Théo Michelot

moveHMM: an R package for the statistical modelling of animal movement data using hidden Markov models

Summary Due to the substantial progress in tracking technology, recent years have seen an explosion in the amount of movement data being collected. This has led to a huge demand for statistical tools that allow ecologists to draw meaningful inference from large tracking data sets. The class of hidden Markov models (HMMs) matches the intuitive understanding that animal movement is driven by underlying behavioural modes and has proven to be very useful for analysing movement data. For data that involve a regular sampling unit and negligible measurement error, these models usually are sufficiently flexible to capture the complex correlation structure found in movement data, yet are computationally inexpensive compared to alternative methods. The R package moveHMM allows ecologists to process GPS tracking data into series of step lengths and turning angles, and to fit an HMM to these data, allowing, in particular, for the incorporation of environmental covariates. The package includes assessment and visualization tools for the fitted model. We illustrate the use of moveHMM using (simulated) movement of the legendary wild haggis Haggis scoticus. Our findings illustrate the role our software, and movement modelling in general, can play in conservation and management by illuminating environmental constraints.

Understanding the ontogeny of foraging behaviour : insights from combining marine predator bio-logging with satellite-derived oceanography in hidden Markov models

The development of foraging strategies that enable juveniles to efficiently identify and exploit predictable habitat features is critical for survival and long-term fitness. In the marine environment, meso- and sub-mesoscale features such as oceanographic fronts offer a visible cue to enhanced foraging conditions, but how individuals learn to identify these features is a mystery. In this study, we investigate age-related differences in the fine-scale foraging behaviour of adult (aged ≥ 5 years) and immature (aged 2–4 years) northern gannets Morus bassanus. Using high-resolution GPS-loggers, we reveal that adults have a much narrower foraging distribution than immature birds and much higher individual foraging site fidelity. By conditioning the transition probabilities of a hidden Markov model on satellite-derived measures of frontal activity, we then demonstrate that adults show a stronger response to frontal activity than immature birds, and are more likely to commence foraging behaviour as frontal intensity increases. Together, these results indicate that adult gannets are more proficient foragers than immatures, supporting the hypothesis that foraging specializations are learned during individual exploratory behaviour in early life. Such memory-based individual foraging strategies may also explain the extended period of immaturity observed in gannets and many other long-lived species.

Maximum penalized likelihood estimation in semiparametric mark-recapture-recovery models

We discuss the semiparametric modeling of mark-recapture-recovery data where the temporal and/or individual variation of model parameters is explained via covariates. Typically, in such analyses a fixed (or mixed) effects parametric model is specified for the relationship between the model parameters and the covariates of interest. In this paper, we discuss the modeling of the relationship via the use of penalized splines, to allow for considerably more flexible functional forms. Corresponding models can be fitted via numerical maximum penalized likelihood estimation, employing cross-validation to choose the smoothing parameters in a data-driven way. Our contribution builds on and extends the existing literature, providing a unified inferential framework for semiparametric mark-recapture-recovery models for open populations, where the interest typically lies in the estimation of survival probabilities. The approach is applied to two real datasets, corresponding to gray herons (Ardea cinerea), where we model the survival probability as a function of environmental condition (a time-varying global covariate), and Soay sheep (Ovis aries), where we model the survival probability as a function of individual weight (a time-varying individual-specific covariate). The proposed semiparametric approach is compared to a standard parametric (logistic) regression and new interesting underlying dynamics are observed in both cases.

Linking resource selection and step selection models for habitat preferences in animals.

The two dominant approaches for the analysis of species-habitat associations in animals have been shown to reach divergent conclusions. Models fitted from the viewpoint of an individual (step selection functions), once scaled up, do not agree with models fitted from a population viewpoint (resource selection functions [RSFs]). We explain this fundamental incompatibility, and propose a solution by introducing to the animal movement field a novel use for the well-known family of Markov chain Monte Carlo (MCMC) algorithms. By design, the step selection rules of MCMC lead to a steady-state distribution that coincides with a given underlying function: the target distribution. We therefore propose an analogy between the movements of an animal and the movements of an MCMC sampler, to guarantee convergence of the step selection rules to the parameters underlying the populations utilization distribution. We introduce a rejection-free MCMC algorithm, the local Gibbs sampler, that better resembles real animal movement, and discuss the wide range of biological assumptions that it can accommodate. We illustrate our method with simulations on a known utilization distribution, and show theoretically and empirically that locations simulated from the local Gibbs sampler give rise to the correct RSF. Using simulated data, we demonstrate how this framework can be used to estimate resource selection and movement parameters.