Macro-ecology of plankton–seabird associations in the North Pacific Ocean

In conjunction with the North Pacific Continuous Plankton Recorder program, we conducted surveys of seabirds from June 2002 to June 2007. Here, we tested the hypotheses of (i) east–west variations in coupled plankton and seabird abundance, and (ii) that surface-feeding and diving seabirds vary in their relationships to primary productivity and mesozooplankton species abundance and diversity. To test these hypotheses, we developed statistical models for 20 species of seabirds and 12 zooplankton taxonomic groups. Seabird density was highly variable between seasons, but was consistently higher in the western than eastern North Pacific. Seabird diversity was greater in the east. Zooplankton abundance did not differ between regions. We found associations at the “bulk” level between seabird density and net primary productivity, but only one association between seabirds and total zooplankton abundance or diversity. However, we found many relationships between seabird species and the abundance of different zooplankton summarized at the genus or family level. Some of these taxonomic relationships reflect direct predator–prey interactions, while others may reflect zooplankton that serve as ecological indicators of other prey, such as micronekton, upon which the birds may feed. Surface or near-surface feeding, mostly piscivorous seabirds, did not differ systematically from diving, mainly planktivorous seabirds in their zooplankton associations. Seabirds apparently respond to zooplankton taxonomic groupings more so than bulk zooplankton characteristics, such as abundance or diversity. Macro-ecological studies of remote marine ecosystems using zooplankton and seabirds as ecological indicators provide a framework for understanding and assessing spatial and temporal variations in these difficult-to-study pelagic environments.

Sardine cycles, krill declines, and locust plagues: revisiting ‘wasp-waist’ food webs

‘Wasp-waist’ systems are dominated by a mid trophic-level species that is thought to exert top-down control on its food and bottom-up control on its predators. Sardines, anchovy, and Antarctic krill are suggested examples, and here we use locusts to explore whether the wasp-waist concept also applies on land. These examples also display the traits of mobile aggregations and dietary diversity, which help to reduce the foraging footprint from their large, localised biomasses. This suggests that top-down control on their food operates at local aggregation scales and not at wider scales suggested by the original definition of wasp-waist. With this modification, the wasp-waist framework can cross-fertilise marine and terrestrial approaches, revealing how seemingly disparate but economically important systems operate.

Implications of streamlining theory for microbial ecology

Whether a small cell, a small genome or a minimal set of chemical reactions with self-replicating properties, simplicity is beguiling. As Leonardo da Vinci reportedly said, 'simplicity is the ultimate sophistication'. Two diverging views of simplicity have emerged in accounts of symbiotic and commensal bacteria and cosmopolitan free-living bacteria with small genomes. The small genomes of obligate insect endosymbionts have been attributed to genetic drift caused by small effective population sizes (Ne). In contrast, streamlining theory attributes small cells and genomes to selection for efficient use of nutrients in populations where Ne is large and nutrients limit growth. Regardless of the cause of genome reduction, lost coding potential eventually dictates loss of function. Consequences of reductive evolution in streamlined organisms include atypical patterns of prototrophy and the absence of common regulatory systems, which have been linked to difficulty in culturing these cells. Recent evidence from metagenomics suggests that streamlining is commonplace, may broadly explain the phenomenon of the uncultured microbial majority, and might also explain the highly interdependent (connected) behavior of many microbial ecosystems. Streamlining theory is belied by the observation that many successful bacteria are large cells with complex genomes. To fully appreciate streamlining, we must look to the life histories and adaptive strategies of cells, which impose minimum requirements for complexity that vary with niche.

Environmental context explains Lévy and Brownian movement patterns of marine predators

An optimal search theory, the so-called Levy-flight foraging hypothesis(1), predicts that predators should adopt search strategies known as Levy flights where prey is sparse and distributed unpredictably, but that Brownian movement is sufficiently efficient for locating abundant prey(2-4). Empirical studies have generated controversy because the accuracy of statistical methods that have been used to identify Levy behaviour has recently been questioned(5,6). Consequently, whether foragers exhibit Levy flights in the wild remains unclear. Crucially, moreover, it has not been tested whether observed movement patterns across natural landscapes having different expected resource distributions conform to the theory's central predictions. Here we use maximum-likelihood methods to test for Levy patterns in relation to environmental gradients in the largest animal movement data set assembled for this purpose. Strong support was found for Levy search patterns across 14 species of open-ocean predatory fish (sharks, tuna, billfish and ocean sunfish), with some individuals switching between Levy and Brownian movement as they traversed different habitat types. We tested the spatial occurrence of these two principal patterns and found Levy behaviour to be associated with less productive waters (sparser prey) and Brownian movements to be associated with productive shelf or convergence-front habitats (abundant prey). These results are consistent with the Levy-flight foraging hypothesis(1,7), supporting the contention(8,9) that organism search strategies naturally evolved in such a way that they exploit optimal Levy patterns.

A new approach for objective identification of turns and steps in organism movement data relevant to random walk modelling

1. A first step in the analysis of complex movement data often involves discretisation of the path into a series of step-lengths and turns, for example in the analysis of specialised random walks, such as Lévy flights. However, the identification of turning points, and therefore step-lengths, in a tortuous path is dependent on ad-hoc parameter choices. Consequently, studies testing for movement patterns in these data, such as Lévy flights, have generated debate. However, studies focusing on one-dimensional (1D) data, as in the vertical displacements of marine pelagic predators, where turning points can be identified unambiguously have provided strong support for Lévy flight movement patterns. 2. Here, we investigate how step-length distributions in 3D movement patterns would be interpreted by tags recording in 1D (i.e. depth) and demonstrate the dimensional symmetry previously shown mathematically for Lévy-flight movements. We test the veracity of this symmetry by simulating several measurement errors common in empirical datasets and find Lévy patterns and exponents to be robust to low-quality movement data. 3. We then consider exponential and composite Brownian random walks and show that these also project into 1D with sufficient symmetry to be clearly identifiable as such. 4. By extending the symmetry paradigm, we propose a new methodology for step-length identification in 2D or 3D movement data. The methodology is successfully demonstrated in a re-analysis of wandering albatross Global Positioning System (GPS) location data previously analysed using a complex methodology to determine bird-landing locations as turning points in a Lévy walk. For this high-resolution GPS data, we show that there is strong evidence for albatross foraging patterns approximated by truncated Lévy flights spanning over 3·5 orders of magnitude. 5. Our simple methodology and freely available software can be used with any 2D or 3D movement data at any scale or resolution and are robust to common empirical measurement errors. The method should find wide applicability in the field of movement ecology spanning the study of motile cells to humans.

Do oceanic loggerhead turtles Caretta caretta associate with oceanographic fronts? Evidence from the Canary Current Large Marine Ecosystem

ABSTRACT: Oceanographic fronts are physical interfaces between water masses that differ in properties such as temperature, salinity, turbidity and chl a enrichment. Bio-physical coupling along fronts can lead to the development of pelagic biodiversity hotspots. A diverse range of marine vertebrates have been shown to associate with fronts, using them as foraging and migration habitats. Elucidation of the ecological significance of fronts generates a better understanding of marine ecosystem functioning, conferring opportunities to improve management of anthropogenic activities in the oceans. This study presents novel insight into the oceanographic drivers of habitat use in a population of marine turtles characterised by an oceanic-neritic foraging dichotomy. Using satellite tracking data from adult female loggerhead turtles nesting at Cape Verde (n = 12), we test the hypothesis that oceanic-foraging loggerheads associate with mesocale (10s – to 100s of km) thermal fronts. We use high-resolution (1 km) composite front mapping to characterise frontal activity in the Canary Current Large Marine Ecosystem (LME) over 2 temporal scales: (1) seasonal front frequency and (2) 7-day front metrics. Our use-availability analysis indicates that oceanic loggerheads show a preference for the highly productive upwelling region between Cape Verde and mainland Africa, an area of intense frontal activity. Within the upwelling region, turtles appear to forage epipelagically around mesoscale thermal fronts, exploiting profitable foraging opportunities resulting from physical aggregation of prey.

Basking sharks and oceanographic fronts: quantifying associations in the north‐east Atlantic

Understanding the mechanisms linking oceanographic processes and marine vertebrate habitat use is critical to effective management of populations of conservation concern. The basking shark Cetorhinus maximus has been shown to associate with oceanographic fronts – physical interfaces at the transitions between water masses – to exploit foraging opportunities resulting from aggregation of zooplankton. However, the scale, significance and variability of these observed associations have not yet been established. Here, we quantify the influence of mesoscale (10s – 100s km) frontal activity on habitat use over timescales of weeks to months. We use animal-mounted archival tracking with composite front mapping via Earth Observation (EO) remote sensing to provide an oceanographic context to individual shark movements. We investigate levels of association with fronts occurring over two spatio-temporal scales, (i) broad-scale seasonally persistent frontal zones and (ii) contemporaneous mesoscale thermal and chl-a fronts. Using random walk simulations and logistic regression within an iterative generalised linear mixed modelling (GLMM) framework, we find that seasonal front frequency is a significant predictor of shark presence. Temporally-matched oceanographic metrics also indicate that sharks demonstrate a preference for productive regions, and associate with contemporaneous thermal and chl-a fronts more frequently than could be expected at random. Moreover, we highlight the importance of cross-frontal temperature change and persistence, which appear to interact to affect the degree of prey aggregation along thermal fronts. These insights have clear implications for understanding the preferred habitats of basking sharks in the context of anthropogenic threat management and marine spatial planning in the northeast Atlantic.

Do oceanic loggerhead turtles Caretta caretta associate with oceanographic fronts? Evidence from the Canary Current Large Marine Ecosystem

ABSTRACT: Oceanographic fronts are physical interfaces between water masses that differ in properties such as temperature, salinity, turbidity and chl a enrichment. Bio-physical coupling along fronts can lead to the development of pelagic biodiversity hotspots. A diverse range of marine vertebrates have been shown to associate with fronts, using them as foraging and migration habitats. Elucidation of the ecological significance of fronts generates a better understanding of marine ecosystem functioning, conferring opportunities to improve management of anthropogenic activities in the oceans. This study presents novel insight into the oceanographic drivers of habitat use in a population of marine turtles characterised by an oceanic-neritic foraging dichotomy. Using satellite tracking data from adult female loggerhead turtles nesting at Cape Verde (n = 12), we test the hypothesis that oceanic-foraging loggerheads associate with mesocale (10s – to 100s of km) thermal fronts. We use high-resolution (1 km) composite front mapping to characterise frontal activity in the Canary Current Large Marine Ecosystem (LME) over 2 temporal scales: (1) seasonal front frequency and (2) 7-day front metrics. Our use-availability analysis indicates that oceanic loggerheads show a preference for the highly productive upwelling region between Cape Verde and mainland Africa, an area of intense frontal activity. Within the upwelling region, turtles appear to forage epipelagically around mesoscale thermal fronts, exploiting profitable foraging opportunities resulting from physical aggregation of prey.

Long-term individual foraging site fidelity – why some gannets don’t change their spots

To follow

Basking sharks and oceanographic fronts: quantifying associations in the north-east Atlantic

Understanding the mechanisms linking oceanographic processes and marine vertebrate habitat use is critical to effective management of populations of conservation concern. The basking shark Cetorhinus maximus has been shown to associate with oceanographic fronts – physical interfaces at the transitions between water masses – to exploit foraging opportunities resulting from aggregation of zooplankton. However, the scale, significance and variability of these observed associations have not yet been established. Here, we quantify the influence of mesoscale (10s – 100s km) frontal activity on habitat use over timescales of weeks to months. We use animal-mounted archival tracking with composite front mapping via Earth Observation (EO) remote sensing to provide an oceanographic context to individual shark movements. We investigate levels of association with fronts occurring over two spatio-temporal scales, (i) broad-scale seasonally persistent frontal zones and (ii) contemporaneous mesoscale thermal and chl-a fronts. Using random walk simulations and logistic regression within an iterative generalised linear mixed modelling (GLMM) framework, we find that seasonal front frequency is a significant predictor of shark presence. Temporally-matched oceanographic metrics also indicate that sharks demonstrate a preference for productive regions, and associate with contemporaneous thermal and chl-a fronts more frequently than could be expected at random. Moreover, we highlight the importance of cross-frontal temperature change and persistence, which appear to interact to affect the degree of prey aggregation along thermal fronts. These insights have clear implications for understanding the preferred habitats of basking sharks in the context of anthropogenic threat management and marine spatial planning in the northeast Atlantic.

Hierarchical random walks in trace fossils and the origin of optimal search behavior

Efficient searching is crucial for timely location of food and other resources. Recent studies show diverse living animals employ a theoretically optimal scale-free random search for sparse resources known as a Lévy walk, but little is known of the origins and evolution of foraging behaviour and the search strategies of extinct organisms. Here we show using simulations of self-avoiding trace fossil trails that randomly introduced strophotaxis (U-turns) – initiated by obstructions such as ¬¬¬self-trail avoidance or innate cueing – leads to random looping patterns with clustering across increasing scales that is consistent with the presence of Lévy walks. This predicts optimal Lévy searches can emerge from simple behaviours observed in fossil trails. We then analysed fossilized trails of benthic marine organisms using a novel path analysis technique and find the first evidence of Lévy-like search strategies in extinct animals. Our results show that simple search behaviours of extinct animals in heterogeneous environments give rise to hierarchically nested Brownian walk clusters that converge to optimal Lévy patterns. Primary productivity collapse and large-scale food scarcity characterising mass extinctions evident in the fossil record may have triggered adaptation of optimal Lévy-like searches. The findings suggest Lévy-like behaviour has been employed by foragers since at least the Eocene but may have a more ancient origin, which could explain recent widespread observations of such patterns among modern taxa.

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish. Behavioral Ecology and Sociobiology, 68, 1995-2003.

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.

Looking backwards to look forwards: the role of natural history in temperate reef ecology

Temperate reefs are superb tractable systems for testing hypotheses in ecology and evolutionary biology. Accordingly there is a rich history of research stretching back over 100 years, which has made major contributions to general ecological and evolutionary theory as well as providing better understanding of how littoral systems work by linking pattern with process. A brief resumé of the history of temperate reef ecology is provided to celebrate this rich heritage. As a community, temperate reef ecologists generally do well designed experiments and test well formulated hypotheses. Increasingly large datasets are being collected, collated and subjected to complex meta-analyses and used for modelling. These datasets do not happen spontaneously – the burgeoning subject of macroecology is possible only because of the efforts of dedicated natural historians whether it be observing birds, butterflies, or barnacles. High-quality natural history and old-fashioned field craft enable surveys or experiments to be stratified (i.e. replicates are replicates and not a random bit of rock) and lead to the generation of more insightful hypotheses. Modern molecular approaches have led to the discovery of cryptic species and provided phylogeographical insights, but natural history is still required to identify species in the field. We advocate a blend of modern approaches with old school skills and a fondness for temperate reefs in all their splendour.