Ian D. Jonsen

Tracking apex marine predator movements in a dynamic ocean

Pelagic marine predators face unprecedented challenges and uncertain futures. Overexploitation and climate variability impact the abundance and distribution of top predators in ocean ecosystems. Improved understanding of ecological patterns, evolutionary constraints and ecosystem function is critical for preventing extinctions, loss of biodiversity and disruption of ecosystem services. Recent advances in electronic tagging techniques have provided the capacity to observe the movements and long-distance migrations of animals in relation to ocean processes across a range of ecological scales. Tagging of Pacific Predators, a field programme of the Census of Marine Life, deployed 4,306 tags on 23 species in the North Pacific Ocean, resulting in a tracking data set of unprecedented scale and species diversity that covers 265,386 tracking days from 2000 to 2009. Here we report migration pathways, link ocean features to multispecies hotspots and illustrate niche partitioning within and among congener guilds. Our results indicate that the California Current large marine ecosystem and the North Pacific transition zone attract and retain a diverse assemblage of marine vertebrates. Within the California Current large marine ecosystem, several predator guilds seasonally undertake north–south migrations that may be driven by oceanic processes, species-specific thermal tolerances and shifts in prey distributions. We identify critical habitats across multinational boundaries and show that top predators exploit their environment in predictable ways, providing the foundation for spatial management of large marine ecosystems.

ROBUST STATE–SPACE MODELING OF ANIMAL MOVEMENT DATA

Remotely sensed tracking data collected on animal movement is vastly un- derutilized due to a lack of statistical tools for appropriate analysis. Features of such data that make analysis particularly challenging include the presence of estimation errors that are non-Gaussian and vary in time, observations that occur irregularly in time, and com- plexity in the underlying behavioral processes. We develop a state-space framework that simultaneously deals with these features and demonstrate our method by analyzing three seal pathway data sets. We show how known information regarding error distributions can be used to improve inference of the underlying process(es) and demonstrate that our frame- work provides a powerful and flexible method for fitting different behavioral models to tracking data.

META‐ANALYSIS OF ANIMAL MOVEMENT USING STATE‐SPACE MODELS

The study of animal movement and behavior is being revolutionized by technology, such as satellite tags and harmonic radar, that allows us to track the movements of individual animals. However, our ability to analyze and model such data has lagged behind the sophisticated collection methods. We review problems with current methods and suggest a more powerful and flexible approach, state-space modeling, and we illustrate how these models can be posed in a meta-analytic framework so that information from individual trajectories may be combined optimally. State-space models enable us to deal with the complexity of modeling animals interacting with their environment but, unlike other methods, they allow simultaneous estimation of measurement error and process noise that are inherent in animal-trajectory data. A Bayesian framework allows us to incorporate important prior information when available and also allows meta-analytic techniques to be incorporated in a straightforward fashion. Meta-analysis enables both individual and broader-level inference from observations of multiple individual pathways. Our approach is powerful because it allows researchers to test hypotheses regarding animal movement, to connect theoretical models to data, and to use modern likelihood-based estimation techniques, all under a single statistical framework.

Response of generalist and specialist insect herbivores to landscape spatial structure

The purpose of this study was to investigate the effect ofchanges in landscape pattern on generalist and specialistinsects. We did this by comparing the species richness andabundance of generalist and specialist herbivorous insects inalfalfa (Medicago sativa, L.) fields on 26 agriculturallandscapes that differed in spatial structure. The insects werefrom the families Curculionidae (Coleoptera), weevils, andCicadellidae (Auchennorhyncha), leafhoppers.We hypothesized that: (1) generalist richness and abundancewould be highest in landscapes with high diversity(Shannon-Wiener); (2) specialist richness and abundance would behighest in landscapes with (i) high percent cover alfalfa and(ii) low mean inter-patch distance. We tested for these effectsafter controlling for the patch-level effects of field size,field age, frequency of disturbance and vegetation texture.The important findings of the study are: (1) generalist richness andabundance increased with increasing landscape diversity and (2)isolation (percent cover alfalfa in the landscape and/or meaninter-patch distance) does not affect specialist insects. Theseresults are significant because they indicate that bothgeneralist and specialist insects may move over much largerdistances than previously thought. This is one of the firststudies to demonstrate a large scale effect of spatial structureon insects across a broad range of landscapes.

Sex-specific, seasonal foraging tactics of adult grey seals (Halichoerus grypus) revealed by state-space analysis

In many large pelagic animals, observing behavior is limited to observation by radio or satellite telemetry. In many cases, discriminating different behaviors from telemetry data has been a key, but often elusive, goal. Here we use state-space models (SSMs) to fit a correlated random walk (CRW) model that switches between two unobserved behavioral states (nominally foraging and traveling) to 41 male and 43 female adult grey seal (Halichoerus grypus) satellite telemetry tracks. The SSM results reveal markedly different spatial behavior between the sexes, fitting well with sexual size dimorphism and known dietary differences, suggesting that the sexes deal with seasonal prey availability and reproductive costs differently. From these results we were also able to produce behaviorally informed habitat use maps, showing a complex and dynamic network of small, intensely used foraging areas. Our flexible SSM approach clearly demonstrates sex-related behavioral differences, fine scale spatial and temporal foraging patterns, and a clearer picture of grey seal ecology and role in the Scotian Shelf ecosystem.

Key Questions in Marine Megafauna Movement Ecology

It is a golden age for animal movement studies and so an opportune time to assess priorities for future work. We assembled 40 experts to identify key questions in this field, focussing on marine megafauna, which include a broad range of birds, mammals, reptiles, and fish. Research on these taxa has both underpinned many of the recent technical developments and led to fundamental discoveries in the field. We show that the questions have broad applicability to other taxa, including terrestrial animals, flying insects, and swimming invertebrates, and, as such, this exercise provides a useful roadmap for targeted deployments and data syntheses that should advance the field of movement ecology.

Effect of habitat patch characteristics on abundance and diversity of insects in an agricultural landscape

ABSTRACT The objective of this study was to test for general effects of patch size, patch isolation, disturbance frequency, and patch life span, on density and diversity of organisms. We sampled predominantly herbivorous insects in 31 alfalfa fields that varied in size, isolation, frequency of disturbance by cutting, and age (number of years planted in alfalfa). Effects on insect relative density and diversity were examined at three taxonomic levels: all insects, eight separate orders, and six legume-specialist weevil species. We found that (a) more isolated alfalfa fields had higher overall insect richness, (b) fields with higher disturbance frequency had lower overall insect richness, and (c) fields of intermediate age had highest insect richness. In some cases these patterns were reflected at lower taxonomic levels, but in many cases they were not. These results are important because they indicate that, although we cannot simultaneously tailor a landscape for each of thousands of species, we may be able to produce desired effects at a more general level.

The influence of matrix habitat on Aphthona flea beetle immigration to leafy spurge patches

Variation in movement ability by insects among different non-habitat (matrix) types may have important implications for both metapopulation dynamics and weed biocontrol practices. We used a mark-recapture experiment to explore the effects of two different matrix habitats (grass vs shrub) on the ability of two species of Aphthona (Chrysomelidae: Coleoptera) flea beetle to immigrate to patches of the invasive weed, leafy spurge. Using generalized linear models, we compared effects of the matrix habitat types, species and sex on observed immigration probabilities. Our analyses demonstrated that one species (A. nigriscutis) had a much higher immigration probability when moving through a grass-dominated matrix than a shrub-dominated matrix whereas immigration probabilities for the second species (A. lacertosa) were similar in both matrix habitats but significantly lower overall than for A. nigriscutis. Furthermore, A. nigriscutis females were more likely to immigrate to spurge patches embedded in a grass matrix than in shrub, whereas the opposite occurred for males. Our results suggest that metapopulation dynamics may be strongly affected by the type(s) of matrix habitat present on a landscape. These effects also suggest that release strategies for weed biocontrol should be tailored according to the structure of the landscape into which releases are planned. In addition, even closely related species can have significantly different movement abilities which will also affect release strategies.

Integrative modelling of animal movement: incorporating in situ habitat and behavioural information for a migratory marine predator

A fundamental goal in animal ecology is to quantify how environmental (and other) factors influence individual movement, as this is key to understanding responsiveness of populations to future change. However, quantitative interpretation of individual-based telemetry data is hampered by the complexity of, and error within, these multi-dimensional data. Here, we present an integrative hierarchical Bayesian state-space modelling approach where, for the first time, the mechanistic process model for the movement state of animals directly incorporates both environmental and other behavioural information, and observation and process model parameters are estimated within a single model. When applied to a migratory marine predator, the southern elephant seal (Mirounga leonina), we find the switch from directed to resident movement state was associated with colder water temperatures, relatively short dive bottom time and rapid descent rates. The approach presented here can have widespread utility for quantifying movement–behaviour (diving or other)–environment relationships across species and systems.

North Atlantic blue and fin whales suspend their spring migration to forage in middle latitudes: building up energy reserves for the journey?

The need to balance energy reserves during migration is a critical factor for most long-distance migrants and an important determinant of migratory strategies in birds, insects and land mammals. Large baleen whales migrate annually between foraging and breeding sites, crossing vast ocean areas where food is seldom abundant. How whales respond to the demands and constraints of such long migrations remains unknown. We applied a behaviour discriminating hierarchical state-space model to the satellite tracking data of 12 fin whales and 3 blue whales tagged off the Azores, to investigate their movements, behaviour (transiting and area-restricted search, ARS) and daily activity cycles during the spring migration. Fin and blue whales remained at middle latitudes for prolonged periods, spending most of their time there in ARS behaviour. While near the Azores, fin whale ARS behaviour occurred within a restricted area, with a high degree of overlap among whales. There were noticeable behavioural differences along the migratory pathway of fin whales tracked to higher latitudes: ARS occurred only in the Azores and north of 56°N, whereas in between these areas whales travelled at higher overall speeds while maintaining a nearly direct trajectory. This suggests fin whales may alternate periods of active migration with periods of extended use of specific habitats along the migratory route. ARS behaviour in blue whales occurred over a much wider area as whales slowly progressed northwards. The tracks of these whales terminated still at middle latitudes, before any behavioural switch was detected. Fin whales exhibited behavioural-specific diel rhythms in swimming speed but these varied significantly between geographic areas, possibly due to differences in the day-night cycle across areas. Finally, we show a link between fin whales seen in the Azores and those summering in eastern Greenland-western Iceland along a migratory corridor located in central Atlantic waters.