State-space models (SSMs) are increasingly used in ecology to model time-series such as animal movement paths and population dynamics. This type of hierarchical model is often structured to account for two levels of variability: biological stochasticity and measurement error. SSMs are flexible. They can model linear and nonlinear processes using a variety of statistical distributions. Recent ecological SSMs are often complex, with a large number of parameters to estimate. Through a simulation study, we show that even simple linear Gaussian SSMs can suffer from parameter- and state-estimation problems. We demonstrate that these problems occur primarily when measurement error is larger than biological stochasticity, the condition that often drives ecologists to use SSMs. Using an animal movement example, we show how these estimation problems can affect ecological inference. Biased parameter estimates of a SSM describing the movement of polar bears (Ursus maritimus) result in overestimating their energy expenditure. We suggest potential solutions, but show that it often remains difficult to estimate parameters. While SSMs are powerful tools, they can give misleading results and we urge ecologists to assess whether the parameters can be estimated accurately before drawing ecological conclusions from their results.
State-space models (SSM) are often used for analyzing complex ecological processes that are not observed directly, such as marine animal movement. When outliers are present in the measurements, special care is needed in the analysis to obtain reliable location and process estimates. Here we recommend using the Laplace approximation combined with automatic differentiation (as implemented in the novel R package Template Model Builder; TMB) for the fast fitting of continuous-time multivariate non-Gaussian SSMs. Through Argos satellite tracking data, we demonstrate that the use of continuous-time t-distributed measurement errors for error-prone data is more robust to outliers and improves the location estimation compared to using discretized-time t-distributed errors (implemented with a Gibbs sampler) or using continuous-time Gaussian errors (as with the Kalman filter). Using TMB, we are able to estimate additional parameters compared to previous methods, all without requiring a substantial increase in computational time. The model implementation is made available through the R package argosTrack.
Tracking of marine animals has increased exponentially in the past decade, and the resulting data could lead to an in-depth understanding of the causes and consequences of movement in the ocean. However, most common marine tracking systems are associated with large measurement errors. Accounting for these errors requires the use of hierarchical models, which are often difficult to fit to data. Using 3 case studies, we demonstrate that Template Model Builder (TMB), a new R package, is an accurate, efficient and flexible framework for modelling movement data. First, to demonstrate that TMB is as accurate but 30 times faster than bsam, a popular R package used to apply state-space models to Argos data, we modelled polar bear Ursus maritimus Argos data and compared the locations estimated by the models to GPS locations of these same bears. Second, to demonstrate how TMB's gain in efficiency and frequentist framework facilitate model comparison, we developed models with different error structures and compared them to find the most effective model for light-based geolocations of rhinoceros auklets Cerorhinca monocerata. Finally, to maximize efficiency through TMB's use of the Laplace approximation of the marginal likelihood, we modelled behavioural changes with continuous rather than discrete states. This new model directly accounts for the irregular sampling intervals characteristic of Fastloc-GPS data of grey seals Halichoerus grypus. Using real and simulated data, we show that TMB is a fast and powerful tool for modelling marine movement data. We discuss how TMB's potential reaches beyond marine movement studies.
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Electronic telemetry is frequently used to document animal movement through time. Methods that can identify underlying behaviors driving specific movement patterns can help us understand how and why animals use available space, thereby aiding conservation and management efforts. For aquatic animal tracking data with significant measurement error, a Bayesian state‐space model called the first‐Difference Correlated Random Walk with Switching (DCRWS) has often been used for this purpose. However, for aquatic animals, highly accurate tracking data are now becoming more common. We developed a new hidden Markov model (HMM) for identifying behavioral states from animal tracks with negligible error, called the hidden Markov movement model (HMMM). We implemented as the basis for the HMMM the process equation of the DCRWS, but we used the method of maximum likelihood and the R package for rapid model fitting. The HMMM was compared to a modified version of the DCRWS for highly accurate tracks, the DCRWSNOME, and to a common HMM for animal tracks fitted with the R package . We show that the HMMM is both accurate and suitable for multiple species by fitting it to real tracks from a grey seal, lake trout, and blue shark, as well as to simulated data. The HMMM is a fast and reliable tool for making meaningful inference from animal movement data that is ideally suited for ecologists who want to use the popular DCRWS implementation and have highly accurate tracking data. It additionally provides a groundwork for development of more complex modeling of animal movement with . To facilitate its uptake, we make it available through the R package .
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