Shifting Nonbreeding Distributions of Migratory Fauna in Relation to Climatic Change
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Global Change Biology (2005) 11, 31–38, doi: 10.1111/j.1365-2486.2004.00876.x Shifting nonbreeding distributions of migratory fauna in relation to climatic change GRAHAM E. AUSTIN andMARK M. REHFISCH British Trust for Ornithology, The Nunnery, Thetford, Norfolk IP24 2PU, UK Abstract The distributions of eight out of nine common species of waders (Charadrii) overwintering on UK estuaries have changed in association with recent climate change. These birds represent a high proportion of various populations from breeding grounds as far apart as Greenland to the west to high-arctic Russia to the east. During warmer winters, smaller proportions of seven species wintered in south-west Britain. The distributions of the smaller species show the greatest temperature dependence. The opposite was found for the largest species and no relationship was found for a particularly site-faithful species. In north-west Europe, the winter isotherms have a broadly north to south alignment, with the east being colder than the west. The average minimum winter temperatures across the UK having increased by about 1.5 1C since the mid-1980s, the temperatures on the east coast during recent winters have been similar to those of the west coast during the mid-1980s. On average, estuaries on the east and south coasts of Britain have muddier sediments than those on the west coast and thus support a higher biomass of the invertebrate prey of waders. We suggest that, with global climatic change, the advantage gained by waders wintering in the milder west to avoid cold weather-induced mortality is diminished. Consequently, more choose to winter in the east and thus benefit from better foraging opportunities. The implications of these results are considered in terms of a site-based approach to wildlife protection used in Europe and elsewhere. Keywords: body weight, conservation, global change, re-distribution, shorebirds, site designation, waders, winter iskotherms Received 27 February 2004; received in revised form and accepted 8 July 2004 The internationally important numbers of waders Introduction and wildfowl wintering in the United Kingdom Meta-analyses have shown a clear effect of climatic (Rehfisch et al., 2003a, b) are attracted there by change on nature (Parmesan & Yohe, 2003; Root et al., productive wetlands and relatively mild winters. These 2003). With climatic change, phenological changes in birds arrive during the late Northern Hemisphere plants (Myneni et al., 1997), birds (Crick & Sparks, 1999; summer from breeding grounds as far apart as Green- Thomas et al., 2001) and amphibians (Beebee, 1995), land (e.g. Red Knot Calidris canutus and Ringed Plover range shifts of plants (Easterling et al., 2000), butterflies Charadrius hiaticula) and Iceland (e.g. Eurasian Oyster- (Parmesan et al., 1999), fish (Finney et al., 2000) and birds catcher Haematopus ostralegus, Ringed Plover, Sander- (Thomas & Lennon, 1999) and changes in the demo- ling Calidris alba and Common Redshank Tringa totanus) graphy of mammals (Coulson et al., 2001) and birds to the west and high-arctic Russia to the east (e.g. Grey (Sillett et al., 2000) have been documented. This paper Plover Pluvialis squatarola) and from wide-ranging presents the first analysis to date to demonstrate changes locations within northern Europe, particularly Fennos- in the winter distribution of migratory fauna with milder candia (e.g. Eurasian Oystercatcher, Ringed Plover, winters, probably to take advantage of better feeding Dunlin Calidris alpina, Eurasian Curlew Numenius conditions and to be nearer breeding grounds. arquata and Common Redshank), but a few species breed within the UK itself (e.g. Eurasian Oystercatcher, Correspondence: Graham E. Austin, Eurasian Curlew and Common Redshank) (Wernham e-mail: [email protected] et al., 2002). Since 1969/1970, most wader populations r 2005 Blackwell Publishing Ltd 31 32 G. E. AUSTIN & M. M. REHFISCH overwintering on UK estuaries have remained fairly relatively high invertebrate densities and in turn, much stable or have increased. However, a general pattern is higher densities of all the species to be considered here evident across many species: numbers on the south- with the exception of Eurasian Oystercatcher (Austin west coast of Britain (Wales and the English counties of et al., 1996; Rehfisch et al., 1997). The majority of Red Avon, Devon, Cornwall and Dorset) have either tended Knot, Sanderling, Grey Plover, Dunlin, Bar-tailed God- to increase less than numbers on the east coast, wit (Limosa lapponica) and Eurasian Oystercatcher and particularly in eastern England (Essex, Suffolk, Norfolk Eurasian Curlew that breed elsewhere but winter in the and Lincolnshire), or have decreased (Austin et al., UK make landfall in the east. Consequently, wintering 2000). These contrasting trends have only been evident on east coast estuaries rather than those further west since the mid-1980s. While the extent and distribution incurs lower migration costs. Also, birds in their first of intertidal habitat has not changed markedly during year of life, which in many of the species considered this period, there have been clear climatic trends, migrate asynchronously to older birds, will have no including those towards milder winters throughout knowledge of estuaries further to the west upon first Britain (Hulme & Jenkins, 1998). In order to predict arrival. UK breeding populations of Eurasian Curlew, future changes and assess the need for habitat Eurasian Oystercatcher, Ringed Plover and Common conservation, it is important to determine whether the Redshank either remain close to their breeding grounds shifts in distributions have resulted from detrimental or shift towards the south-west and west to winter changes in south-west Britain, or from favourable (Wernham et al., 2002). changes in the east of Britain. Hypothesis Choice of wintering area with respect to climate We hypothesize that, while in the past it would have Severe winter weather will have both a direct effect on been relatively more advantageous for individuals of the birds themselves and an effect on their invertebrate certain species to settle further west, relatively lower prey. In most species of wader, individuals remain site prey abundance perhaps coupled with the extra cost faithful with the onset of harsh weather (Myers et al., associated with a longer migration route may have 1979; Davidson & Clark, 1985; Townshend, 1985), which, caused overwintering in south-west Britain to become a coupled with their already relatively high daily energy suboptimal strategy as winters have become more expenditure (Wiersma & Piersma, 1994) and decreased benign. intake rates under such conditions (Goss-Custard et al., Here we consider whether observed trends in wader 1977; Pienkowski, 1981; Zwarts & Wanink, 1993), can distribution with winter weather patterns in the UK are result in substantial mortality when the weather is consistent with this hypothesis. Furthermore, we particularly severe over a prolonged period. In Britain, discuss the implications of our results for conservation such major mortality events occur more often on the management given current scenarios for global climatic relatively colder east coast (Clark, 1982; Davidson & change. Clark, 1985). In a local context, significantly reduced annual survival rates because of severe winter weather Methods have been reported for Common Redshank (Swann & Etheridge, 1989; Insley et al., 1997) and Eurasian Wader counts Oystercatcher (Durell et al., 2001). Despite the tendency towards site faithfulness (Warnock & Takekawa, 1996; Wader numbers have been monitored on all but a few Rehfisch et al., 1996, 2003c), some individuals will move UK estuaries since the winter of 1969/1970 by monthly in response to particularly severe weather (Clark, 1982; co-ordinated counts as part of the Wetland Bird Survey Davidson & Clark, 1985). Species such as Red Knot, (WeBS) (Musgrove et al., 2001). We considered all which are close to their upper limit of metabolic widely distributed, common estuarine species: Ringed expenditure in north-west Europe (Wiersma & Piersma, Plover, Red Knot, Sanderling, Bar-tailed Godwit and 1994), are especially mobile (Davidson & Wilson, 1992). Common Redshank (species that have shown marked declines in numbers in south-west Britain); Grey Plover, Dunlin and Eurasian Curlew (species for which the Choice of wintering area with respect to food availability overall population increase has been more pronounced and migration routes in the east); and Eurasian Oystercatcher (a species for East coast estuaries are generally muddier than those which numbers have remained stable in south-west on the west coast because of differences in estuary Britain) (Austin et al., 2000). We use data from the morphology and tidal influence and thus support winters of 1974/1975 (since when monthly coverage of r 2005 Blackwell Publishing Ltd, Global Change Biology, 11, 31–38, SHIFTING NONBREEDING WADER DISTRIBUTIONS 33 all British estuaries can be considered to be complete) to UK wader populations can vary between years and 1997/1998 (the latest winter for which suitable data differences in distribution could result from the buffer were available) inclusive. effect whereby less attractive sites only support high numbers of waders in years when UK populations are high (Brown, 1969; Fretwell & Lucas, 1970), although Climatic variation given that the increases to the east have been balanced by decreases rather than stability in the west this effect All available data for daily temperature,