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Tern Diet in the UK and Ireland: a Review of Key Prey Species and Potential Impacts of Climate Change

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Tern Diet in the UK and Ireland: a Review of Key Prey Species and Potential Impacts of Climate Change Tern diet in the UK and Ireland: a review of key prey species and potential impacts of climate change October 2017 Elizabeth Green Species and Habitats Officer, RSPB The project "Improving the conservation prospects of the priority species roseate tern throughout its range in the UK and Ireland" is supported by the LIFE Programme of the European Union. LIFE14 NAT/UK/394 ROSEATE TERN Contents Background ............................................................................................................................................. 1 Introduction ............................................................................................................................................ 1 The diet of breeding terns in the UK, Ireland and southern North Sea .................................................. 2 UK, the North Sea ............................................................................................................................... 3 West Scotland ..................................................................................................................................... 7 The Irish Sea ........................................................................................................................................ 7 The Celtic Sea ...................................................................................................................................... 9 The Southern North Sea ...................................................................................................................... 9 The Wadden Sea, south-eastern North Sea ...................................................................................... 10 Summary ........................................................................................................................................... 11 The ecology and status of sandeels, sprat and herring ........................................................................ 12 Sandeels ............................................................................................................................................ 12 Sprat .................................................................................................................................................. 16 Herring .............................................................................................................................................. 21 Mapping prey hotspots and tern resilience .......................................................................................... 24 Prey hotspots .................................................................................................................................... 24 Resilience and vulnerability of tern colonies to prey shortage ........................................................ 29 Impacts of climate-driven environmental change on sandeels, sprat and herring .............................. 31 Observed environmental change ...................................................................................................... 31 Predicted future environmental change ........................................................................................... 33 Summary and recommendations for fisheries management ............................................................... 34 Acknowledgements ............................................................................................................................... 36 References ............................................................................................................................................ 36 Background In order to improve the status of priority species, pressures on populations must be identified and effectively managed. For the roseate tern, a species of high conservation concern in the UK, this requires an improved understanding of the key issues affecting populations in both the breeding and wintering grounds. One such issue is the availability of high-energy food during the breeding season, when terns must collect enough food to sustain themselves and raise their chicks. Due to the small roseate tern population in the UK (~100 pairs) and Ireland (~1,400 pairs), little research has been carried out on the diet of this species. However, the diets of other tern species in this region and the southern North Sea have been more extensively studied and, although inter-specific variation in diet exists, key prey species are fairly consistent across tern species. Improving our understanding of the diets of Arctic, common, little and Sandwich terns will also support the conservation of these species, for which the UK holds internationally important breeding populations. This report, carried out as part of the EU-funded Roseate Tern LIFE Project, reviews current knowledge regarding the diets of the five tern species that breed in the UK and Ireland and the ecology of their key prey species. The impacts that ongoing climate- driven environmental change may have on key fish species are also discussed, and recommendations are made that aim to safeguard these prey populations and the seabirds they support. Introduction Food availability is an important driver of seabird breeding success (Cairns, 1988). High food availability can increase productivity by improving survival or growth rates of chicks or enabling adults to raise a greater number of chicks (Martin, 1987; Monaghan et al., 1989; Robinson and Hamer, 2000). The energy content of food is also a critical factor, and varies between different prey species (Harris and Hislop, 1978) but also within species depending on body condition. Indeed, years of poor body condition of prey have been associated with poor seabird breeding success (Wanless et al., 2005; Österbrom et al., 2006; Österbrom et al., 2008). Such bottom-up effects of forage fish on predators are likely to be strongest when the predator is a specialist that feeds on one or a small number of fish species (Cairns, 1987; Engelhard et al., 2014). One such group of piscivores is the terns. Five tern species breed in the UK: the Arctic tern Sterna paradisaea, common tern Sterna hirundo, little tern Sternula albifrons, roseate tern Sterna dougallii and the Sandwich tern Sterna sandvicensis. Terns target a relatively small number of prey species and generally behave as single prey loaders (although see Dunn, 1972), making them particularly sensitive to changes in food availability and quality (Furness and Tasker, 2000). Tern chicks are generally fed on high energy prey, with low quality species or small individuals being consumed by adults during foraging trips (Dunn, 1972; Ewins, 1985; Wilson et al., 2004; Perrow et al., 2010; Perrow et al., 2011b). However, during periods of low availability of their preferred prey, chicks may be fed on less calorific items. High energy prey tend to be marine fish which have a high protein and fat content, such as sandeels and clupeids (Harris and Hislop, 1978; Massias and Becker, 1990), whereas low energy prey contain more indigestible material and include crustaceans, such as shrimp, and freshwater fish, such as the three-spined stickleback (Massias and Becker, 1990). Experimental studies have shown that chicks fed on a high quality prey (herring) grow quickly while chicks fed on low quality prey (shrimp or sticklebacks) suffer poor growth rates and in some cases even lose weight (Massias and Becker, 1990). This highlights how crucial it is for terns to have access to an abundant supply of high quality prey during the breeding season. 1 Tern diet is likely to be affected by foraging range, as different prey species associate with different areas and habitats (Reay, 1970; Whitehead, 1985; Araujo et al. 2000; Wright et al., 2000). The tern with the shortest foraging range is the smallest-bodied of the five species, the little tern, while the largest- bodied species, the Sandwich tern, has the largest foraging range (Eglington and Perrow, 2014). Thaxter et al. (2012) reviewed studies of foraging distances of seabirds and reported that the maximum foraging range of little terns was 11 km, while that of Arctic, common and roseate was 30 km and Sandwich terns were reported to travel up to 54 km. The mean foraging ranges estimated in the same study were much lower, at 2.1 km for little terns, 7.1 km for Arctic terns, 4.5 km for common terns, 12.2 km for roseate terns and 11.5 km for Sandwich terns. Moderate confidence was associated with the estimates of Arctic, common and Sandwich terns, whereas estimates of little terns and roseate terns were described as low confidence due to a lack of direct measurements (Thaxter et al., 2012). There is substantial variation in estimates of foraging distances between years, colonies, individuals, tidal conditions (Steinen et al., 2000; Perrow et al., 2004; Paiva et al., 2008; Perrow et al., 2010) and even reviews; see Eglington and Perrow (2014) for a comprehensive summary. Therefore it may be inappropriate to consider species as being either strictly inshore or offshore foragers, except perhaps for the little tern (inshore). Diet composition is also influenced by the time of day, the tide and the weather (Veen, 1977; Frick and Becker, 1995; Wendeln, 1997; Stienen et al., 2000; Morris 2012). For example, some diet studies have found that clupeids are most frequently caught in the mornings and evenings, while sandeels are most frequent during the middle of the day (Stienen et al., 2000; Morris 2012). This corresponds with greater availability of clupeids
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