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Sympatric Atlantic Puffins and Razorbills Show Contrasting Responses to Adverse Marine Conditions During Winter Foraging Within the North Sea Katie St

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Sympatric Atlantic Puffins and Razorbills Show Contrasting Responses to Adverse Marine Conditions During Winter Foraging Within the North Sea Katie St St. John Glew et al. Movement Ecology (2019) 7:33 https://doi.org/10.1186/s40462-019-0174-4 RESEARCH Open Access Sympatric Atlantic puffins and razorbills show contrasting responses to adverse marine conditions during winter foraging within the North Sea Katie St. John Glew1* , Sarah Wanless2, Michael P. Harris2, Francis Daunt2, Kjell Einar Erikstad3,4, Hallvard Strøm5, John R. Speakman6,7, Benjamin Kürten8,9 and Clive N. Trueman1 Abstract Background: Natural environments are dynamic systems with conditions varying across years. Higher trophic level consumers may respond to changes in the distribution and quality of available prey by moving to locate new resources or by switching diets. In order to persist, sympatric species with similar ecological niches may show contrasting foraging responses to changes in environmental conditions. However, in marine environments this assertion remains largely untested for highly mobile predators outside the breeding season because of the challenges of quantifying foraging location and trophic position under contrasting conditions. Method: Differences in overwinter survival rates of two populations of North Sea seabirds (Atlantic puffins (Fratercula arctica) and razorbills (Alca torda)) indicated that environmental conditions differed between 2007/08 (low survival and thus poor conditions) and 2014/15 (higher survival, favourable conditions). We used a combination of bird-borne data loggers and stable isotope analyses to test 1) whether these sympatric species showed consistent responses with respect to foraging location and trophic position to these contrasting winter conditions during periods when body and cheek feathers were being grown (moult) and 2) whether any observed changes in moult locations and diet could be related to the abundance and distribution of potential prey species of differing energetic quality. Results: Puffins and razorbills showed divergent foraging responses to contrasting winter conditions. Puffins foraging in the North Sea used broadly similar foraging locations during moult in both winters. However, puffin diet significantly differed, with a lower average trophic position in the winter characterised by lower survival rates. By contrast, razorbills’ trophic position increased in the poor survival winter and the population foraged in more distant southerly waters of the North Sea. Conclusions: Populations of North Sea puffins and razorbills showed contrasting foraging responses when environmental conditions, as indicated by overwinter survival differed. Conservation of mobile predators, many of which are in sharp decline, may benefit from dynamic spatial based management approaches focusing on behavioural changes in response to changing environmental conditions, particularly during life history stages associated with increased mortality. Keywords: Fratercula arctica, Isoscape, Alca torda, Marine spatial management, North Sea, Seabird foraging behaviour, Spatial ecology, Trophic ecology, Moult * Correspondence: [email protected] 1Ocean and Earth Science, University of Southampton, Waterfront Campus, Southampton SO143ZH, UK Full list of author information is available at the end of the article © The Author(s). 2019 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. St. John Glew et al. Movement Ecology (2019) 7:33 Page 2 of 14 Background Models of spatial variations in the stable isotopic com- How animals respond to changing environmental condi- positions of the pelagic food web (isoscapes) have re- tions is a key topic in movement and foraging ecology. cently been constructed for the North Sea [63]. St. John Mobile predators may potentially respond to changes in Glew et al. [58] used inferences from isoscape models, prey availability by moving to find new resources or feather stable isotope data and tag data to estimate at- switching diets [1, 11, 42, 43, 45, 50, 70]. The nature of sea locations and diet in three species of auk (common any such response reflects a balance of risk/reward asso- guillemot (Uria aalge), razorbill (Alca torda) and Atlan- ciated with resource intake, energetic costs and physio- tic puffin (Fratercula arctica), hereafter puffin) during logical demands. The ability of sympatric species with the winter moult period of 2014/15, a period charac- similar ecological niches to respond to environmental terised by overwinter survival rates similar to or higher variation, and the relative nature of their responses, than the 33 year long term average of these auk popula- could profoundly influence patterns of species distribu- tions, and presumably favourable environmental condi- tion, particularly when environmental disturbances tions. Here we compare locational and isotopic data occur during critical life history stages. However, the from puffins and razorbills during winter foraging of challenge of quantifying foraging distribution and diet of 2014/15 with similar data from the winter of 2007/08, sympatric species of mobile predators means that this when overwinter survival was markedly lower than the assertion remains largely untested. long-term average implying that conditions were less Members of the auk family (Alcidae) dominate the favourable. avian community wintering in the North Sea [59]. They Our aims were to 1) test whether differences in over- forage by pursuit diving and the medium and larger winter survival were associated with changes in locations sized species undergo moult of their flight and body within the North Sea and/or trophic position, 2) assess feathers outside the breeding season (hereafter referred whether such responses were similar in the two species to as during winter). During this period of annual fea- and 3) investigate whether any observed changes in loca- ther moult energy demands are high, making them po- tions and diet could be related to the abundance and tentially vulnerable to reduced prey availability and distribution of potential prey species. For this final test severe weather conditions [28, 55]. Like many other sea- we collated data on the distribution and abundance of birds, annual survival rates of adult auks are typically lesser sandeel (Ammodytes marinus), sprat (Sprattus high (c.0.90) with most mortality occurring in winter sprattus) and herring (Clupea harengus), lipid-rich high [28]. Ecological and environmental conditions during quality prey that are known to be common in the diet of winter moult therefore have the potential to influence puffins and razorbills during the breeding and winter the distribution, behaviour and mortality of seabirds, but seasons across the North Sea [26, 27, 37]. We also ex- the majority of studies on environmental influences on tracted distribution and abundance data, as well as diet and distributions of seabirds have focussed on im- isotope measurements for snake pipefish (Entelurus pacts during the breeding season [30, 54, 70]. Consider- aequoreus), as this normally rare and nutritionally poor ably less information is available regarding changes species showed a short-lived population explosion in occurring outside the breeding season. 2007 and 2008, and was found in the diets of a wide Long-term demographic studies of three auk species range of marine predators [24, 64]. breeding at the Isle of May colony in the northwestern North Sea found that annual survival rates varied synchron- Methods ously in association with common environmental proxies Survival rates [35]. However, the extent of population- and individual-level Fieldwork was carried out on the Isle of May National differences in foraging behaviours in response to different Nature Reserve, south-east Scotland (56°11′N, 2°34′W) winter conditions and prey availability remain relatively un- where annual survival rates of adult puffins and razor- explored. Until recently observation of behaviour during bills have been estimated each year since 1984. Breeding winter foraging has been impossible, but the development of adults were caught and marked with a numbered metal bird-borne, light-based geolocators [7, 10, 18, 71]toinfer ring and a unique combination of colour rings (total foraging location, and the use of stable isotope analysis to sample sizes over the study: n = 694 puffins and 215 ra- retrospectively determine nutrient acquisition and trophic zorbills). In each year, visual searches were made for position has provided new opportunities [29, 53]. Combin- marked birds in the areas where they had been ringed ing stable isotope and geolocation methods reveals spatial and in other parts of the colony. The resighting data and trophic information that cannot be retrieved from either were modelled with a standard Cormack-Jolly-Seber approach in isolation, and crucially now enables us to test mark-recapture model, which estimates annual survival whether species respond to contrasting conditions during and recapture probabilities as outlined in Freeman et al. winter foraging by altering location and/or switching diet. [13], and updated to include subsequent years’ data.
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