data observed at sampling sites in a spatial region and repeatedly over several time points. That is, data are repeated measures over time and are potentially correlated across space. The traditional repeated-measures analysis allows for time dependence but assumes that the observations at different sampling sites are mutually independent, which may not be suitable for field data that are correlated across space. In this paper, a nonparametric large-sample inference procedure is developed to assess the time effects while accounting for the spatial dependence using a block bootstrap. For illustration, the methodology is applied to describe the population changes of root-lesion nematodes over time in a production field in Wisconsin.     

Hierarchical Bayesian strategy for modeling correlated compositional data with observed zero counts

This article proposes a hierarchical multivariate conditional autoregressive model applied to a compositional response vector. We particularly focus on situations when the composition is discrete occurring when observations are based on small multinomial counts. We address drawbacks that exist in current modeling approaches for such data. Our hierarchical model will be demonstrated with data used to help manage a commercial sockeye salmon fishery in the Fraser River of British Columbia.     

Statistics for correlated data: phylogenies, space, and time.

Here we give an introduction to the growing number of statistical techniques for analyzing data that are not independent realizations of the same sampling process--in other words, correlated data. We focus on regression problems, in which the value of a given variable depends linearly on the value of another variable. To illustrate different types of processes leading to correlated data, we analyze four simulated examples representing diverse problems arising in ecological studies. The first example is a comparison among species to determine the relationship between home-range area and body size; because species are phylogenetically related, they do not represent independent samples. The second example addresses spatial variation in net primary production and how this might be affected by soil nitrogen; because nearby locations are likely to have similar net primary productivity for reasons other than soil nitrogen, spatial correlation is likely. In the third example, we consider a time-series model to ask whether the decrease in density of a butterfly species is the result of decreases in its host-plant density; because the population density of a species in one generation is likely to affect the density in the following generation, time-series data are often correlated. The fourth example combines both spatial and temporal correlation in an experiment in which prey densities are manipulated to determine the response of predators to their food supply. For each of these examples, we use a different statistical approach for analyzing models of correlated data. Our goal is to give an overview of conceptual issues surrounding correlated data, rather than a detailed tutorial in how to apply different statistical techniques. By dispelling some of the mystery behind correlated data, we hope to encourage ecologists to learn about statistics that could be useful in their own work. Although at first encounter these techniques might seem complicated, they have the power to simplify ecological research by making more types of data and experimental designs open to statistical evaluation.     

Modified Weibull multi-state models for the analysis of animal carcinogenicity data

The Peto test is the standard method of analysis used in carcinogenicity studies to compare tumor incidence in groups of animals. It assumes that tumors are either instantly fatal or have no effect on mortality and requires a judgement of the lethality of each tumor. To avoid this requirement, parametric multi-state models have been proposed. In addition these allow estimation of tumor onset and mortality rates. This paper considers two such models and presents a modification. It is shown that the modified models provide a better fit to carcinogenicity data and simulated data are used to show that the modified models provide a modest increase in test power relative to the Peto test.     

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.     

A hierarchical model for spatial capture-recapture data

Estimating density is a fundamental objective of many animal population studies. Application of methods for estimating population size from ostensibly closed populations is widespread, but ineffective for estimating absolute density because most populations are subject to short-term movements or so-called temporary emigration. This phenomenon invalidates the resulting estimates because the effective sample area is unknown. A number of methods involving the adjustment of estimates based on heuristic considerations are in widespread use. In this paper, a hierarchical model of spatially indexed capture-recapture data is proposed for sampling based on area searches of spatial sample units subject to uniform sampling intensity. The hierarchical model contains explicit models for the distribution of individuals and their movements, in addition to an observation model that is conditional on the location of individuals during sampling. Bayesian analysis of the hierarchical model is achieved by the use of data augmentation, which allows for a straightforward implementation in the freely available software WinBUGS. We present results of a simulation study that was carried out to evaluate the operating characteristics of the Bayesian estimator under variable densities and movement patterns of individuals. An application of the model is presented for survey data on the flat-tailed horned lizard (Phrynosoma mcallii) in Arizona, USA.     

Eigenanalysis of selection ratios from animal radio-tracking data

The development of methods to analyze habitat selection when resources are defined by several categories (e.g., vegetation types) is a topical issue in radio-tracking studies. The White and Garrott statistic, an extension of the widely used test of Neu et al., can be used to determine whether habitat selection is significant. As well, Manly's selection ratio, a particularly useful measure of resource selectivity by resource users, allows detection of the most strongly selected habitat types. However, when both the number of animals and types of habitat are large, the biologist often has to deal with an excessively large number of measures. In this paper we present a new method, the eigenanalysis of selection ratios, that generalizes these two common methods within the framework of eigenanalyses. This method undertakes an additive linear partitioning of the White and Garrott statistic, so that the difference between habitat use and availability is maximized on the first factorial axes. The eigenanalysis of selection ratios is therefore optimal in habitat selection studies. Although we primarily consider the case where the habitat availability is the same for all animals (design II), we also extend this analysis to the case where the habitat availability varies from one animal to another (design III). An application of this method is provided using radio-tracking data collected on 17 squirrels in five habitat types. The results indicate variability in habitat selection, with two groups of animals displaying two patterns of preference. This difference between the two groups is explained by the patch structure of the study area. Because this method is mainly exploratory, and therefore does not rely on any distributional assumption, we recommend its use in studies of habitat selection.     

Analysis of correlated circular and extremal data with a flexible cylindrical distribution

Environmental and Ecological Statistics - In this article, we introduce a flexible cylindrical distribution for modeling and analysis of dependent extremal and directional observations. The...     

Rank regression analysis of correlated water quality data from South East Queensland

With growing population and fast urbanization in Australia, it is a challenging task to maintain our water quality. It is essential to develop an appropriate statistical methodology in analyzing water quality data in order to draw valid conclusions and hence provide useful advices in water management. This paper is to develop robust rank-based procedures for analyzing nonnormally distributed data collected over time at different sites. To take account of temporal correlations of the observations within sites, we consider the optimally combined estimating functions proposed by Wang and Zhu (Biometrika, 93:459–464, 2006) which leads to more efficient parameter estimation. Furthermore, we apply the induced smoothing method to reduce the computational burden. Smoothing leads to easy calculation of the parameter estimates and their variance-covariance matrix. Analysis of water quality data from Total Iron and Total Cyanophytes shows the differences between the traditional generalized linear mixed models and rank regression models. Our analysis also demonstrates the advantages of the rank regression models for analyzing nonnormal data.     

Antispermatogenic effects of artemether: An animal model

Various antimalarial drugs have been shown to exert different adverse effects; however, scanty information is available for artemether-induced potential side effects. The present study assessed effects of artemether on lipid profile, sperm count, and histological features of testes in an animal model. The mean total cholesterol, high-density lipoproteins, low-density lipoproteins, triglyceride, and total proteins in mice-administered artemether were higher compared with controls. The mean sperm counts in mice treated with artemether were reduced when compared with controls. In addition, it was observed that artemether affected the histopathology of seminiferous epithelia and Leydig cells. Evidence indicates that artemether exerts adverse effects in mice testes.     

Regression analysis of correlated circular data based on the multivariate von Mises distribution

Environmental and Ecological Statistics - A regression model for correlated circular data is proposed by assuming that samples of angular measurements are drawn from a multivariate von Mises...     

Bayesian testing for trend in a power model for clustered binary data

Developmental toxicity studies are widely used to investigate the potential risk of environmental hazards. In dose–response experiments, subjects are randomly allocated to groups receiving various dose levels. Tests for trend are then often applied to assess possible dose effects. Recent techniques for risk assessment in this area are based on fitting dose–response models. The complexity of such studies implies a number of non-trivial challenges for model development and the construction of dose-related trend tests, including the hierarchical structure of the data, litter effects inducing extra variation, the functional form of the dose–response curve, the adverse event at dam or at fetus level, the inference paradigm, etc. The purpose of this paper is to propose a Bayesian trend test based on a non-linear power model for the dose effect and using an appropriate model for clustered binary data. Our work is motivated by the analysis of developmental toxicity studies, in which the offspring of exposed and control rodents are examined for defects. Simulations show the performance of the method over a number of samples generated under typical experimental conditions.