The Decline of the Willow Tit in Britain Alex J
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Conservation Priority Species The decline of the Willow Tit in Britain Alex J. G. Lewis, Arjun Amar, Elisabeth C. Charman and Finn R. P. Stewart ABSTRACT Numbers of the British Willow Tit Poecile montanus kleinschmidti have decreased dramatically in recent decades.This paper outlines the three main hypotheses for the decline – increased competition, increased predation and changes in habitat availability – and describes the fieldwork and analyses undertaken to try and understand its cause.The results so far are not conclusive but indicate that the loss of young, damp woodland may have been important. Possible positive management strategies are suggested. Introduction the first study to observe such a high number of Range and habitat natural nests in Britain and thus provides an The British race of the Willow Tit Poecile insight into the species’ breeding ecology. montanus kleinschmidti is found in England, Willow Tits remain in their territories from Wales and parts of southern Scotland, including their first breeding year and start to sing in Dumfries & Galloway and North Lanarkshire January when the weather is good (Maxwell (Clyde). The species as a whole is found 2007). Between April and May, they excavate a throughout much of the Western Palearctic and nest hole in dead wood. The most frequently east through northern Asia as far as Japan, where chosen tree species in the Midlands study area it is found largely in boreal forest with conifers were willow Salix spp. (n=20), Elder Sambucus (Snow 1954; Alerstam et al. 1974; Snow & niger (n=10) (plate 211) and Silver Birch Betula Perrins 1998; Sellers 2002). A recent study in pendula (n=16) (plate 212), but nests were also northern Finland (Siffczyk et al. 2003) showed found in Alder Alnus glutinosa (n=3), Hawthorn that in a ‘mosaic forest landscape’, Willow Tits Crataegus monogyna (n=2), Hazel Corylus preferred mature woodland (and pine bogs) and avellana (n=2), Wild Cherry Prunus avium avoided young stands. In Britain, however, the (n=1) and a conifer (n=1). The nest hole is species is associated almost exclusively with excavated in either a dead tree or a dead part of early successional habitats such as those found a live tree, or occasionally in a felled trunk (plate along river valleys, overgrown flooded gravel- 213). Willow Tits can nest successfully in a trunk pits and wet woodland (Snow & Perrins 1998). with a diameter as small as 5 cm (measured at the part of the trunk corresponding to the base Breeding ecology of the nest chamber), although the mean Three separate but related studies of Willow Tits diameter recorded was 12 cm. were commissioned by the RSPB and took place Willow Tits line their nests with material such in 2005 and 2006, and form the basis of this as feathers, animal hair, thin strips of bark and review. One of these was carried out in the the fluffy seeds from the heads of bulrush Typha Midlands (South Yorkshire, Derbyshire and (plate 214). The female incubates for approx- Nottinghamshire) and involved the location and imately 14 days (Maxwell 2007). Of 56 nests observation of 56 Willow Tit nests during the studied, it was possible to inspect only 20 two breeding seasons (Lewis unpubl.). This was internally (with an endoscope; plate 215) so that 386 © British Birds 102 • July 2009 • 386–393 The decline of the Willow Tit in Britain Alex Lewis Alex Lewis Alex Lewis 211–213. Willow Tit Poecile montanus nests in Elder Sambucus niger (211), Silver Birch Betula pendula (212) and a felled Hawthorn Crataegus monogyna (213), Nottinghamshire, April 2006. clutch size could be determined. Clutch size was remain in the breeding territory all year round. found to be 8.8 ± 0.4 (mean ± 1 s.e.) and this was The few ringing recoveries for this species not related to tree diameter (as measured above). support the finding that Willow Tits are In all cases, other than that described in Lewis & sedentary (but that juveniles in late summer or Daniells (2007), it was impossible to calculate autumn will occasionally move more than 5 km; brood size and number of fledglings owing to: Sellers 2002). (i) the way that the chicks were invariably sitting on top of one another inside the chamber; and Population status (ii) the need to avoid excessive disturbance that An analysis of Common Birds Census (CBC) and prolonged viewing may have caused. Breeding Bird Survey (BBS) data has shown that After the eggs hatch, the chicks remain in the the British Willow Tit underwent a dramatic nest for approximately 18 days (plate 216). Once decline of 88% between 1970 and 2006 (Eaton et fledged (plate 217), they remain in the al. 2008). Ringing data provide further evidence of immediate area for up to 20 days, while they a substantial population decline (Perrins 2003). In continue to be fed by both parents (Maxwell fact, of all the species monitored adequately by the 2007). At the end of the breeding season, British BBS, the Willow Tit showed the greatest decline Willow Tits differ from their European (77%, 1994–2007; Risely et al. 2008). The Repeat counterparts in that they do not form gregarious Woodland Bird Survey, which looked at social groups, neither with conspecifics nor with population changes in woodland habitat only, also other tit species (Perrins 1979; Sellers 2002), and recorded a severe decline, of 70% between the mid British Birds 102 • July 2009 • 386–393 387 The decline of the Willow Tit in Britain 1980s and 2003/04 (Hewson et al. 2007). As well as losing a nest hole following its completion can a reduction in numbers, the Willow Tit has result in breeding failure. Between 1995 and suffered a marked contraction in its range in 2000, Maxwell (2002) monitored 30 Willow Tit Britain and has been lost entirely from many pairs using both natural nests and nestboxes. He counties (Gibbons et al. 1993; Smith et al. 1993). found that only ten of these pairs were success- Owing to its population decline, the Willow Tit ful. Of the 20 unsuccessful pairs, 18 had their appears on the UK’s ‘Red list’ of birds of nest cavity taken over by Blue Tits and two by conservation concern (Eaton et al. 2009) and is a Great Tits. Blue and Great Tit populations have priority species in the UK Biodiversity Action increased in the UK between 1970 and 2006 Plan. However, despite the decline in numbers and (Blue Tits by 33% and Great Tits by 91%) range contraction, Willow Tits still appear to be (Eaton et al. 2008). Consequently, interspecific doing well in parts of the Midlands. Consequently, competition for nest-sites may have increased much of the recent work has focused on these and contributed to the Willow Tit’s decline. apparently stable populations. Predation Potential threats to Willow Tit populations Once a nest hole has been excavated and lined, Research into the decline of the Willow Tit has Willow Tits can still be noisy around their focused on three hypotheses: increased breeding site, leaving them vulnerable to competition, increased predation and habitat detection by Great Spotted Woodpeckers change. Dendrocopos major, which are accomplished at extracting prey from rotten wood (Wesołowski Competition 2002; Fuller et al. 2005). This species will destroy The excavation of a nest hole can be a noisy a Willow Tit nest with ease and take the eggs or activity, as both birds often call repeatedly to chicks (Lewis unpubl.) (plate 218). Willow Tits each other. Typically, the process is also visually are single-brooded and if predation occurs at obvious as it involves much physical activity and the chick stage, the pair is extremely unlikely to continual production of small wood chippings. breed again that year. Even if predation occurs at An excavating Willow Tit pair is thus vulnerable the egg stage, the pair will have limited resources to detection by competitively superior Blue (of time and energy) for another attempt. Cyanistes caeruleus and Great Tits Parus major.If Numbers of Great Spotted Woodpeckers have either of these species finds a Willow Tit nest increased dramatically in the UK (by 314% hole, they can oust the occupants with little between 1970 and 2006; Eaton et al. 2008), and difficulty (Maxwell 2002). Excavating a nest hole Willow Tits may have suffered a corresponding is a time- and energy-consuming process and increase in predation rate. Habitat Willow Tits are traditionally associated with damp, scrubby areas in Britain (Perrins 1979; Snow & Perrins 1998). Despite the steep pop- ulation decline, they can still occur at relatively high densi- ties in some brownfield sites and disused gravel-pits, where such habitat is often charac- teristic (plate 219) (Lewis unpubl.). However, these disused industrial sites have become less common in recent decades, having been lost to (for example) urban Alex Lewis development and agricultural 214. Eight Willow Tit Poecile montanus chicks in a nestbox,Wiltshire, May 2005.The bedding is made up of feathers and thin strips of bark. clearance (Barr et al. 1993). 388 British Birds 102 • July 2009 • 386–393 The decline of the Willow Tit in Britain This apparent wasteland does not have the same unoccupied patches represent unfavourable perceived biodiversity value as more established habitat; in areas which are sparsely populated habitats such as woodland (Mortimer et al. owing to factors other than habitat (such as 2000) and the loss of large areas of such scrub predation), many favourable sites may be unoc- could be driving the Willow Tit’s decline. cupied and thus wrongly classified as unfavourable. A large number of habitat vari- Testing the hypotheses and exploring the ables were quantified (including tree diameter, causes of the decline tree-stem density and tree species composition, Data analysis canopy cover, soil-water content, dead-wood Siriwardena (2004) showed that the major abundance and vegetation cover at varying decline in Willow Tit numbers had occurred in height bands) in both occupied and unoccupied woodland and farmland, and that populations sites, and in areas within each occupied site in wet habitats had remained stable.