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      Modelling the Effects of Prey Size and Distribution on Prey Capture Rates of Two Sympatric Marine Predators

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          Abstract

          Understanding how prey capture rates are influenced by feeding ecology and environmental conditions is fundamental to assessing anthropogenic impacts on marine higher predators. We compared how prey capture rates varied in relation to prey size, prey patch distribution and prey density for two species of alcid, common guillemot ( Uria aalge) and razorbill ( Alca torda) during the chick-rearing period. We developed a Monte Carlo approach parameterised with foraging behaviour from bird-borne data loggers, observations of prey fed to chicks, and adult diet from water-offloading, to construct a bio-energetics model. Our primary goal was to estimate prey capture rates, and a secondary aim was to test responses to a set of biologically plausible environmental scenarios. Estimated prey capture rates were 1.5±0.8 items per dive (0.8±0.4 and 1.1±0.6 items per minute foraging and underwater, respectively) for guillemots and 3.7±2.4 items per dive (4.9±3.1 and 7.3±4.0 items per minute foraging and underwater, respectively) for razorbills. Based on species' ecology, diet and flight costs, we predicted that razorbills would be more sensitive to decreases in 0-group sandeel (Ammodytes marinus) length (prediction 1), but guillemots would be more sensitive to prey patches that were more widely spaced (prediction 2), and lower in prey density (prediction 3). Estimated prey capture rates increased non-linearly as 0-group sandeel length declined, with the slope being steeper in razorbills, supporting prediction 1. When prey patches were more dispersed, estimated daily energy expenditure increased by a factor of 3.0 for guillemots and 2.3 for razorbills, suggesting guillemots were more sensitive to patchier prey, supporting prediction 2. However, both species responded similarly to reduced prey density (guillemot expenditure increased by 1.7; razorbill by 1.6), thus not supporting prediction 3. This bio-energetics approach complements other foraging models in predicting likely impacts of environmental change on marine higher predators dependent on species-specific foraging ecologies.

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          Prey Patch Patterns Predict Habitat Use by Top Marine Predators with Diverse Foraging Strategies

          Spatial coherence between predators and prey has rarely been observed in pelagic marine ecosystems. We used measures of the environment, prey abundance, prey quality, and prey distribution to explain the observed distributions of three co-occurring predator species breeding on islands in the southeastern Bering Sea: black-legged kittiwakes (Rissa tridactyla), thick-billed murres (Uria lomvia), and northern fur seals (Callorhinus ursinus). Predictions of statistical models were tested using movement patterns obtained from satellite-tracked individual animals. With the most commonly used measures to quantify prey distributions - areal biomass, density, and numerical abundance - we were unable to find a spatial relationship between predators and their prey. We instead found that habitat use by all three predators was predicted most strongly by prey patch characteristics such as depth and local density within spatial aggregations. Additional prey patch characteristics and physical habitat also contributed significantly to characterizing predator patterns. Our results indicate that the small-scale prey patch characteristics are critical to how predators perceive the quality of their food supply and the mechanisms they use to exploit it, regardless of time of day, sampling year, or source colony. The three focal predator species had different constraints and employed different foraging strategies – a shallow diver that makes trips of moderate distance (kittiwakes), a deep diver that makes trip of short distances (murres), and a deep diver that makes extensive trips (fur seals). However, all three were similarly linked by patchiness of prey rather than by the distribution of overall biomass. This supports the hypothesis that patchiness may be critical for understanding predator-prey relationships in pelagic marine systems more generally.
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            Differences in prey selection and behaviour during self-feeding and chick provisioning in rhinoceros auklets.

            We determined whether a marine diving bird, the rhinoceros auklet, Cerorhinca monocerata, used different foraging behaviour and collected different prey items for its young than when feeding itself. Foraging behaviour was determined by conducting visual scans, and prey items were sampled by collecting fish delivered to chicks and by collecting fish where auklets were self-feeding, which was verified by two other sources of information. Adult auklets ate small fish (59.1+/-0.5 mm, N=547), including juvenile Pacific sand lance, Ammodytes hexapterus, and Pacific herring, Clupea harengus, but collected larger fish to feed their chicks (95.2+/-1.3 mm, N=321), including primarily Pacific sand lance, Pacific herring, Pacific salmon species, Oncorhynchus spp., and surf smelt, Hypomesus pretiosus. In addition, auklets collected fish for themselves primarily by diving in mixed-species feeding flocks before 1600 hours, whereas they collected fish to feed their chicks by diving solitarily after 1600 hours. This suggests that auklets switched both foraging behaviour and prey selection when collecting fish for self-feeding, compared with when collecting fish for chick provisioning. Several avian studies have documented different diets of adults and chicks, but this is the first research to observe directly and document different foraging behaviour used in adult and chick provisioning. We emphasize the importance of distinguishing between self-feeding and chick provisioning in foraging and life history studies. Copyright 1999 The Association for the Study of Animal Behaviour.
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              Unconventional ventral attachment of time-depth recorders as a new method for investigating time budget and diving behaviour of seabirds.

              We tested the use of commercially available electronic time-depth recorders (TDRs) to quantify activities and thus total time budgets of seabirds. This new method involved first fitting TDRs onto the birds' bellies (not on their backs), and, secondly, analysing continuous recordings of temperature, light and pressure to differentiate activities on land and at sea. The birds studied were 12 common guillemots (Uria aalge) rearing chicks at Hornøya, in northern Norway. The method successfully recorded five different activities: at the colony, flying, diving, and resting or active at the sea surface. Overall, common guillemots spent 68% of their time at the colony and 32% at sea. While at sea, the birds spent the majority (77%) of their time at the surface, during which they were active 64% of the time, and rested only 13%. Birds engaged in the costly behaviours of flying and diving for shorter times (11% and 12% of their time at sea, respectively). The method allowed us to differentiate between two types of trips to sea based on the presence (foraging trips: 77% of the total number of trips) or absence (non-foraging trips: 23%) of dives. On average, foraging trips lasted 3.2 h, but most trips were shorter (<1 h), during which the mean estimated travel distance from the colony was 11 km. Diving occurred in bouts of 7.7+/-6.6 dives (mean +/- S.D.). The mean maximum dive depth was 10.2+/-7.6 m (deepest dive: 37 m), and the mean dive duration and post-dive intervals were 38.7+/-21.3 s (longest dive: 119 s) and 20+/-12 s, respectively. Direct and indirect evidence suggests that common guillemots had no difficulty in finding food during the study period, and that the TDRs had minimal effects on the birds' behaviour and physiology. The method is easy to use in the field and is applicable to many other flying seabird species; it is therefore an efficient way of collecting information on time budgets and diving behaviour in the context of various ecological and monitoring studies.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2013
                15 November 2013
                : 8
                : 11
                : e79915
                Affiliations
                [1 ]School of Biology, University of Leeds, Leeds, United Kingdom
                [2 ]Centre for Ecology & Hydrology, Bush Estate, Penuick, Midlothian, United Kingdom
                [3 ]Centre d'Ecologie Fonctionnelle et Evolutive, UMR 5175 du CNRS, Montpellier, France
                [4 ]Percy FitzPatrick Institute, DST/NRF Centre of Excellence, University of Cape Town, Rondebosch, South Africa
                [5 ]Department of Biology, University of Pisa, Pisa, Italy
                [6 ]Graduate School of Fisheries Sciences, Hokkaido University, Hakodate, Hokkaido, Japan
                University of Waikato (National Institute of Water and Atmospheric Research), New Zealand
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: CBT FD KCH SW. Performed the experiments: CBT FD MPH SB YW KCH SW. Analyzed the data: CBT. Contributed reagents/materials/analysis tools: CBT FD DG MPH SB YW. Wrote the paper: CBT FD KCH SW.

                [¤]

                Current address: British Trust for Ornithology, The Nunnery, Thetford, Norfolk, United Kingdom

                Article
                PONE-D-13-26534
                10.1371/journal.pone.0079915
                3829866
                1d8116cd-675e-4786-99bf-d314547df5fc
                Copyright @ 2013

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 27 June 2013
                : 27 September 2013
                Page count
                Pages: 10
                Funding
                This work was supported by a Case Studentship (NER/S/A/2004/12298) from the Natural Environmental Research Council (NERC), an EU-funded project (FP5 Project ‘Interactions between the marine environment, predators and prey: implications for sustainable sandeel fisheries’ (IMPRESS, Q5RS-2000-30864), and the Joint Nature Conservation Committee (on behalf of Natural England, Scottish Natural Heritage, the Countryside Council for Wales and the Council for Nature Conservation and the Countryside in Northern Ireland). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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