Foraging success of biological Levy flights recorded in situ

It is an open question how animals find food in dynamic natural environments where they possess little or no knowledge of where resources are located. Foraging theory predicts that in environments with sparsely distributed target resources, where forager knowledge about resources’ locations is incomplete, Lévy flight movements optimize the success of random searches. However, the putative success of Lévy foraging has been demonstrated only in model simulations. Here, we use high-temporal-resolution Global Positioning System (GPS) tracking of wandering (Diomedea exulans) and black-browed albatrosses (Thalassarche melanophrys) with simultaneous recording of prey captures, to show that both species exhibit Lévy and Brownian movement patterns. We find that total prey masses captured by wandering albatrosses during Lévy movements exceed daily energy requirements by nearly fourfold, and approached yields by Brownian movements in other habitats. These results, together with our reanalysis of previously published albatross data, overturn the notion that albatrosses do not exhibit Lévy patterns during foraging, and demonstrate that Lévy flights of predators in dynamic natural environments present a beneficial alternative strategy to simple, spatially intensive behaviors. Our findings add support to the possibility that biological Lévy flight may have naturally evolved as a search strategy in response to sparse resources and scant information.

Optimal foraging strategies: Lévy walks balance searching and patch exploitation under a very broad range of conditions

While evidence for optimal random search patterns, known as Lévy walks, in empirical movement data is mounting for a growing list of taxa spanning motile cells to humans, there is still much debate concerning the theoretical generality of Lévy walk optimisation. Here, using a new and robust simulation environment, we investigate in the most detailed study to date (24×10(6) simulations) the foraging and search efficiencies of 2-D Lévy walks with a range of exponents, target resource distributions and several competing models. We find strong and comprehensive support for the predictions of the Lévy flight foraging hypothesis and in particular for the optimality of inverse square distributions of move step-lengths across a much broader range of resource densities and distributions than previously realised. Further support for the evolutionary advantage of Lévy walk movement patterns is provided by an investigation into the 'feast and famine' effect, with Lévy foragers in heterogeneous environments experiencing fewer long 'famines' than other types of searchers. Therefore overall, optimal Lévy foraging results in more predictable resources in unpredictable environments.

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.

The influence of ocean acidification on nitrogen regeneration and nitrous oxide production in the northwest European shelf sea

The assimilation and regeneration of dissolved inorganic nitrogen, and the concentration of N₂O, was investigated at stations located in the NW European shelf sea during June/July 2011. These observational measurements within the photic zone demonstrated the simultaneous regeneration and assimilation of NH₄⁺, NO₂⁻ and NO₃⁻. NH₄⁺ was assimilated at 1.82–49.12 nmol N L⁻¹ h⁻¹ and regenerated at 3.46–14.60 nmol N L⁻¹ h⁻¹; NO₂^- was assimilated at 0–2.08 nmol N L⁻¹ h⁻¹ and regenerated at 0.01–1.85 nmol N L⁻¹ h⁻¹; NO₃⁻ was assimilated at 0.67–18.75 nmol N L⁻¹ h⁻¹ and regenerated at 0.05–28.97 nmol N L⁻¹ h⁻¹. Observations implied that these processes were closely coupled at the regional scale and that nitrogen recycling played an important role in sustaining phytoplankton growth during the summer. The [N₂O], measured in water column profiles, was 10.13 ± 1.11 nmol L⁻¹ and did not strongly diverge from atmospheric equilibrium indicating that sampled marine regions were neither a strong source nor sink of N₂O to the atmosphere. Multivariate analysis of data describing water column biogeochemistry and its links to N-cycling activity failed to explain the observed variance in rates of N-regeneration and N-assimilation, possibly due to the limited number of process rate observations. In the surface waters of five further stations, ocean acidification (OA) bioassay experiments were conducted to investigate the response of NH₄⁺ oxidising and regenerating organisms to simulated OA conditions, including the implications for [N₂O]. Multivariate analysis was undertaken which considered the complete bioassay data set of measured variables describing changes in N-regeneration rate, [N₂O] and the biogeochemical composition of seawater. While anticipating biogeochemical differences between locations, we aimed to test the hypothesis that the underlying mechanism through which pelagic N-regeneration responded to simulated OA conditions was independent of location. Our objective was to develop a mechanistic understanding of how NH₄⁺ regeneration, NH₄⁺ oxidation and N₂O production responded to OA. Results indicated that N-regeneration process responses to OA treatments were location specific; no mechanistic understanding of how N-regeneration processes respond to OA in the surface ocean of the NW European shelf sea could be developed.

Monitoring of a quasi-stationary eddy in the Bay of Biscay by means of satellite, in situ and model results

The presence of a quasi-stationary anticyclonic eddy within the southeastern Bay of Biscay (centred around 44°30′N-4°W) has been reported on various occasions in the bibliography. The analysis made in this study for the period 2003–2010, by using in situ and remote sensing measurements and model results shows that this mesoscale coherent structure is present almost every year from the end of winter-beginning of spring, to the beginning of fall. During this period it remains in an area limited to the east by the Landes Plateau, to the west by Le Danois Bank and Torrelavega canyon and to the northwest by the Jovellanos seamount. All the observations and analysis made in this contribution, suggest that this structure is generated between Capbreton and Torrelavega canyons. Detailed monitoring from in situ and remote sensing data of an anticyclonic quasi-stationary eddy, in 2008, shows the origin of this structure from a warm water current located around 43°42′N-3°30′W in mid-January. This coherent structure is monitored until August around the same area, where it has a marked influence on the Sea Level Anomaly, Sea Surface Temperature and surface Chlorophyll-a concentration. An eddy tracking method, applied to the outputs of a numerical model, shows that the model is able to reproduce this type of eddy, with similar 2D characteristics and lifetimes to that suggested by the observations and previous works. This is the case, for instance, of the simulated MAY04 eddy, which was generated in May 2004 around Torrelavega canyon and remained quasi-stationary in the area for 4 months. The diameter of this eddy ranged from 40 to 60 km, its azimuthal velocity was less than 20 cm s−1, its vertical extension reached 3000–3500 m depth during April and May and it was observed to interact with other coherent structures.

On the temporal consistency of chlorophyll products derived from three ocean-colour sensors

Satellite ocean-colour sensors have life spans lasting typically five-to-ten years. Detection of long-term trends in chlorophyll-a concentration (Chl-a) using satellite ocean colour thus requires the combination of different ocean-colour missions with sufficient overlap to allow for cross-calibration. A further requirement is that the different sensors perform at a sufficient standard to capture seasonal and inter-annual fluctuations in ocean colour. For over eight years, the SeaWiFS, MODIS-Aqua and MERIS ocean-colour sensors operated in parallel. In this paper, we evaluate the temporal consistency in the monthly Chl-a time-series and in monthly inter-annual variations in Chl-a among these three sensors over the 2002–2010 time period. By subsampling the monthly Chl-a data from the three sensors consistently, we found that the Chl-a time-series and Chl-a anomalies among sensors were significantly correlated for >90% of the global ocean. These correlations were also relatively insensitive to the choice of three Chl-a algorithms and two atmospheric-correction algorithms. Furthermore, on the subsampled time-series, correlations between Chl-a and time, and correlations between Chl-a and physical variables (sea-surface temperature and sea-surface height) were not significantly different for >92% of the global ocean. The correlations in Chl-a and physical variables observed for all three sensors also reflect previous theories on coupling between physical processes and phytoplankton biomass. The results support the combining of Chl-a data from SeaWiFS, MODIS-Aqua and MERIS sensors, for use in long-term Chl-a trend analysis, and highlight the importance of accounting for differences in spatial sampling among sensors when combining ocean-colour observations.

The future of the northeast Atlantic benthic flora in a high CO2 world

Seaweed and seagrass communities in the northeast Atlantic have been profoundly impacted by humans, and the rate of change is accelerating rapidly due to runaway CO2 emissions and mounting pressures on coastlines associated with human population growth and increased consumption of finite resources. Here, we predict how rapid warming and acidification are likely to affect benthic flora and coastal ecosystems of the northeast Atlantic in this century, based on global evidence from the literature as interpreted by the collective knowledge of the authorship. We predict that warming will kill off kelp forests in the south and that ocean acidification will remove maerl habitat in the north. Seagrasses will proliferate, and associated epiphytes switch from calcified algae to diatoms and filamentous species. Invasive species will thrive in niches liberated by loss of native species and spread via exponential development of artificial marine structures. Combined impacts of seawater warming, ocean acidification, and increased storminess may replace structurally diverse seaweed canopies, with associated calcified and noncalcified flora, with simple habitats dominated by noncalcified, turf-forming seaweeds.

Simple shade plots aid better long-term choices of data pre-treatment in multivariate assemblage studies

Shade plots, simple visual representations of abundance matrices from multivariate species assemblage studies, are shown to be an effective aid in choosing an overall transformation (or other pre-treatment) of quantitative data for long-term use, striking an appropriate balance between dominant and less abundant taxa in ensuing resemblance-based multivariate analyses. Though the exposition is entirely general and applicable to all community studies, detailed illustrations of the comparative power and interpretative possibilities of shade plots are given in the case of two estuarine assemblage studies in south-western Australia: (a) macrobenthos in the upper Swan Estuary over a two-year period covering a highly significant precipitation event for the Perth area; and (b) a wide-scale spatial study of the nearshore fish fauna from five divergent estuaries. The utility of transformations of intermediate severity is again demonstrated and, with greater novelty, the potential importance seen of further mild transformation of all data after differential down-weighting (dispersion weighting) of spatially clumped' or schooled' species. Among the new techniques utilized is a two-way form of the RELATE test, which demonstrates linking of assemblage structure (fish) to continuous environmental variables (water quality), having removed a categorical factor (estuary differences). Re-orderings of sample and species axes in the associated shade plots are seen to provide transparent explanations at the species level for such continuous multivariate patterns.

Multiple environmental controls on phytoplankton growth strategies determine adaptive responses of the N : P ratio

The controls on the 'Redfield' N:P stoichiometry of marine phytoplankton and hence the N:P ratio of the deep ocean remain incompletely understood. Here, we use a model for phytoplankton ecophysiology and growth, based on functional traits and resource-allocation trade-offs, to show how environmental filtering, biotic interactions, and element cycling in a global ecosystem model determine phytoplankton biogeography, growth strategies and macromolecular composition. Emergent growth strategies capture major observed patterns in marine biomes. Using a new synthesis of experimental RNA and protein measurements to constrain per-ribosome translation rates, we determine a spatially variable lower limit on adaptive rRNA:protein allocation and hence on the relationship between the largest cellular P and N pools. Comparison with the lowest observed phytoplankton N:P ratios and N:P export fluxes in the Southern Ocean suggests that additional contributions from phospholipid and phosphorus storage compounds play a fundamental role in determining the marine biogeochemical cycling of these elements.

Lipid productivity and cell wall ultrastructure of six strains of Nannochloropsis: Implications for biofuel production and downstream processing

Microalgae are generating considerable interest for third generation biodiesel production. However, appropriate strain selection is proving challenging due to the significant variation in cellular physiology, metabolic potential and genetics observed even amongst strains deemed morphologically similar. Six strains of Nannochloropsis from the CCAP culture collection were assessed for their lipid productivity and cellular structure, as proxies for oil production and harvesting ease, to assess their suitability as biodiesel production platforms. Differences in growth rate and lipid accumulation across the strains were observed. Nannochloropsis oculata strain 849/7 showed significantly reduced doubling time compared to Nannochloropsis salina strain 849/3, whilst Nannochloropsis oceanica 849/10 produced the highest lipid content. In addition the six strains could be differentiated into 3 distinct classes based on their cell wall thickness, which varied across the strains from 63 to 119 nm and which is independent of both species and geographical isolation location. The importance of these variations in ultrastructure and physiology for biodiesel production is discussed.

Coastal Zone Ecosystem Services: From science to values and decision making; a case study

This research is concerned with the following environmental research questions: socio-ecological system complexity, especially when valuing ecosystem services; ecosystems stock and services flow sustainability and valuation; the incorporation of scale issues when valuing ecosystem services; and the integration of knowledge from diverse disciplines for governance and decision making. In this case study, we focused on ecosystem services that can be jointly supplied but independently valued in economic terms: healthy climate (via carbon sequestration and storage), food (via fisheries production in nursery grounds), and nature recreation (nature watching and enjoyment). We also explored the issue of ecosystem stock and services flow, and we provide recommendations on how to value stock and flows of ecosystem services via accounting and economic values respectively. We considered broadly comparable estuarine systems located on the English North Sea coast: the Blackwater estuary and the Humber estuary. In the past, these two estuaries have undergone major land-claim. Managed realignment is a policy through which previously claimed intertidal habitats are recreated allowing the enhancement of the ecosystem services provided by saltmarshes. In this context, we investigated ecosystem service values, through biophysical estimates and welfare value estimates. Using an optimistic (extended conservation of coastal ecosystems) and a pessimistic (loss of coastal ecosystems because of, for example, European policy reversal) scenario, we find that context dependency, and hence value transfer possibilities, vary among ecosystem services and benefits. As a result, careful consideration in the use and application of value transfer, both in biophysical estimates and welfare value estimates, is advocated to supply reliable information for policy making.

Enhancing the Global Ocean Observing System to meet evidence based needs for the ecosystem-based management of coastal ecosystem services

Ecosystem-based approaches (EBAs) to managing anthropogenic pressures on ecosystems, adapting to changes in ecosystem states (indicators of ecosystem health), and mitigating the impacts of state changes on ecosystem services are needed for sustainable development. EBAs are informed by integrated ecosystem assessments (IEAs) that must be compiled and updated frequently for EBAs to be effective. Frequently updated IEAs depend on the sustained provision of data and information on pressures, state changes, and impacts of state changes on services. Nowhere is this truer than in the coastal zone, where people and ecosystem services are concentrated and where anthropogenic pressures converge. This study identifies the essential indicator variables required for the sustained provision of frequently updated IEAs, and offers an approach to establishing a global network of coastal observations within the framework of the Global Ocean Observing System. The need for and challenges of capacity-building are highlighted, and examples are given of current programmes that could contribute to the implementation of a coastal ocean observing system of systems on a global scale. This illustrates the need for new approaches to ocean governance that can achieve coordinated integration of existing programmes and technologies as a first step towards this goal.

Permanent draft genomes of four new coccolithoviruses: EhV-18, EhV-145, EhV-156 and EhV-164

Coccolithoviruses infect the marine coccolithophorid microalga Emiliania huxleyi. Here, we describe the genomes of four new coccolithoviruses isolated from UK coastal locations. Of particular interest, EhV-18 and EhV-145 encode serine palmitoyltransferase function via two distinct genes, whereas all other coccolithoviruses have SPT as a gene fusion of LCB1/LCB2 domains.

A massive phytoplankton bloom induced by an ecosystem-scale iron fertilization experiment in the equatorial Pacific Ocean

The seeding of an expanse of surface waters in the equatorial Pacific Ocean with low concentrations of dissolved iron triggered a massive phytoplankton bloom which consumed large quantities of carbon dioxide and nitrate that these microscopic plants cannot fully utilize under natural conditions. These and other observations provide unequivocal support for the hypothesis that phytoplankton growth in this oceanic region is limited by iron bioavailability.

Air-sea gas exchange

This chapter contains sections titled: Introduction Air-Sea Gas Exchange Models and Theory Laboratory Studies of Air-Water Gas Exchange Large-Scale Estimates of Air-Sea Gas Transfer Local Techniques and Measurements Micrometeorological Techniques and Measurements Parameterizations of Air-Sea Gas Transfer Future Work