Recent developments in animal tracking technology have permitted the collection of detailed data on the movement paths of individuals from many species. However, analysis methods for these data have ...not developed at a similar pace, largely due to a lack of suitable candidate models, coupled with the technical difficulties of fitting such models to data. To facilitate a general modeling framework, we propose that complex movement paths can be conceived as a series of movement strategies among which animals transition as they are affected by changes in their internal and external environment. We synthesize previously existing and novel methodologies to develop a general suite of mechanistic models based on biased and correlated random walks that allow different behavioral states for directed (e.g., migration), exploratory (e.g., dispersal), area-restricted (e.g., foraging), and other types of movement. Using this "toolbox" of nested model components, multistate movement models may be custom-built for a wide variety of species and applications. As a unified state-space modeling framework, it allows the simultaneous investigation of numerous hypotheses about animal movement from imperfectly observed data, including time allocations to different movement behavior states, transitions between states, the use of memory or navigation, and strengths of attraction (or repulsion) to specific locations. The inclusion of covariate information permits further investigation of specific hypotheses related to factors driving different types of movement behavior. Using reversible-jump Markov chain Monte Carlo methods to facilitate Bayesian model selection and multi-model inference, we apply the proposed methodology to real data by adapting it to the natural history of the grey seal (
Halichoerus grypus
) in the North Sea. Although previous grey seal studies tended to focus on correlated movements, we found overwhelming evidence that bias toward haul-out or foraging locations better explained seal movement than did simple or correlated random walks. Posterior model probabilities also provided evidence that seals transition among directed, area-restricted, and exploratory movements associated with haul-out, foraging, and other behaviors. With this intuitive framework for modeling and interpreting animal movement, we believe that the development and application of custom-made movement models will become more accessible to ecologists and non-statisticians.
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.
Hidden Markov models: Pitfalls and opportunities in ecology Glennie, Richard; Adam, Timo; Leos‐Barajas, Vianey ...
Methods in ecology and evolution,
January 2023, 2023-01-00, 20230101, 2023-01-01, Letnik:
14, Številka:
1
Journal Article
Recenzirano
Odprti dostop
Hidden Markov models (HMMs) and their extensions are attractive methods for analysing ecological data where noisy, multivariate measurements are made of a hidden, ecological process, and where this ...hidden process is represented by a sequence of discrete states. Yet, as these models become more complex and challenging to understand, it is important to consider what pitfalls these methods have and what opportunities there are for future research to address these pitfalls.
In this paper, we review five lesser known pitfalls one can encounter when using HMMs or their extensions to solve ecological problems: (a) violation of the snapshot property in continuous‐time HMMs; (b) biased inference from hierarchical HMMs when applied to temporally misaligned processes; (c) sensitive inference from using random effects to partially pool across heterogeneous individuals; (d) computational burden when using HMMs to approximate models with continuous state spaces; and (e) difficulty linking the hidden process to space or environment.
This review is for ecologists and ecological statisticians familiar with HMMs, but who may be less aware of the problems that arise in more specialised applications. We demonstrate how each pitfall arises, by simulation or example, and discuss why this pitfall is important to consider. Along with identifying the problems, we highlight potential research opportunities and offer ideas that may help alleviate these pitfalls.
Each of the methods we review are solutions to current ecological research problems. We intend for this paper to heighten awareness of the pitfalls ecologists may encounter when applying these more advanced methods, but we also hope that by highlighting future research opportunities, we can inspire ecological statisticians to weaken these pitfalls and provide improved methods.
Recent technological advances have permitted the collection of detailed animal location and ancillary biotelemetry data that facilitate inference about animal movement and associated behaviors. ...However, these rich sources of individual information, location, and biotelemetry data, are typically analyzed independently, with population-level inferences remaining largely post hoc. We describe a hierarchical modeling approach, which is able to integrate location and ancillary biotelemetry (e.g., physiological or accelerometer) data from many individuals. We can thus obtain robust estimates of (1) population-level movement parameters and (2) activity budgets for a set of behaviors among which animals transition as they respond to changes in their internal and external environment. Measurement error and missing data are easily accommodated using a state-space formulation of the proposed hierarchical model. Using Bayesian analysis methods, we demonstrate our modeling approach with location and dive activity data from 17 harbor seals (
Phoca vitulina
) in the United Kingdom. Based jointly on movement and diving activity, we identified three distinct movement behavior states: resting, foraging, and transit, and estimated population-level activity budgets to these three states. Because harbor seals are known to dive for both foraging and transit (but not usually for resting), we compared these results to a similar population-level analysis utilizing only location data. We found that a large proportion of time steps were mischaracterized when behavior states were inferred from horizontal trajectory alone, with 33% of time steps exhibiting a majority of dive activity assigned to the resting state. Only 1% of these time steps were assigned to resting when inferred from both trajectory and dive activity data using our integrated modeling approach. There is mounting evidence of the potential perils of inferring animal behavior based on trajectory alone, but there fortunately now exist many flexible analytical techniques for extracting more out of the increasing wealth of information afforded by recent advances in biologging technology.
The cosmological utility of galaxy cluster catalogues is primarily limited by our ability to calibrate the relation between halo mass and observable mass proxies such as cluster richness, X-ray ...luminosity or the Sunyaev-Zeldovich signal. Projection effects are a particularly pernicious systematic effect that can impact observable mass proxies; structure along the line of sight can both bias and increase the scatter of the observable mass proxies used in cluster abundance studies. In this work, we develop an empirical method to characterize the impact of projection effects on redMaPPer cluster catalogues. We use numerical simulations to validate our method and illustrate its robustness. We demonstrate that modeling of projection effects is a necessary component for cluster abundance studies capable of reaching $\approx 5\%$ mass calibration uncertainties (e.g. the Dark Energy Survey Year 1 sample). Specifically, ignoring the impact of projection effects in the observable--mass relation --- i.e. marginalizing over a log-normal model only --- biases the posterior of the cluster normalization condition $S_8 \equiv \sigma_8 (\Omega_{\rm m}/0.3)^{1/2}$ by $\Delta S_8 =0.05$, more than twice the uncertainty in the posterior for such an analysis.
The population densities of leopards vary widely across their global range, influenced by prey availability, intraguild competition and human persecution. In Asia, particularly the Middle East and ...the Caucasus, they generally occur at the lower extreme of densities recorded for the species. Reliable estimates of population density are important for understanding their ecology and planning their conservation. We used a photographic spatial capture-recapture (SCR) methodology incorporating animal movement to estimate density for the endangered Persian leopard Panthera pardus saxicolor in three montane national parks, northeastern Iran. We combined encounter history data arising from images of bilaterally asymmetrical left- and right-sided pelage patterns using a Bayesian spatial partial identity model accommodating multiple "non-invasive" marks. We also investigated the effect of camera trap placement on detection probability. Surprisingly, considering the subspecies' reported low abundance and density based on previous studies, we found relatively high population densities in the three national parks, varying between 3.10 ± SD 1.84 and 8.86 ± SD 3.60 individuals/100 km
. The number of leopards detected in Tandoureh National Park (30 individuals) was larger than estimated during comparable surveys at any other site in Iran, or indeed globally. Capture and recapture probabilities were higher for camera traps placed near water resources compared with those placed on trails. Our results show the benefits of protecting even relatively small mountainous areas, which accommodated a high density of leopards and provided refugia in a landscape with substantial human activity.
Abundance and population density are fundamental pieces of information for population ecology and species conservation, but they are difficult to estimate for rare and elusive species. Mark-resight ...models are popular for estimating population abundance because they are less invasive and expensive than traditional mark-recapture. However, density estimation using mark-resight is difficult because the area sampled must be explicitly defined, historically using ad hoc approaches. We developed a spatial mark-resight model for estimating population density that combines spatial resighting data and telemetry data. Incorporating telemetry data allows us to inform model parameters related to movement and individual location. Our model also allows <100% individual identification of marked individuals. We implemented the model in a Bayesian framework, using a custom-made Metropolis-within-Gibbs Markov chain Monte Carlo algorithm. As an example, we applied this model to a mark-resight study of raccoons (
Procyon lotor
) on South Core Banks, a barrier island in Cape Lookout National Seashore, North Carolina, USA. We estimated a population of 186.71 ± 14.81 individuals, which translated to a density of 8.29 ± 0.66 individuals/km
2
(mean ± SD). The model presented here will have widespread utility in future applications, especially for species that are not naturally marked.
We present density split statistics, a framework that studies lensing and counts-in-cells as a function of foreground galaxy density, thereby providing a large-scale measurement of both 2-point and ...3-point statistics. Our method extends our earlier work on trough lensing and is summarized as follows: given a foreground (low redshift) population of galaxies, we divide the sky into subareas of equal size but distinct galaxy density. We then measure lensing around uniformly spaced points separately in each of these subareas, as well as counts-in-cells statistics (CiC). The lensing signals trace the matter density contrast around regions of fixed galaxy density. Through the CiC measurements this can be related to the density profile around regions of fixed matter density. Together, these measurements constitute a powerful probe of cosmology, the skewness of the density field and the connection of galaxies and matter. In this paper we show how to model both the density split lensing signal and CiC from basic ingredients: a non-linear power spectrum, clustering hierarchy coefficients from perturbation theory and a parametric model for galaxy bias and shot-noise. Using N-body simulations, we demonstrate that this model is sufficiently accurate for a cosmological analysis on year 1 data from the Dark Energy Survey.
Ecology Letters (2010) 13: 659-674 Analytical methods accounting for imperfect detection are often used to facilitate reliable inference in population and community ecology. We contend that similar ...approaches are needed in disease ecology because these complicated systems are inherently difficult to observe without error. For example, wildlife disease studies often designate individuals, populations, or spatial units to states (e.g., susceptible, infected, post-infected), but the uncertainty associated with these state assignments remains largely ignored or unaccounted for. We demonstrate how recent developments incorporating observation error through repeated sampling extend quite naturally to hierarchical spatial models of disease effects, prevalence, and dynamics in natural systems. A highly pathogenic strain of avian influenza virus in migratory waterfowl and a pathogenic fungus recently implicated in the global loss of amphibian biodiversity are used as motivating examples. Both show that relatively simple modifications to study designs can greatly improve our understanding of complex spatio-temporal disease dynamics by rigorously accounting for uncertainty at each level of the hierarchy.
The recent surge in the development and application of species occurrence models has been associated with an acknowledgment among ecologists that species are detected imperfectly due to observation ...error. Standard models now allow unbiased estimation of occupancy probability when false negative detections occur, but this is conditional on no false positive detections and sufficient incorporation of explanatory variables for the false negative detection process. These assumptions are likely reasonable in many circumstances, but there is mounting evidence that false positive errors and detection probability heterogeneity may be much more prevalent in studies relying on auditory cues for species detection (e.g., songbird or calling amphibian surveys). We used field survey data from a simulated calling anuran system of known occupancy state to investigate the biases induced by these errors in dynamic models of species occurrence. Despite the participation of expert observers in simplified field conditions, both false positive errors and site detection probability heterogeneity were extensive for most species in the survey. We found that even low levels of false positive errors, constituting as little as 1% of all detections, can cause severe overestimation of site occupancy, colonization, and local extinction probabilities. Further, unmodeled detection probability heterogeneity induced substantial underestimation of occupancy and overestimation of colonization and local extinction probabilities. Completely spurious relationships between species occurrence and explanatory variables were also found. Such misleading inferences would likely have deleterious implications for conservation and management programs. We contend that all forms of observation error, including false positive errors and heterogeneous detection probabilities, must be incorporated into the estimation framework to facilitate reliable inferences about occupancy and its associated vital rate parameters.
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