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Importance of Sitka Spruce Picea Sitchensis Seed and Garden Bird-Feeders for Siskins Carduelis Spinus and Coal Tits Periparus Ater

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Importance of Sitka Spruce Picea Sitchensis Seed and Garden Bird-Feeders for Siskins Carduelis Spinus and Coal Tits Periparus Ater Bird Study (2007) 54, 236–247 Importance of Sitka Spruce Picea sitchensis seed and garden bird-feeders for Siskins Carduelis spinus and Coal Tits Periparus ater AILSA J. MCKENZIE1,2*, STEVE J. PETTY3, MIKE P. TOMS4 and ROBERT W. FURNESS1 1Institute of Biomedical and Life Sciences, Graham Kerr Building, University of Glasgow, Glasgow, G12 8QQ, UK, 2School of Biology and Psychology, Ridley Building, Newcastle University, Newcastle upon Tyne, NE1 7RU, UK, 3Ecology Division, Forest Research, Northern Research Station, Roslin, Midlothian, EH25 9SY, UK and 4British Trust for Ornithology, The Nunnery, Thetford, Norfolk, IP24 2PU, UK Capsule Siskins and Coal Tits were present in gardens more often in years of poor Sitka Spruce cone crops, with synchrony across the UK. Aims To examine how the relative abundances of Siskins and Coal Tits in a garden in western Scotland varied annually with the size of the spruce cone crop, and to determine whether this pattern was similar nationally. Methods A ten-year mist-netting data set from Tarbet, Argyll and Bute was used to test the hypothesis that some species utilize supplementary food in gardens more often in years with poor cone crops than in years when crops are large. Annual peaks in the relative abundances of four species (two that ate conifer seed and two that did not) in Tarbet were correlated with a Sitka Spruce masting index. BTO Garden BirdWatch data from six regions of the UK were correlated with the Tarbet data, the cone index and each other to determine the extent of synchrony. Results Both Siskins and Coal Tits were present more often in the Tarbet garden in years with poor cone crops, with relative abundances significantly negatively correlated with cone crop for both species (r = –0.739 and –0.832, respectively). Correlations between regions using BTO data showed abundances of species in gardens to be broadly similar across the UK. Independent scores of cone abundance in Kielder and Galloway were correlated. Correlations between BTO data, cone index and Tarbet garden data were mostly high and almost all in the same direction, also indicating a degree of synchrony in cone production across the UK. Conclusion Both Siskins and Coal Tits appear to be influenced by changes in cone abundance, switching to feed on supplementary food in gardens more often in years with few cones than in mast years. While such a relationship has been shown previously for Siskins, for Coal Tits this is a novel result. Over the last 80 years, the area of forest in the UK has Petty & Avery 1990). Conversely, many woodland increased from 5.0% of the land surface in 1924 to species have benefited, including some with previously 11.5% in 2004 (Forestry Commission 2004). Most of restricted distributions, or even species that had ceased this increase has been accomplished by establishing to breed in Britain (Jardine 1993, Knox 1993, Petty forests of non-native species on marginal agricultural 1996). land in the uplands (Petty & Avery 1990). The estab- Sitka Spruce Picea sitchensis, a native of northwest lishment of such habitats has been criticized because of America, has been one of the main tree species used the negative impact on several moorland bird species, in Britain’s reforestation programme because of its some of high conservation value (Nature Conservancy exceptional growth rate and high quality timber Council 1986, Stroud et al. 1987, Thompson et al. 1988, (Cahalan 1987, Rook 1992). Today it constitutes 29% of the total forest area of the UK, and 47% of the for- *Correspondence author. est area in Scotland (Forestry Commission 2004). It Email: [email protected] produces prolific cone crops at intervals of three to © 2007 British Trust for Ornithology Value of Sitka Spruce seed for Siskins and Coal Tits 237 five years that appear to be synchronized over large broadleaved forests (Perrins 1979), but similar to geographical areas (Matthews 1955, Philipson, 1987, Marsh Tits Poecile palustris and Willow Tits Poecile Petty et al. 1995, Lurz et al. 2000). Sitka Spruce has montanus (Perrins 1979). relatively small seeds compared to widespread conifer The practice of feeding birds in gardens has become species native to Europe such as Scots Pine Pinus more popular over the last 30 years, yet it is only now sylvestris, Norway Spruce Picea abies and European that its effects are being considered scientifically Larch Larix deciduas (Staines et al. 1987). Sitka Spruce (Cannon 1999, Bland et al. 2004). It has been suggested is also one of the first conifers to shed its seed in that when natural foods are scarce, birds will utilize autumn. Therefore, it is of considerable importance to supplementary foods to a much greater extent than better understand how the potentially enormous food when natural foods are plentiful (Brittingham & resource on Sitka Spruce is utilized by Britain’s wildlife, Temple 1988, 1992a, Chamberlain et al. 2005). This as many characteristics differ from those of native was found to be the case for Siskins, with virtually no European conifers with which conifer seed-eating birds using feeders in good cone years (Shaw 1990). species have co-evolved. Years when the highest numbers of Siskins were In northern Europe, many birds that exploit conifer observed in the forest coincided with years when the seed exhibit rapid population growth rates in years with cone crop was particularly good (Shaw 1990). a good cone crop, due to extended breeding seasons and Although a similar relationship has been suggested in low mortality (Newton 1972). In poor cone years, a Coal Tits, no analysis has been undertaken to investi- range of adaptations have evolved to overcome periods gate whether they too switch between cones and when conifer seed is unavailable, such as nomadic garden feeders depending on cone abundance. movements (Newton 1970), scatter-hoarding of conifer In this paper, we use a large data set of birds mist- seed (Moreno et al. 1988, Perrins 1979) and switching netted in a garden in western Scotland (Tarbet), a to other foods (Newton 1972). location surrounded by extensive Sitka Spruce forests, In Britain, an increasing range of bird species has to investigate relationships between cone crops and the been shown to exploit Sitka Spruce seed (Shaw & occurrence of Siskins and Coal Tits at bird-feeders. Our Livingstone 1991). Two of the best known examples hypothesis was that both species would use feeders in are Common Crossbill Loxia curvirostra and Siskin greater numbers when the cone crop was poor. As the Carduelis spinus, both of which have greatly increased data set contained information on a large number of their breeding ranges in Britain in response to the other species, a further two species (Blue Tit and Great increasing area of cone-producing forest, with numbers Tit) were used as controls. It was predicted that dramatically increasing in good cone years (Jardine the abundance of both would be unrelated to Sitka 1993, Knox 1993). A number of studies have shown Spruce cone crop, as both species feed predominantly that population size and breeding performance of both in deciduous woodlands (Perrins 1979). species are closely linked to the coning frequencies of As cone crops appear to be synchronized over large spruce, both in Britain and mainland Europe geographical areas, it is likely that the cone data used (Reinikainen 1937, Newton 1972, Petty et al. 1995, in this study were representative of coning frequencies Virkkala 2004). A range of other bird species utilize over a larger area, comprising at least northern England Sitka Spruce seed when it is abundant, including Coal and southern Scotland. To investigate this possibility, Tits Periparus ater. Coal Tits have also increased their we used the British Trust for Ornithology’s (BTO) breeding range in Britain, as conifer forests are their Garden BirdWatch (GBW) data from six regions in the preferred breeding habitat, although the species occurs UK to see if peaks in reporting rate of the four species widely in other habitats outside the breeding season. (Coal Tit, Blue Tit, Great Tit and Siskin) were Similarly to other tits, Coal Tits feed largely on insects synchronized with the mist-netting data from Tarbet during the breeding season, but exploit seeds at other and the cone crop index. The GBW data were times of the year and have been observed feeding on independent of the mist-netting records and were Sitka Spruce seeds, usually from cones on trees (Perrins collected by a large number of people over an extensive 1979, Shaw & Livingstone 1991). They are known to area, thus permitting regional patterns to be explored. scatter-hoard seeds in conifer forests (Brotons & The likely effect of there being significantly less Sitka Haftorn 1999, Brotons 2000a, 2000b), unlike the Spruce present in the south of England compared to ubiquitous Blue Tit Cyanistes caeruleus and Great Tit northern Britain were incorporated into the analysis by Parus major, species characteristic of temperate using a general linear model (GLM). © 2007 British Trust for Ornithology, Bird Study, 54, 236–247 238 A.J. McKenzie et al. MATERIALS AND METHODS Kielder Forest, Northumberland (Petty et al. 1995). A similar assessment was carried out in Galloway by G. Study area and catching protocol Shaw from 1982 to 2001 after which time R.W. Furness Birds were caught in a mist-net and ringed by R.W. continued recording in Argyll. Petty and Shaw estab- Furness in a garden in Tarbet, near Loch Lomond in lished criteria for a cone crop index running from zero western Scotland (56°12′N, 4°42′W), between January to 10 (see Table 1 for definitions of each increment). 1995 and October 2004. The net was set close to bird- The index used in this analysis comprised the Petty feeders and captured birds were removed, identified, data during 1994/5–1997/8, the Shaw data during sexed and aged (if possible), ringed or ring read, 1998/9–2001/2 and the Furness data during 2002/3 and weighed and released.
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