A framework for integrating inferred movement behavior into disease risk models.

Movement behavior is an important contributor to habitat selection and its incorporation in disease risk models has been somewhat neglected. The habitat preferences of host individuals affect their probability of exposure to pathogens. If preference behavior can be incorporated in ecological niche models (ENMs) when data on pathogen distributions are available, then variation in such behavior may dramatically impact exposure risk. Here we use data from the anthrax endemic system of Etosha National Park, Namibia, to demonstrate how integrating inferred movement behavior alters the construction of disease risk maps. We used a Maximum Entropy (MaxEnt) model that associated soil, bioclimatic, and vegetation variables with the best available pathogen presence data collected at anthrax carcass sites to map areas of most likely Bacillus anthracis (the causative bacterium of anthrax) persistence. We then used a hidden Markov model (HMM) to distinguish foraging and non-foraging behavioral states along the movement tracks of nine zebra (Equus quagga) during the 2009 and 2010 anthrax seasons. The resulting tracks, decomposed on the basis of the inferred behavioral state, formed the basis of step-selection functions (SSFs) that used the MaxEnt output as a potential predictor variable. Our analyses revealed different risks of exposure during different zebra behavioral states, which were obscured when the full movement tracks were analyzed without consideration of the underlying behavioral states of individuals. Pathogen (or vector) distribution models may be misleading with regard to the actual risk faced by host animal populations when specific behavioral states are not explicitly accounted for in selection analyses. To more accurately evaluate exposure risk, especially in the case of environmentally transmitted pathogens, selection functions could be built for each identified behavioral state and then used to assess the comparative exposure risk across relevant states. The scale of data collection and analysis, however, introduces complexities and limitations for consideration when interpreting results.

Going through the motions: incorporating movement analyses into disease research.

Though epidemiology dates back to the 1700s, most mathematical representations of epidemics still use transmission rates averaged at the population scale, especially for wildlife diseases. In simplifying the contact process, we ignore the heterogeneities in host movements that complicate the real world, and overlook their impact on spatiotemporal patterns of disease burden. Movement ecology offers a set of tools that help unpack the transmission process, letting researchers more accurately model how animals within a population interact and spread pathogens. Analytical techniques from this growing field can also help expose the reverse process: how infection impacts movement behaviours, and therefore other ecological processes like feeding, reproduction, and dispersal. Here, we synthesise the contributions of movement ecology in disease research, with a particular focus on studies that have successfully used movement-based methods to quantify individual heterogeneity in exposure and transmission risk. Throughout, we highlight the rapid growth of both disease and movement ecology and comment on promising but unexplored avenues for research at their overlap. Ultimately, we suggest, including movement empowers ecologists to pose new questions, expanding our understanding of host-pathogen dynamics and improving our predictive capacity for wildlife and even human diseases.

Suite of simple metrics reveals common movement syndromes across vertebrate taxa.

BACKGROUND: Because empirical studies of animal movement are most-often site- and species-specific, we lack understanding of the level of consistency in movement patterns across diverse taxa, as well as a framework for quantitatively classifying movement patterns. We aim to address this gap by determining the extent to which statistical signatures of animal movement patterns recur across ecological systems. We assessed a suite of movement metrics derived from GPS trajectories of thirteen marine and terrestrial vertebrate species spanning three taxonomic classes, orders of magnitude in body size, and modes of movement (swimming, flying, walking). Using these metrics, we performed a principal components analysis and cluster analysis to determine if individuals organized into statistically distinct clusters. Finally, to identify and interpret commonalities within clusters, we compared them to computer-simulated idealized movement syndromes representing suites of correlated movement traits observed across taxa (migration, nomadism, territoriality, and central place foraging). RESULTS: Two principal components explained 70% of the variance among the movement metrics we evaluated across the thirteen species, and were used for the cluster analysis. The resulting analysis revealed four statistically distinct clusters. All simulated individuals of each idealized movement syndrome organized into separate clusters, suggesting that the four clusters are explained by common movement syndrome. CONCLUSIONS: Our results offer early indication of widespread recurrent patterns in movement ecology that have consistent statistical signatures, regardless of taxon, body size, mode of movement, or environment. We further show that a simple set of metrics can be used to classify broad-scale movement patterns in disparate vertebrate taxa. Our comparative approach provides a general framework for quantifying and classifying animal movements, and facilitates new inquiries into relationships between movement syndromes and other ecological processes.

Paradigms for parasite conservation.

Parasitic species, which depend directly on host species for their survival, represent a major regulatory force in ecosystems and a significant component of Earth's biodiversity. Yet the negative impacts of parasites observed at the host level have motivated a conservation paradigm of eradication, moving us farther from attainment of taxonomically unbiased conservation goals. Despite a growing body of literature highlighting the importance of parasite-inclusive conservation, most parasite species remain understudied, underfunded, and underappreciated. We argue the protection of parasitic biodiversity requires a paradigm shift in the perception and valuation of their role as consumer species, similar to that of apex predators in the mid-20th century. Beyond recognizing parasites as vital trophic regulators, existing tools available to conservation practitioners should explicitly account for the unique threats facing dependent species. We built upon concepts from epidemiology and economics (e.g., host-density threshold and cost-benefit analysis) to devise novel metrics of margin of error and minimum investment for parasite conservation. We define margin of error as the risk of accidental host extinction from misestimating equilibrium population sizes and predicted oscillations, while minimum investment represents the cost associated with conserving the additional hosts required to maintain viable parasite populations. This framework will aid in the identification of readily conserved parasites that present minimal health risks. To establish parasite conservation, we propose an extension of population viability analysis for host-parasite assemblages to assess extinction risk. In the direst cases, ex situ breeding programs for parasites should be evaluated to maximize success without undermining host protection. Though parasitic species pose a considerable conservation challenge, adaptations to conservation tools will help protect parasite biodiversity in the face of an uncertain environmental future.

Patch-use dynamics by a large herbivore.

BACKGROUND: An adaption of the optimal foraging theory suggests that herbivores deplete, depart, and finally return to foraging patches leaving time for regrowth [van Moorter et al., Oikos 118:641-652, 2009]. Inter-patch movement and memory of patches then produce a periodic pattern of use that may define the bounds of a home range. The objective of this work was to evaluate the underlying movements within home ranges of elk (Cervus elaphus) according to the predictions of this theory. Using a spatial temporal permutation scan statistic to identify foraging patches from GPS relocations of cow elk, we evaluated return patterns to foraging patches during the 2012 growing season. Subsequently, we used negative binomial regression to assess environmental characteristics that affect the frequency of returns, and thereby characterize the most successful patches. RESULTS: We found that elk return to known patches regularly over a season, on average after 15.4 (±5.4 SD) days. Patches in less-rugged terrain, farther from roads and with high productivity were returned to most often when controlling for the time each patch was known to each elk. CONCLUSIONS: Instead of diffusion processes often used to describe animal movement, our research demonstrates that elk make directed return movements to valuable foraging sites and, as support for Van Moorter et al.'s [Oikos 118:641-652, 2009] model, we submit that these movements could be an integral part of home-range development in wild ungulates.