Review of Resource Requirements for Target Woodland Bird Species

Home , Common redstart, Lesser redpoll, Nightingale (software), Spotted flycatcher, Thrush nightingale, Tree pipit

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APPENDIX 1

Review of resource requirements for target woodland bird species

Summary

1. This review summarises published information on the resource requirements and habitat associations of 17 target species which represents woodland birds of conservation priority and species that use the lower vegetation layers most likely to be affected by deer browsing. These species were Willow Tit, Marsh Tit, Willow Warbler, Garden Warbler, Song Thrush, Spotted Flycatcher, Nightingale, Dunnock, Tree Pipit, Lesser Redpoll and Bullfinch. The remaining species were Nightjar, Lesser Spotted Woodpecker, Wood Warbler, Redstart, Pied Flycatcher and Hawfinch. The review provides information on habitat structure and resource use to supplement results from an analyses of habitat associations from an extensive dataset of 300 woodland plots, particularly for scarce species which occurred on few plots.

2. Information sources included scientific journals, RSPB research reports and Books. Journal articles were identified using Web of Science, Zoological Record and Google Scholar search engines with the current species scientific names and recent synonyms as search terms. Further sources were RSPB reports on habitat requirements, ornithological handbooks and species monographs. For each target species the literature was searched to provide information on their nesting, feeding and territory requirements as well as broader habitat associations and behaviour which is likely to be relevant in determining responses to changes in woodland structure.

3. Four species (Nightjar, Wood Warbler, Willow Warbler and Tree Pipit) nest on or near the ground. Six species (Garden Warbler, Song Thrush, Nightingale, Dunnock, Lesser Redpoll and Bullfinch) nest in a dense shrub layer. Seven species (Lesser Spotted Woodpecker, Willow Tit, Marsh Tit, Spotted Flycatcher, Redstart, Pied Flycatcher and Hawfinch) were canopy or hole nesters.

4. All species except Bullfinch and Hawfinch predominately feed on invertebrates during the breeding season and also feed their nestlings invertebrates. Garden Warbler, Song Thrush, Nightingale and Spotted Flycatcher also feed on fruits outside the breeding season, especially prior and during migration. Willow Tit, Marsh Tit, Lesser Redpoll and Dunnock diets also contain a considerable proportion of seeds outside the breeding season. Bullfinch and Hawfinch feed mostly on seeds throughout the year, but also feed on buds, flowers and invertebrates when seeds are scarce in spring. Lesser Spotted Woodpecker, Wood Warbler and Willow Tit feed in the canopy while Dunnock, Song

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Thrush, Redstart, Nightingale and Tree Pipit forage mainly on the ground. Garden Warblers mainly forage in the shrub layer. Nightjar and Spotted Flycatcher are mainly aerial foragers, Nightjar in the open and Spotted Flycatcher within the canopy. Lesser Redpoll feeds on seeds and invertebrates mainly in the tree canopy but also on the ground when all seed has fallen. Bullfinch, Marsh Tit and Pied Flycatcher forage in a wide range of heights from the field layer to the canopy depending on food sources at different times of year.

5. In the breeding season, Nightjar, Willow Tit (in the UK), Willow Warbler, Garden Warbler, Song Thrush, Dunnock, Tree Pipit and Lesser Redpoll all show a preference for younger stages of woodland growth. Lesser Spotted Woodpecker, Wood Warbler, Marsh Tit, Spotted Flycatcher, Pied Flycatcher and Redstart show a preference for mature woodland while Bullfinch uses a range of different woodland types. Lesser Spotted Woodpecker, Marsh Tit, Wood Warbler, Garden Warbler, Nightingale and Pied Flycatcher prefer broadleaved woodland while other species used both broadleaved, conifer and mixed woodland.

6. Large scale timber harvesting such as clearfell and coppice favour populations of Nightjar, Willow Warbler, Garden Warbler, Song Thrush, Nightingale, Dunnock, Tree Pipit and Bullfinch. However, it can also negatively affect Marsh Tit and Spotted Flycatcher populations. Canopy thinning at silvicultural levels has only small effects on the target bird species. Effects of thinning may depend on woodland type and may only show significant effects with high levels of canopy reduction. Increased deer populations most strongly affect species which use the low shrub and field layer, in particular Willow Tit, Willow Warbler and Nightingale. High levels of deer browsing have been shown to negatively affect Willow Warbler, Garden Warbler, Song Thrush, Nightingale, Dunnock and Bullfinch. Higher Reeves’ muntjac density has been shown to negatively affect abundance of Willow Warbler and Song Thrush, and higher Roe deer density has been shown to negatively affect abundance of Willow Tit and Nightingale. Nightjar, Wood Warbler, Redstart, Pied Flycatcher and Tree Pipit may benefit from higher levels of deer browsing but evidence is currently lacking. (See Appendix 2 for a review of the effects of woodland management and deer browsing on birds).

7. The following tables give a brief summary of habitat and resource requirements for each species, split into species associated with young woodland (Table A) and those associated with mature woodland (Table B).

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Summary Table A Habitat attributes and species requirement for target bird species associated with young woodland. Species are listed alphabetically.

Bullfinch Dunnock Garden warbler Nest construction Cup of fine twigs lined with grass and roots Cup of twigs and roots, lined with moss Grass and herbs lined with grass and hair feathers and hair Nest location In dense woody cover 1-2m from ground In dense low woody vegetation <3.5m. In low dense vegetation 0.5-1.5m When present, young spruce trees are favoured nest sites. Adult food Primarily seeds, but take buds , flowers and Mainly small ground dwelling Mainly invertebrates during breeding season. invertebrates when seed scarce in Spring invertebrates, e.g. beetles, spiders, Berries are important for fattening before springtails. Also small seeds in winter e.g. migration nettle & dock. Nestling food Initially invertebrates with increasing plant Mainly beetles, spiders, flies and Primarily caterpillars material (mainly seeds) with age caterpillars. Foraging methods Take seeds from perches off the ground. Feed Mainly picking items from ground surface. Forage by gleaning from foliage in at all heights from field layer to canopy. Also taking inverts from foliage in summer broadleaved trees and shrubs, 0.5-6m from or digging in soft soil in winter. ground. Territory attributes Non-territorial. Move between breeding Territories are centred on areas of dense Territories selected for dense low shrubs attempts to track food supplies shrub or young conifers. particularly bramble and birch Breeding habitat Wide range of habitats with trees and shrubs. Occur in a wide range of habitats with Prefer broadleaves and young stages of forest Dense shrub layer essential. dense shrub cover. Will occur in all types of growth. They will use older stands of forest but only where dense low cover is broadleaf with dense shrub layer and avoid available. mature conifers. A dense low shrub layer is essential. They are more likely in more wooded landscapes and away from woodland edges. Post fledging Unknown Unknown Unknown Seasonal differences Move between habitats to track locally Largely sedentary but move from uplands During migration select areas with high abundant seed sources in winter fruiting shrub diversity UK Distribution Widespread throughout UK Widespread throughout UK Widespread in most of UK, scarce in northern Scotland and Ireland

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Summary Table A Continued.

Lesser redpoll Nightingale Nightjar Nest construction Open cup of twigs lined with hair feather or Nest of dead leaves lined with grass hair & Eggs laid on moss or bare ground plant material feathers Nest location In small tree or shrub Nest on or near the ground, <30cm from Nest located in woodland clearing, often near ground. Nest usually in the dense field layer a solitary tree. Nest on flat ground and avoid at the edge of a dense shrubby thicket, slopes. Adult food Mainly invertebrates during breeding A wide variety of small invertebrates taken, Large flying insects predominately moths and season. During winter they feed on small predominately ants and beetles. Also feed on beetles tree seeds, e.g. birch, alder, larch, spruce, berries prior to and during migration. and herbaceous seeds. Nestling food Invertebrates Same as adult food Same as adult. Foraging methods Forage mainly in trees Primarily foraging for invertebrates on the Forage by aerial hawking. Forage mostly over ground or in leaf litter under dense cover. open semi natural habitats, avoid mature pine plantations and intensive agriculture. Territory attributes Unknown Territories centred on patches of dense Territories include open space and forest shrubs. Typically having a dense outer layer of edge. Open areas with low vegetation height foliage reaching the ground, with dense field preferred including heathland, clearfells, layer vegetation at the shrub edges. The roads rides and peatlands. interior of the shrub patch will have an open structure with bare ground. Breeding habitat They prefer conifer habitats to broadleaf Habitat is at low altitude mostly <60m amsl. Breeding habitat includes both forest and within conifers there is a strong Habitat is dense wetland scrub or young clearings and heathland. Habitat quality selection for young spruce 5-15 years. woodland including coppice. It must have related to amount of edge between forest Within restock plantations there is a high density of shrub foliage with some bare and open space and prefer heathland close to selection for higher conifer composition. ground below. Prefer scrub on damp soils woods. with high organic matter Post fledging Unknown male looks after fledglings while female starts young are independent two weeks after next clutch. Young independent after 15-30 fledging days. Seasonal differences Leave upland conifer plantations in winter. African migrant African migrant UK Distribution Scarce in southern and eastern England Only occurs in south and east England patchy distribution centred on larger areas of heathland and lowland conifer plantations

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Summary Table A Continued.

Tree pipit Willow tit Willow warbler Nest construction Cup of grass lined with hair and fine plant material Primary nest hole excavator in dead wood. Nest Dome of grass lined with feathers. lined with bark fur, feathers Nest location Nest is on the ground usually hidden in a grass Nest trees are mainly small diameter birch or On the ground in grass or low shrubs tussock. Orientation of nest entrance reduces shrubs e.g. hawthorn, sallow, elder. Holes usually exposure to the sun and prevailing winds <1.5m from ground Adult food Wide variety of invertebrates Mainly invertebrates but also eat seeds in Wide variety of invertebrates, including flies, winter, e.g. spruce, pine, and labiate species. caterpillars and spiders. Also feed on nectar and pollen in early spring. Prey changes from flying insects to caterpillars after bud burst. Nestling food Invertebrates, caterpillars early in season and later Invertebrates, mainly caterpillars, flies and Wide variety of invertebrates broods also fed on spiders grasshoppers and flies spiders Foraging methods Forage for insects on the ground. Occasionally On continent they feed in lower tree canopy. Most prey is taken from leaf surface either foraging in trees, particularly during caterpillar Unknown in UK, but likely to be in dense scrub. from in flight or perched. Prior to bud burst outbreaks. they will also probe in buds and catch insects in flight. Foraging is mostly in the lower half of tree crowns. Territory attributes Trees needed for song perches, occasionally use Unknown. Territories are centred on woodland with a wires or banks if trees missing in otherwise mean canopy height 1-10m, optimal 5-7m. suitable habitat. Densities of trees vary greatly but Territories are also associated with the generally avoid areas with >60% tree cover. presence of birch trees. Breeding habitat Occur in a wide variety of habitats with low tree In UK ,they occur in wetlands, woodland and They occur in young plantations, density e.g. upland woodland margins, heathland, farmland. Primarily in areas of scrub with higher broadleaved and conifer woodland and conifer clearfells, chestnut coppice. Also, occur in soil moisture and denser shrub cover at 2-4m. scrub. They prefer early successional stages. low density woodland e.g. native pine woods and More stable populations in more wooded In conifers they prefer pine woodland and upland oak woods. In conifer plantations optimal landscapes. Continental populations also breed occur in areas with some birch or other habitat is 2-5 year regrowth. in conifer forest with birch. broadleaves. They occur in small woods but they prefer larger patches of suitable habitat in large woods. Post fledging move up to 600m from nest within week of Dependent on parents for 7-11 days. Stay on Nestlings gradually increase ranging fledging natal territory for further 1-2 weeks before distances from nest site with time. Young dispersing. Settle on winter territory within six dispersed more than 1km after 1 month. weeks of fledging. Seasonal differences African migrant Resident on territory year round. In winter they African migrant have larger overlapping home ranges. UK Distribution Scarce in lowland Britain, absent from Ireland Core range in south Wales, midlands and north Widespread across UK east of England, scarce elsewhere.

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Summary Table B Habitat attributes and species requirement for target bird species associated with mature woodland. Species are listed alphabetically. Redstart Lesser spotted woodpecker Hawfinch Nest construction Cup of grass, roots and moss lined with feathers Primary nest hole excavator. Holes excavated in Base of sticks with a cup of roots and grass, and fur dead wood decorated with lichens Nest location Large holes in trees and buildings, < 3m from In dead limbs within live canopy or dead trees. Nest in trees shrubs and creepers. No apparent ground Branch cavities are usually on the underside near selection for nest location. They occur in a wide the trunk in limbs of 15-50 cm diameter. Nests variety of tree and shrub species at all heights and are normally in upper half of tree. In UK there is many locations on the tree. a selection for Oak as nest tree. Adult food Mainly a variety of invertebrates, but take some Wide variety of invertebrates taken throughout Kernels of tree and shrub seeds, e.g. cherry, berries on migration. the year. hornbeam beech yew, holly. When seed scarce in spring they feed on tree buds, flowers and caterpillars. Nestling food Wide range of invertebrates, particularly Caterpillars and other invertebrates Primarily caterpillars caterpillars, beetles spiders and ants Foraging methods Mostly takes prey from the ground, hunting from Mostly forage on smaller twigs and branches, Prefer to feed in canopy only feeding on fallen perches. Also gleans from many surfaces and aerial taking invertebrates from under bark. In the seeds in late winter. sallies. breeding season switches to surface gleaning from bark and foliage. Mostly forage on broadleaf trees, in England predominantly Oak. Territory attributes Territories centred on areas with bare ground or Breeding territories are 20-50 ha. Weakly territorial, largely defending female and sparse vegetation only fixed to an area of woodland once nest building started. Breeding habitat They are more likely to breed in larger areas of They are only found in lowland broadleaved Mainly associated with woodlands containing woodland >5ha. Associated with woodlands with woodland and favour large woods in a well mature seeding trees, either as parkland, open canopy, sparse shrub layer and field layer wooded landscape. In the UK, they are plantation or semi natural woodland. Tree with bare ground or sparse vegetation. Higher associated with oak dominated woodland with diversity is likely to be important but weak densities in broadleaf than conifer. abundant deadwood. Unmanaged woodland is evidence. better than managed. Post fledging Fledglings independent after 2 weeks, stay Eruptive dispersal of juveniles in autumn on Leave nesting area within a few days. together for 1 month and start migration after 7-8 continental Europe. weeks. Seasonal differences African migrant In winter they range over several hundred Use similar woodlands all year, but will fly many hectares but remain in broadleaved woodland. kilometres to roost in dense conifers during winter. UK Distribution Mainly in north and west Britain, absent from Only occurs in lowlands of England and Wales Very local distribution in England, Wales and Ireland Scotland, scarce in most regions.

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Summary Table B Continued.

Marsh tit Pied flycatcher Song thrush Nest construction In small diameter holes, moss cup lined with fur Secondary hole nester, which in many areas Cup of grass and roots lined with mud. prefers man made nest boxes. Holes used have approximately circular entrance >3.5 cm diameter Nest location Nest holes are in broadleaved trees. they are Most nest holes >2m from the ground. Often Nest located 1-5m from ground in tree or shrub. mostly naturally occurring holes in live trees and in woodpecker holes in the trunks of a wide variety Usually in dense vegetation, often conifers such as the lower half of the tree. of tree species. spruce or juniper. Adult food Small invertebrates of wide variety of taxa during A wide variety of invertebrates. In breeding season diet is predominately the breeding season. In winter this is invertebrates. In autumn and winter they will supplemented with seeds from shrubs and additionally feed on berries and fruit. herbaceous plants. Nestling food Predominantly caterpillars and spiders Primarily caterpillars but the proportion in the Mainly invertebrates, including earthworms, snails, diet declining through the season, being replaced insect larvae, and beetles with spiders, ants and beetles. Foraging methods Invertebrates are taken from upper two thirds of Foraging techniques vary with woodland type Feed on the ground taking invertebrates from the the trees by gleaning from bark and twigs. Fruit and prey type. Feeding on the ground, gleaning surface or probing in soil. Will also glean seeds are removed from the pulp and seed coat from trunks and foliage and aerial sallies all caterpillars if abundant and catch flying insects removed. Temporary abundant seeds are cached. frequently used. during dry weather. Forage within 50m of shrub cover. Territory attributes Territories are 2.5-8 ha. They avoid extensive areas Territories are established in mature woodland Select for areas with greater amount of dense of young tree growth. They select for areas of with low shrub cover. In coniferous woodland shrub cover, e.g. hedgerows or young spruce. maximum canopy height with dense shrub layer. their territories have at least 45% pine. Avoid areas near bird of prey nest sites. Breeding habitat More likely to breed in woods >10 ha. They prefer Breeding habitat includes both mature Breed in a wide variety of habitats with shrubby mature broadleaved woodland but will nest in pine broadleaved and conifer woodland with thin cover, e.g. farmland, gardens, woodland. Strong and mixed conifer woodland. They select for shrub cover. Higher densities are found in preference for woodland edges and more frequent woods with higher canopy height and cover with a broadleaves. Scots pine preferred in conifers. in small woods. Also, breed in conifer plantations, dense shrub layer. Avoid small woods <1 ha. preferably 5-15 yr spruce. Post fledging Fledglings are independent after two weeks then Fledglings stay in territory up to 27 days then Juveniles disperse <20km within few months of disperse to find their winter territory within disperse into surrounding habitats. fledging. another two weeks. Seasonal differences They are sedentary but winter ranges are larger African migrant Use similar habitats in winter. Short distance than breeding territory. partial migrant. UK Distribution Largely confined to lowlands of England and Wales Largely confined to Wales, western England and Widespread throughout UK south west Scotland

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Summary Table B Continued.

Spotted flycatcher Wood warbler Nest construction Grass moss and spider web cup lined with feathers Dome of grass and other vegetation lined with feathers Nest location Nest is on buildings and trees often concealed in a On the ground in grass tussock, leaf litter or under fallen cavity or within creepers. Nest height 1.5-10m. branches. Adult food Mainly feed on flying insects preferring larger flies. A wide variety of invertebrates manly winged insects, Also take berries before and during migration. caterpillars and spiders. Will also take some small berries eg. elder and bramble before migration. Nestling food Preferentially fed large flies but also a range of Mainly caterpillars, winged insects and spiders other invertebrates. Foraging methods Two principal foraging methods areal sallies and Mainly forage in the canopy gleaning from leaves and foliage gleaning. Gleaning used when insect branches. They will also take insects in flight. activity low. Territory attributes Select for areas with large trees and open space. Territories are situated in woodland with high canopy cover and low density of shrub layer. Lower occupation of territories in high rodent years on the continent. Breeding habitat Broad habitats include gardens, farmland and Highest densities in oak woodland which is occupied first, woodland. They breed in both mature broadleaf but also use a variety of broadleaved conifer and mixed and semi-natural pine woods. Structural attributes woodland types. Habitat has high canopy cover and include large mature trees, gaps in the canopy and density, low shrub density and is often situated on slopes. low shrub density. Post fledging Unknown Can remain as family groups up to four weeks. Seasonal differences African migrant African migrant UK Distribution Widespread throughout UK Largely confined to western and upland Britain

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Contents Summary ...... 1 Introduction ...... 10 Aims of the review ...... 10 Species included in the review...... 10 Species requirements included in the reviews ...... 10 Methods ...... 11 Results ...... 11 Overview of the requirements of key bird species and the effects of forest management and deer populations ...... 11 Species nesting on or near the ground ...... 15 Nightjar ...... 15 Tree Pipit ...... 21 Wood Warbler ...... 25 Willow Warbler ...... 29 Species nesting in a dense shrub layer ...... 35 Garden Warbler ...... 35 Song Thrush ...... 39 Nightingale ...... 44 Dunnock ...... 49 Lesser Redpoll ...... 53 Bullfinch ...... 55 Hole nesters and species nesting in the tree canopy...... 59 Lesser Spotted Woodpecker ...... 59 Willow Tit ...... 64 Marsh Tit ...... 68 Spotted Flycatcher ...... 72 Redstart ...... 76 Pied Flycatcher ...... 81 Hawfinch ...... 88

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Introduction

Aims of the review For all target species this review aims to identify species requirements expected to be of interest in interpreting responses to change in habitat due to forest management and deer browsing. This information can be used for interpolating effects of management and deer browsing on vegetation structure based on vegetation changes recorded, and to likely species-specific responses of species that occurred infrequently during field survey. The information will also be used for interpreting results from the field survey to suggest mechanisms explaining any effects recorded. The review also aims to identify knowledge gaps in understanding requirements for these species. It is also intended to catalogue research projects with datasets for these species previously carried out by RSPB and BTO.

Species included in the review. The species included in this review are the 17 species agreed to be target species for the contract. Those in the species scope of the contract are declining woodland birds of conservation concern. Target species for the field survey are species likely to be influenced by excessive deer browsing and forest management and also likely to occur in sufficient woodlands to allow detailed analysis. All species are widespread woodland birds with clear known preferences for different woodland structures.

1. Nightjar 2. Lesser Spotted Woodpecker 3. Willow Tit 4. Marsh Tit 5. Wood Warbler 6. Willow Warbler 7. Garden Warbler 8. Song Thrush 9. Spotted Flycatcher 10. Nightingale 11. Redstart 12. Pied Flycatcher 13. Dunnock 14. Tree Pipit 15. Lesser Redpoll 16. Bullfinch 17. Hawfinch

Species requirements included in the reviews Alterations to vegetation due to forest management and deer browsing have the potential to affect bird habitats in a number of ways. Changes in vegetation may alter nest site suitability, and affect the abundance and availability of food through foraging preferences and behaviour. Substantial change in habitat may affect either suitability for breeding or suitability outside the main breeding season. Each species review summarises information found under the seven headings below and is concluded with an assessment of likely positive or negative impacts from forest management and deer browsing based on the

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separate review of the impacts of forest management and deer browsing on woodland bird habitats.

Categories of information searched for in the review: 1. Nest site selection/requirements 2. Territory selection/attributes 3. Main food items 4. Foraging methods 5. Breeding habitat 6. Post fledging behaviour/requirements 7. Seasonal differences in habitat for resident species 8. Effects of forest management and deer on habitat

Methods Published literature was searched for using Web of Science and Zoological Record online search engines. For each species the current scientific name and any synonyms used in past 50 years were the search terms entered in the search engines. Resulting titles and abstracts were screened to identify papers likely to contain suitable information and details of these were stored in EndNote bibliographic management software. As well as species-specific studies, information of relative abundance in different types of forest were taken from studies of forest bird communities. Results were tabulated by individual species. Finally information from research reports and other known unpublished sources were incorporated. All studies from the UK and continental Europe that were published in English or had good English language summaries and table headings were considered. Where different subspecies occur on continental Europe and the requirements apparently differ from those of the subspecies present in the UK, the information is incorporated in the review but made clear that requirements may differ between races.

Results

Overview of the requirements of key bird species and the effects of forest management and deer populations

Nest site selection and requirements Four target species are ground nesters, six nest in a dense shrub layer and seven are canopy or hole nesters. Of the ground nesting species, Wood Warbler was the only species which exclusively nests under closed canopy woodland also requiring the presence of a small tree or shrub close to nests for perching. Nightjar, Willow Warbler and Tree Pipit typically nest in young woodlands and clearings. Although forest management could have a detrimental impact during forestry operations Nightjar, Willow Warbler and Tree Pipit all benefit from the open conditions created by clearfell, group fell, coppicing or from new and secondary planting. The benefit for Willow Warbler occurs after broadleaved scrub develops. Nightjar and Tree Pipit directly benefit from open ground created by clearfelling followed by replanting, in both broadleaved and coniferous woodlands. No studies show negative effects of deer on nest site selection through nest trampling although this was only examined for Wood Warbler, and all may benefit from deer browsing as this prolongs the

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availability of open areas and early scrub stages, although this was not specifically examined by any studies.

Dense scrub nesting species were not obligate woodland nesters as they also nest in other habitat types, but within woodlands mainly occupy broadleaved woodland of a variety of ages. Nightingale have the most specialised nesting requirement as nests need to be located near to taller vegetation to provide cover for fledged young. Coppice management is typically most beneficial 3-10 years after cutting, for Willow Warbler and Nightingale, but after this time canopy closure makes habitat less suitable for nesting. Deer browsing has been shown to negatively affect Willow Tit, Song Thrush, Nightingale and Dunnock, particularly where browsing is heavy enough to alter understory vegetation structure, although for Willow Tit and Song Thrush the mechanisms have not been investigated.

Canopy and hole nesting species are obligate woodland species with a strong but not exclusive preference for mature broadleaved woodland. The exception to this was Willow Tit which occupy low dense scrub woodland and nest in holes at lower heights. Although hole nesters nest in dead wood none did exclusively; Lesser Spotted Woodpecker nest mainly in dead limbs but of live trees, and so standing dead wood is not a fundamental nesting requirement. Coppiced and young or replanted woodland is unsuitable for hole nesting species unless reasonable densities of standard trees or old coppice stools are present . All except Willow Tit nest above deer browsing height and so are not likely to be significantly affected by deer.

Territory selection Factors influencing territory selection vary considerably between species, with target species showing preferences for both open and closed canopy woodlands, woodlands with intermediate cover and woodland with open spaces at ground or canopy levels. Six species show strong preference for young woodland, and five for mature woodland. Nine species show strong preference for broadleaved woodland, and Nightjar and Lesser Redpoll less strong associations with coniferous woodland. No species was exclusive to woodland areas of any particular tree species and general associations found probably reflect characteristics of study sites rather than actual preferences. In the UK however Lesser Spotted Woodpecker and Pied Flycatcher have a strong association with oak woodland, but this is less strong in continental studies.

Forest management plays a significant role in creating preferred territories for four species. Tree Pipits preferentially select territories within young plantations 2-5 years old. Nightjar show strong selection for new plantings 10-15 years old. Willow Warbler and Nightingale preferentially select territories in coppiced woodland 3-10 after harvesting. To a lesser extent Song Thrush and Redstart benefit from edges created by forestry management including clearfell and thinning. Heavy deer browsing may potentially benefit species requiring an open understory and bare ground such as Nightjar, Wood Warbler, Redstart and Pied Flycatcher.

Main food items All species except Bullfinch predominately feed on adult and larval invertebrates during the breeding season, and also feed nestlings on invertebrates. The most specialised species are

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Nightjar, which has a very strong preference for adult moths, and Lesser Spotted Woodpecker which shows strong relationships with a few cyclically occurring Lepidoptera species in continental Europe. Caterpillars are a significant food for Lesser Spotted Woodpecker, Wood Warbler, Pied Flycatcher and Hawfinch but caterpillars are substituted with other invertebrates as the supply declines during the breeding season. This is especially true in broadleaved woodland. Five target species (four of which are migratory) fed on fruits, mainly post-breeding, and there was some evidence indicating the importance of fruit in diet for mass gain prior to autumn migration. A wide variety of fruits are taken by these species with Bramble a frequently recorded species. Only Bullfinch feed on seeds throughout the year, only rarely feeding on invertebrates.

The effect of management and deer on food availability and diet was not explicitly tested by any study. Management resulting in little understory is likely to be detrimental to availability of fruits and clearfell is likely to reduce the availability of saproxylic invertebrates and invertebrates with life stages requiring dead wood. Deer browsing may indirectly affect availability of many invertebrates, particularly species requiring herb, shrub and tree species of the understory which may be depleted by deer. It is possible that changes in available foliage resulting from browsing may affect availability of invertebrates and berries but such relationships were not examined in the studies found.

Foraging methods Three species mainly feed in the canopy, four mainly on the ground, two mainly aerially, one mainly in the scrub layer and four have no height preference. In addition, Nightingale feed on the ground and in low vegetation, requiring the availability of both for nesting. Foraging requirements for most species depend to some extent on woodland type and seasonality, and for some species weather conditions, with for example a shift from aerial invertebrate feeding to gleaning and ground foraging in cooler or wetter conditions when flying invertebrates are less available. Only Song Thrush is a frequent soil feeder although Redstart and Nightingale required areas of bare ground. Lesser Spotted Woodpecker most utilised dead wood, but mainly fed from foliage and branches of live trees. Lesser Redpoll, Bullfinch and to a lesser extent Hawfinch are most reliant on seeds as food and therefore forage more widely and seasonally to track available seed resources.

Foraging requirements can be affected by forest management and deer mainly by altering vegetation structure. Management has the greatest effect by altering canopy cover with some species requiring a closed canopy, and others requiring a more open canopy, or scrub and understory that can only develop with increased light levels associated with a more open canopy. Density of understory vegetation affects foraging habitat for Nightingale. Nightjar foraging habitat is created through clearfelling and the presence and maintenance of wide forest tracks.

Breeding habitat Lesser Spotted Woodpecker and Nightingale were found almost exclusively at lower elevations in the UK. Four species showed trends with woodland size, Lesser Spotted Woodpecker, Marsh Tit and Pied Flycatcher mainly only occurred in large woodlands with Song Thrush and Dunnock mainly more common in small woodlands. Lesser Spotted Woodpecker, Marsh Tit and Garden Warbler tended to be found in more wooded

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landscapes. Nightjar, Lesser Redpoll and Tree Pipit had strong associations with young planted coniferous woodland. Only one species, Willow Tit, was associated with woodlands with high soil moisture content. There was a suggestion that soil pH indirectly influenced Nightingale breeding habitat. Nightjar, Willow Tit (in the UK, in contrast to continental populations), Willow Warbler, Garden Warbler, Song Thrush, Dunnock, and Tree Pipit showed a preference for younger stages of woodland development. Lesser Spotted Woodpecker, Wood Warbler, Marsh Tit, Spotted Flycatcher, Pied Flycatcher and Redstart showed a preference for mature woodland. Lesser Spotted Woodpecker, Marsh Tit, Wood Warbler, Garden Warbler, Nightingale and Pied Flycatcher prefer broadleaved woodland while other species used both broadleaved, conifer and mixed woodland. Lesser Redpoll showed a preference for Spruce dominated plantations.

Breeding habitat at the woodland scale is most likely to be influenced by management which results in a mosaic of woodland blocks at different age/developmental stages. Species most influenced are species with associations with young woodlands stages or coppice; Nightjar, Willow Warbler, Garden Warbler, Nightingale and Tree Pipit. Predominantly coniferous woodlands are likely to result in most species being absent or scarce except for Nightjar, Tree Pipit and Lesser Redpoll. Mechanisms of the landscape effect of deer on breeding habitats of target birds are unknown with only correlative evidence available, although this clearly links declines of species in regions where deer densities have increased over the past few decades, mainly in south east and central England.

Post-fledging requirements Very little information is available on post-fledging requirements of the target species. This paucity of information makes inferring requirements in relation to forest management and deer browsing very difficult. Nightingale is the only species shown to be affected, with a requirement for taller vegetation to provide cover from predators to nestlings which fledge several days before capable of flight. The close proximity of a variety of structures and types of vegetation required by Nightingale may be affected by management methods that remove any one of these. No effects of deer on post-fledging requirements can be inferred from the literature found.

Seasonal differences for resident species Of the eight resident species few seasonal differences in habitat preference are reported. Dunnock move to lower elevations in the winter, as do some Lesser Redpoll from upland conifer habitats. Bullfinch show seasonal differences as they are wider ranging, tracking seasonal food sources and so are not strongly territorial. No forest management or deer related factors were found to influence seasonal habitat relationships. Back to contents

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Species nesting on or near the ground

Nightjar Caprimulgus europaeus

1. Nest site selection/requirements Nightjars nest on the ground, with eggs laid on bare ground or moss without a scrape, although very occasionally slightly above ground level on dead vegetation or a tree stump (Wiseman, 2006, Cross et al., 2006, Cresswell and Alexander, 1992). Nest sites are usually situated on flat ground, either in flat depressions or elevated plateau, and apparently suitable habitat situated on slopes is avoided (Cross et al., 2006). Nest sites are situated within small open spaces, in one study averaging 1m2 (Burgess et al., 1990), or larger clearings within or adjacent to coniferous plantations(Cross et al., 2006, Alexander and Cresswell, 1990) and are often associated with a single nearby tree which serves as a male song post (Burgess et al., 1990). Nest sites may be associated with ericaceous vegetation, and in some studies with sparse bracken cover (Burgess et al., 1990, Cross et al., 2006, Berry, 1979). In England, 15% of nests located by radio tracking adult females were in plantations containing trees >5 m high (Cresswell, 1996). Although disturbance at the nest by humans and dogs reduces nest success there does not seem to be any avoidance of disturbance as nests were not located further away from tracks, with the distance from nests to nearest track being highly variable, on average 60.9 m (Dolman, 2011, Langston et al., 2007a). High vegetation around the nest increased the chances of birds being flushed from the nest because the sitting adult cannot as readily identify the source of disturbance (Langston et al., 2007a).

2. Territory selection/attributes Territory size is difficult to identify but two studies give average territory size of 3.6 and 13 ha (Bult, 2002, Stasiak et al., 2013). Nightjars exhibit some site fidelity although little detail is given in the studies found (Cresswell, 1993).

Territories are situated in areas containing open woodland or heathland, particularly early successional birch and Scots pine or young coniferous plantations (Gribble, 1983, Jensen and Jacobsen, 1996, Conway et al., 2007, Morris et al., 1994, Stasiak et al., 2013). Home ranges of three radio tracked birds in Switzerland included 20-50% forest cover (Sierro et al., 2001) and 57% of calling males in an English study were located in planted woodland (Langston et al., 2007b). Territories encompass open space, including open space within woodlands, which could consist of heathland, new or restocked conifer plantation, wide tracks within mature woodlands or open space resulting from peatcutting, clearfelling, windfall, burning or grazing, and in England Nightjar also occur in chestnut coppice (Blueml, 2004, Conway et al., 2007, Verstraeten et al., 2011, Stasiak et al., 2013). The minimum size of open space within territories has been quantified in many studies but results are highly variable. Studies find territories require a minimum open area of between 2-10 ha (Ravenscroft, 1989, Cross et al., 2006, Jensen, 2010) with occurrence more likely where open spaces are larger or where more open patches occur (Scott et al., 1998, Bright et al., 2007, Stasiak et al., 2013). The minimum heathland patch size required for more than one territory in southern England was 1.5 ha (Bright et al., 2007) and

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in Wales across four study sites mean area of open space was 23 ha (Cross et al., 2006). Plantations are preferred if <10 years old and are unlikely to be used if older than 15 years old, although some studies find territories situated in much older plantations where the crop has largely failed and widely spaced trees (Bowden and Green, 1994, Scott et al., 1998, Ravenscroft, 1989). In Denmark 68% of territories were located in new clearfells or restocked plantations (Jensen, 2010), in Holland most were situated in young plantations (Bult, 2002) and in France 58% were in wet heathland, 29% in dry heathland or sandy areas and 13% in forests, mainly pine forests (Blueml, 2004). Territories containing open spaces of a circular or square shape seem preferred (Wichmann, 2004, Ravenscroft, 1989). It is suggested that open spaces of a more irregular shape develop scrub and denser ground flora more rapidly, at the expense of bare ground, and so are suitable to Nightjars for fewer years.

Occupancy is positively related to areas with ground cover comprising vegetation with little height or vertical structure (fine debris, moss and short grasses) but also with areas of longer grasses, bracken and shrubs which relates to plantation age (Bowden and Green, 1994). Similarly in a Polish forest, territories were found to have lower densities of field layer vegetation than random areas and were dominated by moss, lichens and grass (Stasiak et al., 2013). The same English study found territories were more common where soil Ph was higher although the opposite was found in another English study (Scott et al., 1998).

Availability of song perches within territories is important, and these consist of a tree or several scattered trees, one study finding Nightjar prefer trees where the lower crown edge is relatively high and use a dead branch below the canopy to sing (‘churr’) from (Wichmann, 2004). Another study demonstrated that population size can be increased by encouraging these conditions through management, along with other relevant woodland, open space and bare ground management (Burgess et al., 1990).

3. Main food items Diet is dominated by adult moths during the breeding season (Cleere, 1999, Sierro et al., 2001, Barbaro and Battisti, 2011, Bowden and Green, 1994). Faecal samples from adult Nightjars in England found 79% of items were moths with Diptera, elaterid beetles, Curculioniods and chafer imago also found (Bowden and Green, 1994). In a Swiss study of adult Nightjar, adult moths constituted 81-93% of prey mass, with 5- 18.7% Coleoptera and <1% Diptera and Hymenoptera, while nestlings were fed a similar proportion of moths but 6.5% Neuroptera (Sierro et al., 2001). In this study prey choice reflected relative abundance of these taxa and found that size of prey increases seasonally during the breeding season although small items are taken throughout. In southern Europe, Nightjar feeds on and can be a significant predator of Pine processionary moth during summer emergence (Barbaro and Battisti, 2011).

4. Foraging methods Nightjars forage for flying invertebrates, by aerial hawking in open habitats including clearings, forest tracks and forest edges. Most foraging activity occurs around dusk

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and dawn with little activity between midnight and 3 am (Alexander and Cresswell, 1990, Cleere, 1999, Cross et al., 2006, Bowden and Green, 1994). Open spaces are utilised by foraging Nightjars to detect prey silhouettes against twilit or moonlit skies with prey approached from below (Jackson, 2003, Wichmann, 2004). Nightjars also forage by moth-catching from a perch during a short flight lasting only a few seconds, a behaviour most prevalent in warmer temperatures and outside the time spent around dusk and dawn when foraging by sustained flight is preferred (Jackson, 2003, Cresswell, 1996). Deciduous or mixed woodlands including wooded gardens and orchards are preferred foraging habitat (Sierro et al., 2001, Alexander and Cresswell, 1990, Cresswell, 1996) with a study in Switzerland recording 74% of foraging locations in oak scrubland, 15% in vineyards and 9% in pine forest. Dense pine forests are avoided as unsuitable for aerial hawking, even when prey abundance is higher (Sierro et al., 2001) and one English study found foraging Nightjars avoided conifer plantations and arable or improved grassland and foraged little in Calluna dominated heathland (Alexander and Cresswell, 1990). Bowden and Green (1994) found similar foraging habitat choice but found grassy heaths were strongly selected along with young coniferous plantations (particularly those 3-5 years old). Another English study found preference for deciduous woodland including orchards, but they also foraged at saltmarsh and wetland habitats, particularly wet meadows, and rarely on heathland (Cresswell, 1996).

Nightjars are efficient predators, observed catching up to 18–19 prey items per minute and to capture more than one insect at a time on African non-breeding grounds (Jackson, 2003). Nestlings were typically provisioned by adults every 10-15 minutes in a Welsh study, although adults sometimes spend up to 6 hours away from territories foraging (Cross et al., 2006). Many radio tracking studies also find foraging in areas away from nest sites, up to 7 km distant and 3.1 km on average, suggesting they travel to exploit a few rich feeding areas over consecutive days (Alexander and Cresswell, 1990, Cresswell, 1996), although in other studies foraging was mainly within 1 km of territory centres (Bowden and Green, 1994).

5. Breeding habitat Density of singing males is highly variable and influenced by habitat. Densities are higher in forest clearings and young plantations than in heathland (Ravenscroft, 1989), and in heathland situated within 500m of woodland (Liley and Clarke, 2003). Density is lower in the vicinity of housing and urban development (Liley and Clarke, 2003, Langston et al., 2007a). Highest densities found were up to 16 territories/km2 in Welsh heathland (Cross et al., 2006), 15-16 territories /km2 in German heathland (Tomasini and Kneis, 2004) and 10-14 territories /km2 in Dutch young plantations (Bult, 2002). Lowest densities found were 0.88 territories/ km2 in Poland (Stasiak et al., 2013), 1.3 territories /km2 (range 0.9-2.1) in Denmark (Jensen, 2010), 2.6 territories /km2 at the landscape scale in German dry pine/heathland (Tomasini and Kneis, 2004), 5.25 territories /km2 in Wales (Cross et al., 2006) and 9.8 territories /km2 in southern England (Bright et al., 2007).

Most Nightjars in southeast and southwest England occur on dry heathland, but elsewhere in the UK most occur in coniferous forests (Conway et al., 2007). Wet

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heathland is also utilised (Blueml, 2004, Conway et al., 2007) although occurrence in wet heathland was positively influenced by proximity to a mosaic of wooded and open habitats (Blueml, 2004). In Italy Nightjar presence is positively associated with forest edges and shrubs (Bartolommei et al., 2013). Habitat selection was more based on presence of open space and proximity to woodland than on species composition. Distribution can be affected by habitat fragmentation with occurrence more likely with increasing area and number of patches of heathland in an area of southern England (Bright et al.).

6. Post fledging behaviour/requirements Second nesting attempts are common, potentially with a new mate (Cresswell and Alexander, 1992). Young become independent about two weeks post fledging (Schuchmann, 1999). No other information was found on post fledging behaviour or requirements.

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Key habitat requirement for Nightjar are areas of open space for centring territories on, in association with woodlands and woodland edges for foraging, and isolated trees for territorial behaviours. These conditions may be spatially separated by up to at least 6 km. Nightjars are abundant in large scale harvesting systems such as clearfelling, and are also abundant in new and restocked coniferous plantations. Presence becomes less likely once post management growth stages become too dense and scrubby when the availability of open spaces diminishes. In plantations, occupancy is much reduced after 10-15 years post planting or felling. Shape of open spaces may influence suitability with irregular shaped open spaces becoming dense and scrubby quicker than circular or square shaped spaces. Rotational clearfell forests will typically provide a favourable continuity of new clearfell and restocked areas, with approximately 30-40% of habitat suitable at any one time (Langston et al., 2007b). There is some evidence that chemical weed treatment positively influences nightjar density, particularly from increased chemical treatment in older (6-10 year) plantations, but not from mechanical weed control (Bowden and Green, 1994).

High deer browsing may be beneficial to Nightjar populations because of their requirement for open space and bare ground for both nesting and foraging. Nightjars are predominantly aerial feeders so deer browsing is unlikely to have an effect of foraging habitat.

9. References Alexander, I. & Cresswell, B. 1990. Foraging by Nightjars Caprimulgus europaeus away from their nesting areas. Ibis 132: 568-574. Barbaro, L. & Battisti, A. 2011. Birds as predators of the pine processionary moth (Lepidoptera: Notodontidae). Biological Control 56: 107-114.

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Bartolommei, P., Mortelliti, A., Pezzo, F. & Puglisi, L. 2013. Distribution of nocturnal birds (Strigiformes and Caprimulgidae) in relation to land-use types, extent and configuration in agricultural landscapes of Central Italy. Rendiconti Lincei-Scienze Fisiche E Naturali 24: 13-21. Berry, R. 1979. Nightjar habitats and breeding in East Anglia. British Birds 72: 207- 218. Blueml, V. 2004. Distribution, population size and habitat preference of the European nightjar (Caprimulgus europaeus) in Lower Saxony 2003: results of a statewide census. Vogelkundliche Berichte aus Niedersachsen 36: 131-162. Bowden, C.G.R. & Green, R.E. 1994. The ecology of nightjars on pine plantations in Thetford Forest. RSPB. Bright, J., Langston, R.H.W., Bierman, S. & Pearce-Higgins. The effect of habitat fragmentation on distribution of nightjars Caprimulgus europaeus breeding on heathland in England. RSPB. Bright, J.A., Langston, R.H.W. & Bierman, S. 2007. Habitat associations of nightjar Caprimulgus europaeus breeding on heathland in England. RSPB Research Report, 25. Bult, H. 2002. Nightjars Caprimulgus europaeus in the SW Netherlands. Limosa 75: 91-102. Burgess, N.D., Evans, C.E. & Sorensen, J. 1990. The management of lowland heath for Nightjars at Minsmere, Suffolk, Great Britain. Journal of Environmental Management 31: 351-359. Cleere, N. 1999. Family caprimulgidae (Nightjars). In Handbook of Birds of the World Volume 5. (Eds. J. Del Hoyo, A. Elliott & J. Sargatal), Vol. 5, pp. 302-386. Barcelona: Lynx Edicions. Conway, G., Wotton, S., Henderson, I., Langston, R., Drewitt, A. & Currie, F. 2007. Status and distribution of European Nightjars Caprimuigus europaeus in the UK in 2004. Bird Study 54: 98-111. Cresswell, B. 1993. Site and habitat preferences of nesting and roosting nightjars. In Stour Ringing Group Annual Report. Vol. 1992, pp. 33-35. Stour Ringing Group. Cresswell, B. 1996. Nightjars - some aspects of their behaviour and conservation. British Wildlife 7: 297-304. Cresswell, B. & Alexander, I. 1992. Activity patterns of foraging nightjars (Caprimulgus europaeus). In Wildlife telemetry: remote monitoring and tracking of animals. (Eds. I. Priede & S. Swift), pp. 642-647. California: University of California. Cross, T., Lewis, J., Morgan, C. & Rees, D. 2006. European nightjar (Caprimulgus europaeus) breeding success and foraging behaviour in upland coniferous forests in mid-Wales. Brecknock Wildlife Trust/Forestry Commision. Dolman, P.M. 2011. Woodlark and Nightjar recreational disturbance and nest predator study 2008 and 2009. Norwich: University of East Anglia. Gribble, F.C. 1983. Nightjars in Britain and Ireland in 1981. Bird Study 30: 165-176. Jackson, H.D. 2003. A review of foraging and feeding behaviour, and associated anatomical adaptations, in Afrotropical nightjars. Ostrich 74: 187-204. Jensen, N.O. 2010. The population of nightjar in NW Jutland 1994-2007 and its future prospects. Dansk Ornitologisk Forenings Tidsskrift 104: 12-21.

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Jensen, N.O. & Jacobsen, L.B. 1996. The breeding population of nightjar Caprimulgus europaeus in Denmark, 1992-95. Dansk Ornitologisk Forenings Tidsskrift 90: 93- 98. Langston, R.H.W., Liley, D., Murison, G., Woodfield, E. & Clarke, R.T. 2007a. What effects do walkers and dogs have on the distribution and productivity of breeding European Nightjar Caprimulgus europaeus? Ibis 149: 27-36. Langston, R.H.W., Wotton, S.R., Conway, G.J., Wright, L.J., Mallord, J.W., Currie, F.A., Drewitt, A.L., Grice, P.V., Hoccom, D.G. & Symes, N. 2007b. Nightjar Caprimulgus europaeus and Woodlark Lullula arborea - Recovering species in Britain? Ibis 149: 250-260. Liley, D. & Clarke, R.T. 2003. The impact of urban development and human disturbance on the numbers of nightjar Caprimulgus europaeus on heathlands in Dorset, England. Biological Conservation 114: 219-230. Morris, A., Burges, D., Fuller, R.J., Evans, A.D. & Smith, K.W. 1994. The status and distribution of Nightjars Caprimulgus europaeus in Britain in 1992 - A report to the British Trust for Ornithology. Bird Study 41: 181-191. Ravenscroft, N.O.M. 1989. The status and habitat of the Nightjar Caprimulgus europaeus in coastal Suffolk. Bird Study 36: 161-169. Schuchmann, K. 1999. European nightjar. In Handbook of Birds of the World. (Eds. J. del Hoyo, A. Elliott & J. Sargatal), Vol. 5, pp. 653. Barcelona: Lynx Edicions. Scott, G.W., Jardine, D.C., Hills, G. & Sweeney, B. 1998. Changes in Nightjar Caprimulgus europaeus populations in upland forests in Yorkshire. Bird Study 45: 219-225. Sierro, A., Arlettaz, R., Naef-Daenzer, B., Strebel, S. & Zbinden, N. 2001. Habitat use and foraging ecology of the nightjar (Caprimulgus europaeus) in the Swiss Alps: towards a conservation scheme. Biological Conservation 98: 325-331. Stasiak, K., Grzywaczewski, G., Gustaw, W. & Cios, S. 2013. Effect of the forest structure on the number and territory size of nightjar Caprimulgus europaeus. Sylwan 157: 306-312. Tomasini, J. & Kneis, P. 2004. Habitat structure and breeding density of the nightjar (Caprimulgus europaeus) in the Gohrischheide Zeithain-Altenau, northern Saxony/southern Brandenburg. Acta Ornithoecologica 5: 3-13. Verstraeten, G., Baeten, L. & Verheyen, K. 2011. Habitat preferences of European Nightjars Caprimulgus europaeus in forests on sandy soils. Bird Study 58: 120- 129. Wichmann, G. 2004. Habitat use of nightjar (Caprimulgus europaeus) in an Austrian pine forest. Journal of Ornithology 145: 69-73. Wiseman, E.J. 2006. European nightjar nesting on tree stump. British Birds 99: 267.

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Tree Pipit Anthus trivialis

1. Nest site selection/requirements Tree Pipit nest on the ground with nests mostly hidden in grass tussocks (van Hecke, 1979, Loske, 1987, Burton, 2006, Burton, 2009). The nest entrance tends to face in an easterly direction (van Hecke, 1979, Loske, 1987, Burton, 2006), with most early nests facing south of east and later nests facing north of east (Burton, 2006). This orientation provides shade from sun and reduced exposure to prevailing wind direction. The type of vegetation that nests are located in can also change between early and late nests as vegetation structure changes with growth during the season. In one study early nests were in marsh vegetation and later ones in grassy banks (Meury, 1989) while in another early nests were in Purple moor grass and later ones in ericaceous species (van Hecke, 1979).

2. Territory selection/attributes Mean territory size is 1.2-1.5 ha, ranging from 0.25-4.0 ha (Meury, 1989, Burton, 2007, Kumstatova et al., 2004). Low pairing success is associated with territories of <1 ha in the Brecks (Burton, 2007). Territory size can vary between years and with breeding densities; smaller territories are defended at higher breeding densities (van Hecke, 1979, Meury, 1989, Burton, 2007). Trees for song perches are required within territories (Meury, 1989, Kouki et al., 1992, Burton, 2007, Moga et al., 2009) and in areas lacking trees but otherwise containing suitable habitat banks, wires and shrubs are used (Szczur, 2000, Kumstatova et al., 2004). Tree cover varies greatly between studies and territories; in most cases treeless areas and areas with tree cover >60% are avoided (Loske, 1987, Kouki et al., 1992, Moga et al., 2009). In denser forest, territories are at higher densities along boundaries between forest and open habitats but there are no such edge effects in more open wooded habitats (Huebner, 2009, Moga et al., 2009).

3. Main food items Nestling diet is entirely invertebrates with caterpillars being the main food items early in the breeding season and crickets and grasshoppers later in the season, with other invertebrates forming a significant part of the diet including spiders, flies, aphids and leafhoppers (Meury, 1991, Krištín and Exnerová, 1994).

4. Foraging methods Foraging is mainly on the ground, with some tree foraging, but rarely in shrubs (Ulfstrand, 1976, Forbes-Watson, 1983, Kouki et al., 1992, Hogstad, 2005). Both food availability and habitat structure affect where Tree Pipits forage; during November moth outbreaks in Norway there was increased foraging in birch trees (Hogstad, 2005). In Switzerland, the preferred feeding habitat was marshland which held highest invertebrate densities, and they only started foraging on grass fields when marshland vegetation became too tall and dense (Meury, 1991).

5. Breeding habitat In the UK, Tree Pipits are mainly associated with newly harvested and young regrowth of conifer plantations and chestnut coppice, heathland, and open

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woodland of upland margins (Currie and Bamford, 1981, Bibby et al., 1985, Fuller and Moreton, 1987, Szczur, 2000, Fuller et al., 2006, Burton, 2007, Hewson et al., 2011). They also occur in semi natural and open pine forests (Summers, 2007, Taylor and Summers, 2009). Within conifer plantations and coppice, the highest breeding densities are found in 1-5 year old regrowth and they are also found at lower densities in newly harvested areas and in older plantations up to 11 years old (Currie and Bamford, 1981, Fuller and Moreton, 1987, Baguette et al., 1994, Burton, 2007). The optimal habitat in conifer plantations is 2-5 year old regrowth with Tree Pipits preferentially occupying this age class first in spring and having higher nest survival compared to older and younger plantations (Burton, 2009).

On continental Europe they also occupy a similarly wide range of broad habitats including peatlands, grasslands, marshland, heathland, broadleaf and conifer forest (van Hecke, 1979, Helle, 1985, Kouki et al., 1992, Baguette et al., 1994, Gjerde and Saetersdal, 1997, Glutz von Blotzheim, 2000, Paquet et al., 2006, Huebner, 2009).

Most habitats occupied have low density trees or are open woodland; dense forest with >80% tree cover is avoided (Loske, 1987, Smith et al., 1987, Szczur, 2000, Smart et al., 2007, Moga et al., 2009, Edenius, 2011). Habitat used by Tree Pipits includes a wide range of ground vegetation types with different structures including areas of high vegetation cover, sparse cover, dry grasslands, moist grasslands, heathlands, Bracken and marshland (van Hecke, 1979, Currie and Bamford, 1981, Glutz von Blotzheim, 2000, Szczur, 2000, Fuller et al., 2006). A common factor of habitats used is they are of low intensity management with varied structure and high invertebrate abundance (Meury, 1989, Meury, 1991, Glutz von Blotzheim, 2000). Low shrub cover is also important Tree Pipit habitat (Bibby et al., 1985, Loske, 1987, Müller, 2005, Smart et al., 2007, Moga et al., 2009).

6. Post fledging behaviour/requirements Young Tree Pipits move up to 600 m from nests within the first week of the post- fledging period (van Hecke, 1979).

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Tree Pipits select habitat with low shrub cover and with tree canopy cover of <60%. Although Tree Pipits both nest and feed on the ground, they will use a wide range of ground vegetation types as long as they provide dense cover for nests and high invertebrate abundance. Tree Pipits also rely on low density of trees for song perches. Within woodlands, Tree Pipit habitat is temporary created by clearfell and coppicing, particularly on sandy soils or in the uplands. The retention of some standing trees on cleared coupes is important for Tree Pipits to be able to make use of the immediate post felling stages until young trees reach a height suitable for song perches (2 m).

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Deer browsing is unlikely to negatively affect Tree Pipit habitat and it has been suggested may improve Tree Pipit habitat through creating a grassier field layer, although no evidence was found to support for this.

9. References Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Burton, N.H.K. 2006. Nest orientation and hatching success in the tree pipit Anthus trivialis. Journal of Avian Biology 37: 312-317. Burton, N.H.K. 2007. Influences of restock age and habitat patchiness on Tree Pipits Anthus trivialis breeding in Breckland pine plantations. Ibis 149: 193-204. Burton, N.H.K. 2009. Reproductive success of Tree Pipits Anthus trivialis in relation to habitat selection in conifer plantations. Ibis 151: 361-372. Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Edenius, L. 2011. Short-term effects of wildfire on bird assemblages in old pine- and spruce-dominated forests in northern Sweden. Ornis Fennica 88: 71-79. Forbes-Watson, A.D. 1983. Arboreal leaf-gleaning by Tree pipit. British Birds 76: 535- 535. Fuller, R.J., Atkinson, P.W., Garnett, M.C., Conway, G.J., Bibby, C.J. & Johnstone, I.G. 2006. Breeding bird communities in the upland margins (ffridd) of Wales in the mid-1980s. Bird Study 53: 177-186. Fuller, R.J. & Moreton, B.D. 1987. Breeding bird populations of Kentish sweet chestnut (Castanea sativa) coppice in relation to age and structure of the coppice. Journal of Applied Ecology 24: 13-27. Gjerde, I. & Saetersdal, M. 1997. Effects on avian diversity of introducing spruce Picea spp. plantations in the native pine Pinus sylvestris forests of western Norway. Biological Conservation 79: 241-250. Glutz von Blotzheim, U.N. 2000. Impact of eutrophication on distribution and demography of water pipit Anthus spinoletta and tree pipit Anthus trivialis in the prealps of central Switzerland. Ornithologische Beobachter 97: 343-347. Helle, P. 1985. Habitat selection of breeding birds in relation to forest succession in northeastern Finland. Ornis Fennica 62: 113-123. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Hogstad, O. 2005. Numerical and functional responses of breeding passerine species to mass occurrence of geometrid caterpillars in a subalpine birch forest: a 30- year study. Ibis 147: 77-91. Huebner, A. 2009. Die Habitatwahl des Baumpiepers Anthus trivialis – eine Analyse mittels GIS. Vogelwarte 47: 165-170. Kouki, J., Niemi, G.J. & Rajasarkka, A. 1992. Habitat associations of breeding peatland passerine species in eastern Finland. Ornis Fennica 69: 126-140.

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Krištín, A. & Exnerová, A. 1994. On the diet and breeding biology of Tree Pipit (Anthus trivialis) and Black Redstart (Phoenicurus ochruros). Sylvia 30: 64-71. Kumstatova, T., Brinke, T., Tomkova, S., Fuchs, R. & Petrusek, A. 2004. Habitat preferences of tree pipit (Anthus trivialis) and meadow pipit (A-pratensis) at sympatric and allopatric localities. Journal of Ornithology 145: 334-342. Loske, K.H. 1987. On the habitat selection of the tree pipit (Anthus trivialis). Journal Fur Ornithologie 128: 33-47. Meury, R. 1989. Siedlungsdichte und Raumnutzung des Baumpiepers Anthus trivialis im inselartig verteilten Habitat des aargauischen Reusstals. Ornithologische Beobachter 86: 105-135. Meury, R. 1991. Zur nahrungsokologie des Baumpiepers Anthus trivialis in einem landwirtschaftsgebiet des Schweizeriscen Mittellandes. Ornithologische Beobachter 88: 57-72. Moga, C.I., Hartel, T. & Ollerer, K. 2009. Ancient oak wood-pasture as a habitat for the endangered tree pipit Anthus trivialis. Biologia 64: 1011-1015. Müller, J. 2005. Bird communities as indicators for woodland structures in oak woods. Ornithologische Beobachter 102: 15-32. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Smart, J., Taylor, E., Amar, A., Smith, K., Bierman, S., Carpenter, J., Grice, P., Currie, F. & Hewson, C. 2007. Habitat associations of woodland birds: implications for woodland management for declining species. In RSPB Research Report No., Vol. 26. Sandy: RSPB. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Summers, R.W. 2007. Stand selection by birds in Scots pinewoods in Scotland: the need for more old-growth pinewood. Ibis, 149 (suppl 2): 175-182. Szczur, J. 2000. The 1999 Nottinghamshire Tree pipit survey. The Birds of Nottinghamshire 1999: 145-153. Taylor, S.D. & Summers, R.W. 2009. Breeding numbers and stand type preferences of Redstarts Phoenicurus phoenicurus and Tree Pipits Anthus trivialis in a Scots Pine Pinus sylvestris wood. Bird Study 56: 120-126. Ulfstrand, S. 1976. Feeding niches of some passerine birds in a south Swedish coniferous plantation in winter and summer. Ornis Scandinavica 7: 21-27. van Hecke, P. 1979. Verhalten, Nest und Neststandort des Baumpiepers (Anthus t. trivialis). Journal Fur Ornithologie 120: 265-279.

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Final report WC0793/CR0485 (29.04.14)

Wood Warbler Phylloscopus sibilatrix

1. Nest site selection/requirements Wood Warblers nest on the ground, most often under tussocks of grass or sedges within the herb layer or leaf litter and also under tree roots and boulders (Grendelmeier, 2011, Lippek, 2009). Nest sites are associated with more open understory, and a plant, shrub or small tree <50 cm high which provides a perch close to the nest (Grendelmeier, 2011, Mallord et al., 2012). Nests are often situated on slopes (Mallord et al., 2012, Goodfellow, 1987). Nest predation is influenced to some extent by vegetation and nest concealment, with lower predation of nests concealed by vegetation at the micro-scale (Grendelmeier, 2011).

2. Territory selection/attributes Territory size is up to 3.6 ha but average size is smaller (0.5 ha) in areas of high density (Clement, 2006). In continental studies adult Wood Warblers exhibit variable but typically low site fidelity, with 0-16% of adults returning to previous breeding sites across 10 studies (Wesolowski et al., 2009). Site fidelity is higher in the UK, 28%, (Norman, 1994) and in Germany 96% returned within 40 km of original territories of which 56% returned within 1.7 km (Lippek, 2009). Between year and within year breeding dispersal movements of >100 km were reported (Herremans, 1993, Lippek, 2009). Territory occupation was influenced by abundance of rodents in Poland, with lower occupation of woodlands in years with high rodent abundance (Wesolowski et al., 2009).

Territories are situated within parts of woodlands with high proportions of canopy cover (Fuller, 2000, Delahaye and Vandevyvre, 2008, Mallord et al., 2012, Marti, 2007). Studies in Belgium, Switzerland and Finland indicate that intra-specific attraction influences territory location, with territory clustering (Herremans, 1993, Tiainen et al., 1983, Grendelmeier, 2011).

3. Main food items Diet comprises predominantly small invertebrates from a wide range of taxa. Diet fed to nestlings is predominantly caterpillars and winged insects, and also spiders, although the relative proportions of these varies between studies and changes seasonally and with chick age (Maziarz and Wesolowski, 2010). Winged insects are frequently fed to nestlings, 63% of food items in Germany (Lippek, 1996), 23% in Poland (Maziarz and Wesolowski, 2010) and 17% in Sweden (Temrin et al., 1997). Panorpa and Tipulidae were the most frequently fed winged insects (Maziarz and Wesolowski, 2010, Lippek, 1996). Caterpillars constituted 34% of nestling diet in Germany (Lippek, 1996), 45% in Poland (Maziarz and Wesolowski, 2010) and 53% in Sweden (Temrin et al., 1997). Spiders constituted 23% of nestling diet in Poland and were provided in greater proportions to younger nestlings (Maziarz and Wesolowski, 2010). Diet includes fruit such as Bramble and Elder in autumn (Clement, 2006).

4. Foraging methods Wood Warblers forage for invertebrates mainly in the forest canopy and undergrowth by actively inspecting leaves and branches, but also hover and make

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sallies after flying invertebrates (Clement, 2006). They usually forage alone, or in pairs, except on passage when they may also forage in small or mixed species flocks (Clement, 2006).

5. Breeding habitat In Britain Wood Warblers preferentially choose oak woodland (Mallord et al., 2012, Bellamy, 1986) but are also found in a wide variety of broadleaved, coniferous and mixed woodland types throughout their range (Herremans, 1993, Wesolowski and Maziarz, 2009, Delahaye and Vandevyvre, 2008, Bellamy, 1986, Bibby, 1989, Currie and Bamford, 1981, Currie and Bamford, 1982, Nilsson, 1979, Haapenen, 1966). Habitat selection is more based on vegetation structure than species composition, with a structural profile including high canopy cover and density, low understorey density containing a scattering of perches provided by low branches or thin shrubs and a low but moderate density of ground cover (Mallord et al., 2012, Delahaye and Vandevyvre, 2008, Bellamy, 1986, Cody, 1978). Wood Warbler density in Wales was positively associated with slope and negatively associated with subcanopy cover 0.5- 2 m high (Mallord et al., 2012, Bellamy, 1986).

6. Post fledging behaviour/requirements Young are fed by parents for 2-3 days after fledging and may remain in family groups for up to 4 weeks (Clement, 2006). No other information was found on post fledging behaviour or requirements.

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Key habitat requirements for Wood Warbler are mature closed canopy woodland with a relatively open understory. Wood Warblers are more abundant in small scale harvesting systems such as group fell and selective felling although presence is unlikely in areas with a dense or shrubby understory such as early growth stages post management. A few studies suggest that foliage feeders are negatively affected by thinning management which may affect Wood Warbler (Virkkala, 1987, de la Montaña et al., 2006).

High deer browsing may be beneficial to Wood Warbler populations because of their open understory nesting requirements. Wood Warblers feed primarily in the canopy so deer browsing is unlikely to have an effect on foraging habitat. No nest trampling by deer was recorded in a detailed nest predator study (Mallord et al, 2012).

9. References Bellamy, P.E. 1986. Habitat preferences of Wood warblers in North Wales. Vol. MSc, pp. 68. Bangor: University of Wales. Bibby, C.J. 1989. A survey of breeding Wood warblers Phylloscopus sibilatrix in Britain, 1984-1985. Bird Study 36: 56-72. Clement, P. 2006. Wood warbler. In Handbook of Birds of the World. (Eds. J. del Hoyo, A. Elliott & Christie), Vol. 11, pp. 653. Barcelona: Lynx Edicions.

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Cody, M.L. 1978. Habitat selection and interspecific territoriality among the Sylviid warblers of England and Sweden. Ecological Monographs 48: 351-396. Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Currie, F.A. & Bamford, R. 1982. The value to birdlife of retaining small conifer stands beyond normal felling age within forests. Quarterly Journal of Forestry 76: 153- 159. de la Montaña, E., Rey-Benayas, J.M. & Carrascal, L.M. 2006. Response of bird communities to silvicultural thinning of Mediterranean maquis. Journal of Applied Ecology 43: 651-659. Delahaye, L. & Vandevyvre, X. 2008. Is the wood warbler an indicator species for the quality of Ardennes deciduous forest? Aves (Liege) 45: 3-14. Fuller, R.J. 2000. Influence of treefall gaps on distributions of breeding birds within interior old-growth stands in Bialowieza Forest, Poland. The Condor: 267-274. Goodfellow, P. 1987. Wood Warblers - Sixty years ago. Devon Birds 40: 27-32. Grendelmeier, A. 2011. The enigmatic decline of the Wood Warbler Phylloscopus sibilatrix: nest predation and habitat characteristics. Vol. MSc. Haapenen, A. 1966. Bird fauna of in Finnish forests in relation to forest succession II. Annales Zoologici Fennici 3: 176-200. Herremans, M. 1993. Clustering of territories in the Wood Warbler Phylloscopus sibilatrix. Bird Study 40: 12-23. Lippek, W. 1996. Prey availability and nestling diet of wood warblers Phylloscopus sibilatrix in forests of eastern Northrhine-Westphalia, Germany. Vogelwelt 117: 29-38. Lippek, W. 2009. Notes on breeding biology and site fidelity of Wood Warbler Phylloscopus sibilatrix in Westphalia-Lippe (Germany). Vogelwelt 130: 165-174. Mallord, J.W., Charman, E.C., Cristinacce, A. & Orsman, C.J. 2012. Habitat associations of Wood Warblers Phylloscopus sibilatrix breeding in Welsh oakwoods. Bird Study 59: 403-415. Marti, J. 2007. Habitat characteristics of the wood warbler Phylloscopus sibilatrix territories in the canton of Glarus, Switzerland. Ornithologische Beobachter 104: 45-52. Maziarz, M. & Wesolowski, T. 2010. Timing of breeding and nestling diet of Wood Warbler Phylloscopus sibilatrix in relation to changing food supply. Bird Study 57: 540-552. Nilsson, S.G. 1979. Density and species richnessof some forest bird communities in Southern Sweden. Oikos 33: 392-401. Norman, D. 1994. The return rate of adult male wood warblers Phylloscopus sibilatrix to a peripheral breeding area. Ringing and Migration 15: 79-83. Temrin, H., Brodin, A., Åkerström, O. & Stenius, S. 1997. Parental investment in monogamous pairs of Wood Warblers (Phylloscopus sibilatrix). Journal of Ornithology 138: 93-101. Tiainen, J., Vickholm, M., Pakkala, T., Piiroinen, J. & Virolainen, E. 1983. The habitat and spatial relations of breeding Phylloscopus warblers and the goldcrest Regulus regulus in southern Finland. In Annales Zoologici Fennici. Vol. 20, pp. 1-12. Virkkala, R. 1987. Effects of forest management on birds breeding in northern Finland. Annales Zoologici Fennici 24: 281-294.

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Wesolowski, T. & Maziarz, M. 2009. Changes in breeding phenology and performance of Wood Warblers Phylloscopus sibilatrix in a primeval forest: a thirty-year perspective. Acta Ornithologica 44: 69-80. Wesolowski, T., Rowinski, P. & Maziarz, M. 2009. Wood Warbler Phylloscopus sibilatrix: a nomadic insectivore in search of safe breeding grounds? Bird Study 56: 26-33.

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Final report WC0793/CR0485 (29.04.14)

Willow Warbler Phylloscopus trochilus

1. Nest site selection/requirements Nest on or near the ground in low vegetation, including the base of shrubs or in grass and herb vegetation (Cramp and Perrins, 1993, Fuller, 1995). The nest is a dome structure constructed primarily of grass and lined with feathers (Elts, 2005).

2. Territory selection/attributes The main attribute of Willow Warbler territories is canopy height and they are found in a wide variety of woodland types with a low canopy height (Cramp and Perrins, 1993, Fuller, 1995). Quantitative descriptions of canopy height range from 0.6-10 m with an optimal mean height of 5-7 m (Cody, 1978, Bellamy et al., 2009, Saether, 1983). Mean canopy height combines measures of maximum canopy height and canopy cover with Willow Warblers selecting areas of both low maximum canopy height and canopy cover, as well as a low cover of bare ground (Smart et al., 2007). Territories are found to be both positively and negatively associated with forest edges by different studies (Hansson, 1983, Tiainen et al., 1983, Kouki et al., 1992, Fuller, 1995, Patterson et al., 1995, Mason, 2001). Differences between studies are likely to relate to structural differences due to the type of forest and type of edge. High forest with eco-tonal edges have higher densities on edges, while forest containing gaps within it, or hard forest edges with arable farmland, have higher densities away from edges. Territories can also be strongly associated with the presence of birch trees (Mason, 1997). Song perches are near the tops of prominent trees, and in one study these were only in pines rather than in broadleaved trees (Kouki et al., 1992).

3. Main food items During the breeding season, Willow Warblers feed mainly on a wide variety of arthropods (Cramp and Perrins, 1993). Prey items are largely taken in proportion to availability from tree leaves, and include Hymenoptera, flies, aphids, mayflies and spiders (Arvidsson and Klaesson, 1986, Hogstad, 2005, Krupa, 2004). In years of geometrid moth outbreaks, particularly Epirrita spp., caterpillars of these are preferentially taken and constitute most of nestling diet (Arvidsson and Klaesson, 1986, Hogstad, 2005). After bud burst, male Willow Warblers shift from feeding on flying insects to aphids and caterpillars (Nystrom, 1991).

On migration Willow Warblers remain mainly insectivorous taking small and medium size beetles and flies but avoiding larger prey (Marchetti et al., 1998). They do not feed on fruit during autumn migration (Boddy, 1991). During spring migration and in early spring after arrival on breeding grounds they also feed on nectar and pollen from willow and poplar catkins (Nystrom, 1997, Johnson and Johnson, 2007).

4. Foraging methods Most prey is captured from leaves either by gleaning, from a perch or flying to pick off the leaf surface, but they rarely hover or catch insects in flight (Angell-Jacobsen, 1980, Nilsson and Ebenman, 1981, Nystrom, 1991, Forstmeier and Kessler, 2001, Bursky et al., 2004). Flycatching is primarily a behaviour shown by males when they

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first arrive on breeding territories before bud burst (Nystrom, 1991). Hovering is more frequent as a foraging method for females, which are lighter and shorter winged (Nystrom, 1991). In an experimental study, Willow Warblers were found to preferentially feed in trees subject to insect attack to the foliage, probably using leaf defence chemicals as the selection cue (Mantyla et al., 2004). In early spring before bud burst, Willow Warblers often forage by probing closed buds of trees and bushes (Amies, 1989). Foraging is mainly confined to tree crown foliage and Willow Warblers rarely forage on the ground or from shrubs (Ulfstrand, 1976, Angell- Jacobsen, 1980, Saether, 1983, Hogstad, 2005, Nystrom, 1991). Within tree crowns, foraging occurs on the outer twigs and leaves away from the trunk (Nilsson and Ebenman, 1981, Saether, 1983, Ulfstrand, 1976, Bursky et al., 2004). Willow Warblers can have a preference for foraging in the lower or mid crown of trees, with heights of foraging birds in the range 4-10 m (Cody, 1978, Angell-Jacobsen, 1980, Nilsson and Ebenman, 1981, Saether, 1983, Kouki et al., 1992, Hogstad, 2005).

A number of studies record preferences for foraging in particular tree species; however preferences are not consistent between studies. Preferred tree species for feeding are birch (Hogstad, 2005, Arvidsson and Klaesson, 1986), deciduous trees (Kouki et al., 1992, Nilsson and Ebenman, 1981) and pine (Ulfstrand, 1976). These differences may relate to geographical location and tree species available in study woodlands, and it is likely that foraging is concentrated on tree species with higher densities of insects in the foliage (Nystrom, 1997).

5. Breeding habitat Willow Warblers breed in a wide variety of woodland types including scrub, conifer, broadleaf and young forest (Moss et al., 1979, Smith et al., 1987, Fuller, 1995, Lavers et al., 2005). A UK wide study found occurrence most likely in woodland with mid- stages of understory growth, at open and closed thicket stage (Hewson et al., 2011). A number of studies record preferences for different types of forest. In Scandinavia, birch, deciduous and mixed forest are preferred over conifer forest (Tiainen et al., 1983, Haila et al., 1996, Hogstad, 1999).Within conifer woodlands pines are favoured over spruce (Gjerde and Saetersdal, 1997), and within broadleaved forest they are more frequent in oak than beech forests (Baguette et al., 1994). Breeding in conifer forest mostly occurs in young stages of plantation growth (Moss et al., 1979, Currie and Bamford, 1981, Baguette et al., 1994, Sweeney et al., 2010). In young conifer plantations abundance is higher in second rotation forests compared to first rotation (Sweeney et al., 2010, Currie and Bamford, 1981), where they prefer older pre- thicket stands at lower altitudes with a high density of trees including some broadleaves (Bibby et al., 1985). In broadleaved woodland, Willow Warblers also select early successional woodlands with low canopy height, low density of understory at 2-4 m, higher moss cover and more dead trees (Baguette et al., 1994, Smart et al., 2007). They also occur in older mature broadleaf stands where lower canopy cover allows the growth of a higher density of vegetation in the low shrub layer (Smith et al., 1987, Baguette et al., 1994).

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Proximity to roads and intensity of road traffic have been shown to reduce suitability of structurally suitable woodland habitat (Kuitunen et al., 1998, Reijnen et al., 1997, Reijnen and Foppen, 1994).

When breeding in patches of young growth within larger forests there is an indication that they may prefer larger areas, although the form of this relationship and the smallest patch size that is suitable is unknown (Paquet et al., 2006, Bellamy et al., 2009).

In fragmented habitats wood size may affect habitat occupancy, with greater occupation than expected in small woods (mean size occupied is 3.29 ha, (Hinsley et al., 1996) or greater abundance in small woods (Mason, 2001)). However in these studies all woodland sizes were relatively small with few > 10 ha in each study.

6. Post fledging behaviour/requirements Post-fledging dispersal distances increase with time since fledging. In the first 20 days juveniles were recorded at distances from 0-120 m from their nest, after 20-30 days they had moved 0-300 m and after 30 days 200-800 m. No young were retrapped after 33 days, having dispersed outside of the study area (Tiainen, 1983).

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Although Willow Warblers breed in a wide variety of woodland types with low canopy cover and canopy height, a key habitat in the UK is early successional woodland. Both clearfell and coppicing provide large areas of suitable habitat which has the highest Willow Warbler densities 3-10 years after harvesting or replanting. Smaller gaps of the type created by more selective felling regimes are less used than larger areas of young woodland (Bellamy et al., 2009, Paquet et al., 2006, Hinsley et al., 2009), therefore a change from clearfell and replanting to more continuous cover forestry is likely to reduce availability of suitable habitat. Within young woodland, the presence of non-crop species of trees is important for providing feeding habitat. The presence of birch is often associated with higher numbers of Willow Warblers, and in young upland conifer plantations they are more likely to be present where broadleaf species such as birch and willow are more abundant (see sections 2 and 5 above). Therefore cleaning operations to reduce weedy tree species from within the crop are likely to be detrimental to Willow Warbler populations.

At the UK scale Willow Warbler abundance is negatively affected by densities of Reeves’ muntjac (Newson et al., 2012). Deer selectively forage in young woodland and have a large impact on tree and shrub growth below 2 m. Therefore, high numbers of deer can lead to a reduction in habitat quality for Willow Warbler. This has been demonstrated for mixed coppice where higher numbers of Willow Warbler are found in areas protected from deer browsing (Holt et al., 2011, Gill and Fuller, 2007) and similar effects are likely in other woodland habitats. Deer browsing can change the composition and structure of the field layer where Willow Warblers nest,

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and therefore has the potential to affect nest site availability and nest predation vulnerability. However, Willow Warblers use a range of different nest sites, some of which are negatively affected by deer browsing (e.g. in Bramble) and others are positively affected (e.g. coarse grass tussocks), so predicting the effect of deer on nesting sites is not straightforward.

9. References Amies, P. 1989. Ivy berries and birds in spring. Bristol Ornithology 20: 183-184. Angell-Jacobsen, B. 1980. Overlap in Feeding Pattern between Willow Warbler Phylloscopus trochilus and Brambling Fringilla montifringilla in Two Forest Habitats in Western Norway. Ornis Scandinavica 11: 146-154. Arvidsson, B. & Klaesson, P. 1986. Territory size in a Willow warbler Phylloscopus trochilus population in mountain birch forest in Swedish Lapland. Ornis Scandinavica 17: 24-30. Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica, 15: 275-288. Bellamy, P.E., Hill, R.A., Rothery, P., Hinsley, S.A., Fuller, R.J. & Broughton, R.K. 2009. Willow Warbler Phylloscopus trochilus habitat in woods with different structure and management in southern England. Bird Study 56: 338-348. Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Boddy, M. 1991. Some aspects of frugivory by bird populations using coastal dune scrub in Lincolnshire. Bird Study 38: 188-199. Bursky, O.V., Kontorshchikov, V.V. & Batova, O.N. 2004. Foraging stereotypes of the willow warbler Phylloscopus trochilus and the chiffchaff P. collybita. Zoologicheskii Zhurnal 83: 839-850. Cody, M.L. 1978. Habitat selection and interspecific territoriality among the Sylviid warblers of England and Sweden. Ecological Monographs 48: 351-396. Cramp, S. & Perrins, C.M. (1993) Handbook of the birds of Europe, the Middle East and North Africa: The birds of the Western Palearctic. Vol. VI Warblers, Oxford: Oxford University Press. Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Elts, J. 2005. On the nest material of the willow warbler: a quantitative analysis. Hirundo 18: 31-33. Forstmeier, W. & Kessler, A. 2001. Morphology and foraging behaviour of Siberian Phylloscopus warblers. Journal of Avian Biology 32: 127-138. Fuller, R.J. (1995) Bird life of woodland and forest. , Cambridge: Cambridge University Press. Gill, R.M.A. & Fuller, R.J. 2007. The effects of deer browsing on woodland structure and songbirds in lowland Britain. Ibis 149: 119-127. Gjerde, I. & Saetersdal, M. 1997. Effects on avian diversity of introducing spruce Picea spp. plantations in the native pine Pinus sylvestris forests of western Norway. Biological Conservation 79: 241-250.

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Haila, Y., Nicholls, A.O., Hanski, I.K. & Raivio, S. 1996. Stochasticity in bird habitat selection: Year-to-year changes in territory locations in a boreal forest bird assemblage. Oikos 76: 536-552. Hansson, L. 1983. Bird numbers across edges between mature conifer forest and clearcuts in central Sweden. Ornis Scandinavica 14: 97-103. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1996. Influences of population size and woodland area on bird species distributions in small woods. Oecologia 105: 100-106. Hinsley, S.A., Hill, R.A., Fuller, R.J., Bellamy, P.E. & Rothery, P. 2009. Bird species distributions across woodland canopy structure gradients. Community Ecology 10: 99-110. Hogstad, O. 1999. Territorial behaviour of non-breeding Willow Warblers Phylloscopus trochilus during the breeding season. Ibis 141: 493-496. Hogstad, O. 2005. Numerical and functional responses of breeding passerine species to mass occurrence of geometrid caterpillars in a subalpine birch forest: a 30-year study. Ibis 147: 77-91. Holt, C.A., Fuller, R.J. & Dolman, P.M. 2011. Breeding and post-breeding responses of woodland birds to modification of habitat structure by deer. Biological Conservation 144: 2151-2162. Johnson, H. & Johnson, P. 2007. Nectar-, pollen- and oil-feeding. British Birds 100: 624-625. Kouki, J., Niemi, G.J. & Rajasarkka, A. 1992. Habitat associations of breeding peatland passerine species in eastern Finland. Ornis Fennica 69: 126-140. Krupa, M. 2004. Food of the willow warbler Phylloscopus trochilus nestlings: differences related to the age of nestlings and sex of feeding parents. Acta Ornithologica (Warsaw) 39: 45-51. Kuitunen, M., Rossi, E. & Stenroos, A. 1998. Do highways influence density of land birds? Environmental Management 22: 297-302. Lavers, C., McCullagh, M.J. & Fuller, R.J. 2005. Combining archive territory mapping data and aerial photography to investigate bird–habitat relationships: a case study from the Lincolnshire coast. Bird Study 52: 314-322. Mantyla, E., Klemola, T. & Haukioja, E. 2004. Attraction of willow warblers to sawfly- damaged mountain birches: novel function of inducible plant defences? Ecology Letters 7: 915-918. Marchetti, C., Locatelli, D.P., Van Noordwijk, A.J. & Baldaccini, N.E. 1998. The effects of prey size on diet differentiation of seven passerine species at two spring stopover sites. Ibis 140: 25-34. Mason, C.F. 1997. Association between Willow Warbler Phylloscopus trochilus territories and birch in woodlands in southeastern England. Ibis 139: 411-412. Mason, C.F. 2001. Woodland area, species turnover and the conservation of bird assemblages in lowland England. Biodiversity and Conservation 10: 495-510. Moss, D., Taylor, P.N. & Easterbee, N. 1979. The effects on song-bird populations of upland afforestation with spruce. Forestry 52: 129-150.

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Newson, S.E., Johnston, A., Renwick, A.R., Baillie, S.R. & Fuller, R.J. 2012. Modelling large-scale relationships between changes in woodland deer and bird populations. Journal of Applied Ecology 49: 278-286. Nilsson, S.G. & Ebenman, B. 1981. Density Changes and Niche Differences in Island and Mainland Willow Warblers Phylloscopus trochilus at a Lake in Southern Sweden. Ornis Scandinavica 12: 62-67. Nystrom, K.G.K. 1991. On sex-specific foraging behaviour in the Willow warbler, Phylloscopus trochilus. Canadian Journal of Zoology-Revue Canadienne De Zoologie 69: 462-470. Nystrom, K.G.K. 1997. Food density, song rate, and body condition in territory- establishing willow warblers (Phylloscopus trochilus). Canadian Journal of Zoology-Revue Canadienne De Zoologie 75: 47-58. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Patterson, I.J., Ollason, J.G. & Doyle, P. 1995. Bird populations in upland spruce plantations in northern Britain. Forest Ecology and Management 79: 107-131. Reijnen, R. & Foppen, R. 1994. The Effects of Car Traffic on Breeding Bird Populations in Woodland .1. Evidence of Reduced Habitat Quality for Willow Warblers (Phylloscopus-Trochilus) Breeding Close to a Highway. Journal of Applied Ecology 31: 85-94. Reijnen, R., Foppen, R. & Veenbaas, G. 1997. Disturbance by traffic of breeding birds: Evaluation of the effect and considerations in planning and managing road corridors. Biodiversity and Conservation 6: 567-581. Saether, B.E. 1983. Habitat selection, foraging niches and horizontal spacing of Willow warbler Phylloscopus-trochilus and Chiffchaff Phylloscopus-collybita in an area of sympatry. Ibis 125: 24-32. Smart, J., Taylor, E., Amar, A., Smith, K., Bierman, S., Carpenter, J., Grice, P., Currie, F. & Hewson, C. 2007. Habitat associations of woodland birds: implications for woodland management for declining species. In RSPB Research Report No., Vol. 26. Sandy: RSPB. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Sweeney, O.F.M., Wilson, M.W., Irwin, S., Kelly, T.C. & O'Halloran, J. 2010. Breeding bird communities of second-rotation plantations at different stages of the forest cycle. Bird Study 57: 301-314. Tiainen, J. 1983. Dynamics of a Local Population of the Willow Warbler Phylloscopus trochilus in Southern Finland. Ornis Scandinavica 14: 1-15. Tiainen, J., Vickholm, M., Pakkala, T., Piiroinen, J. & Virolainen, E. 1983. The habitat and spatial relations of breeding Phylloscopus warblers and the goldcrest Regulus regulus in southern Finland. Annales Zoologici Fennici 20: 1-12. Ulfstrand, S. 1976. Feeding niches of some passerine birds in a south Swedish coniferous plantation in winter and summer. Ornis Scandinavica 7: 21-27.

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Species nesting in a dense shrub layer

Garden Warbler Sylvia borin

1. Nest site selection/requirements Nest is located 0.5-1.5 m high in dense shrubby growth and is made of grass and herbs, lined with fine grasses and hair (Cramp and Perrins, 1993).

2. Territory selection/attributes Within orchards mean territory size was 0.19 ha in one study (Gottschalk, 1995). Breeding birds are site faithful, with males returning to breed within a mean of 60 m from the previous year’s territory, and females 130 m (Widmer, 1996). Territories are often selected to contain areas dominated by birch, and in UK dense low shrub cover particularly Bramble (Kouki et al., 1992, Smart et al., 2007).

3. Main food items During the breeding season Garden Warblers are mainly insectivorous and nestling diet is primarily caterpillars (Garcia, 1989). Garden Warblers start including fruit as part of diet as soon as they become available during July and August (Boddy, 1991, Eggers, 2000). A wide range of small berried fruits are eaten (Eggers, 2000, Jordano, 1987, Simons and Bairlein, 1990, Boddy, 1991, Hernandez, 2009), although a preference is shown for Elderberry when available (Boddy, 1991, Simons and Bairlein, 1990, Hernandez, 2009). Captive experiments show that frugivory is necessary for mass gain prior to migration, as a high protein diet inhibits mass gain (Bairlein, 1998). Of fruits tested Elder gave the highest mass gain (Simons and Bairlein, 1990).

During spring migration diet is mainly invertebrates, particularly crawling insects such as beetles and ants (Marchetti et al., 1998, Schwilch et al., 2001). They also consume nectar and berries such as Ivy when available (Schwilch et al., 2001, Amies, 1989).

4. Foraging methods Garden Warblers forage by gleaning in trees and shrubs 0.5-6 m from the ground (Cody, 1978, Kouki et al., 1992). They primarily forage on broadleaved trees and shrubs and occasionally in pine (Kouki et al., 1992).

5. Breeding habitat In the UK, Garden Warblers are primarily a species of broadleaved woodland but will also breed in young stages of conifer plantations (Currie and Bamford, 1981, Bibby et al., 1985, Smith et al., 1987, Smart et al., 2007). A UK scale study found occurrence most likely in woodlands with an open thicket stage understory (Hewson et al., 2011). Within this habitat they select for a high density of low shrub cover, particularly Bramble, and areas with a higher cover of birch trees, and low density of oaks (Smart et al., 2007, Smith et al., 1987, Currie and Bamford, 1981, Bibby et al., 1985). Garden Warblers are more likely to breed in woodlands within more wooded

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landscapes, and away from external woodland edges (Hinsley et al., 1995, Mason, 2001).

On continental Europe, they also prefer broadleaved woodland and avoid mature conifer dominated forest (Kouki et al., 1992, Morozov, 1992, Paquet et al., 2006, Müller, 2005, Baguette et al., 1994). In all forest types they occur most frequently in young stands <15-20 years old (Baguette et al., 1994, Paquet et al., 2006). They breed at higher densities with denser shrub cover at 0.5-4 m (Camprodon and Brotons, 2006).

6. Post fledging behaviour/requirements No information was found.

7. Seasonal differences in habitat for resident species On autumn migration, Garden Warblers use scrub with a high diversity of fruiting shrubs (Boddy, 1991).

8. Effects of forest management and deer on habitat Garden Warblers are strongly associated with broadleaf woodland and dense low shrub layers. Where they occur in young conifer plantations they are strongly associated with the presence of Bramble and birch. Any management that reduces canopy cover and promotes low shrub growth will benefit Garden Warbler populations. Such operations include clearfell, coppice, groupfell and thinning. Alternatively management that reduces shrub cover such as weeding of young crops will reduce habitat for Garden Warblers.

Deer browsing has largest effects in the low shrub layers of woodland which are used both for nesting and feeding by Garden Warblers. High levels of deer browsing are therefore likely to have a strong negative effect on Garden Warbler populations, although no such relationships were detected in a UK scale project (Newson et al., 2012). However this has been shown for coppice woodland, where the younger stages of growth had significantly more Garden Warblers when deer were excluded (Gill and Fuller, 2007, Holt et al., 2011). Any reduction in fruiting shrubs may adversely affect their ability to gain weight prior to migration.

9. References Amies, P. 1989. Ivy berries and birds in spring. Bristol Ornithology 20: 183-184. Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Bairlein, F. 1998. The effect of diet composition on migratory fuelling in Garden Warblers Sylvia borin. Journal of Avian Biology 29: 546-551. Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Boddy, M. 1991. Some aspects of frugivory by bird populations using coastal dune scrub in Lincolnshire. Bird Study 38: 188-199.

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Camprodon, J. & Brotons, L. 2006. Effects of undergrowth clearing on the bird communities of the Northwestern Mediterranean Coppice Holm oak forests. Forest Ecology and Management 221: 72-82. Cody, M.L. 1978. Habitat selection and interspecific territoriality among the Sylviid warblers of England and Sweden. Ecological Monographs 48: 351-396. Cramp, S. & Perrins, C.M. (1993) Handbook of the birds of Europe, the Middle East and North Africa: The birds of the Western Palearctic. Vol. VI Warblers, Oxford: Oxford University Press. Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Eggers, S. 2000. Compensatory frugivory in migratory Sylvia warblers: geographical responses to season length. Journal of Avian Biology 31: 63-74. Garcia, E. (1989) The blackcap and the garden warbler, Aylesbury: Shire. Gill, R.M.A. & Fuller, R.J. 2007. The effects of deer browsing on woodland structure and songbirds in lowland Britain. Ibis 149: 119-127. Gottschalk, T. 1995. Habitatanalyse von singervogelarten unter verwendung eines geographischen informationsystems (GIS). Vogelwelt 116: 273-284. Hernandez, A. 2009. Birds and guelder rose Viburnum opulus: selective consumption and dispersal via regurgitation of small-sized fruits and seeds. Plant Ecology 203: 111-122. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1995. Habitat and landscape factors influencing the presence of individual breeding bird species in woodland fragments. Journal of Avian Biology 26: 94-104. Holt, C.A., Fuller, R.J. & Dolman, P.M. 2011. Breeding and post-breeding responses of woodland birds to modification of habitat structure by deer. Biological Conservation 144: 2151-2162. Jordano, P. 1987. Frugivory, external morphology and digestive system in mediterranean sylviid warblers Sylvia spp. Ibis 129 (suppl 1): 175-189. Kouki, J., Niemi, G.J. & Rajasarkka, A. 1992. Habitat associations of breeding peatland passerine species in eastern Finland. Ornis Fennica 69: 126-140. Marchetti, C., Locatelli, D.P., Van Noordwijk, A.J. & Baldaccini, N.E. 1998. The effects of prey size on diet differentiation of seven passerine species at two spring stopover sites. Ibis 140: 25-34. Mason, C.F. 2001. Woodland area, species turnover and the conservation of bird assemblages in lowland England. Biodiversity and Conservation 10: 495-510. Morozov, N.S. 1992. Breeding forest birds in the Valdai uplands, north-west Russia - assemblage composition, interspecific associations and habitat amplitudes. Annales Zoologici Fennici 29: 7-28. Müller, J. 2005. Bird communities as indicators for woodland structures in oak woods. Ornithologische Beobachter 102: 15-32. Newson, S.E., Johnston, A., Renwick, A.R., Baillie, S.R. & Fuller, R.J. 2012. Modelling large-scale relationships between changes in woodland deer and bird populations. Journal of Applied Ecology 49: 278-286.

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Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Schwilch, R., Mantovani, R., Spina, F. & Jenni, L. 2001. Nectar consumption of warblers after long-distance flights during spring migration. Ibis 143: 24-32. Simons, D. & Bairlein, F. 1990. Neue Aspekte zur zugzeitlichen frugivorie der Gartengrasmucke (Sylvia borin). Journal Fur Ornithologie 131: 381-401. Smart, J., Taylor, E., Amar, A., Smith, K., Bierman, S., Carpenter, J., Grice, P., Currie, F. & Hewson, C. 2007. Habitat associations of woodland birds: implications for woodland management for declining species. In RSPB Research Report No., Vol. 26. Sandy: RSPB. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Widmer, M. 1996. Phenology, breeding density and population ecology of the garden warbler Sylvia borin in a subalpine habitat of the central Swiss Alps. Journal Fur Ornithologie 137: 479-501.

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Final report WC0793/CR0485 (29.04.14)

Song Thrush Turdus philomelos

1. Nest site selection/requirements Song Thrush nests are constructed from a base of grass and roots and lined with mud or softened wood pulp, and it is likely that the presence of wet areas or water may be necessary for nest construction (Tomialojc, 1992).

The nest is normally located 1-5 m above the ground in a tree or shrub (Schnack, 1991, Spaar and Hegelbach, 1994, Edula, 1997, Kelleher and O'Halloran, 2007). Where Norway spruce is available it is strongly selected as a nesting tree (Spaar and Hegelbach, 1994, Edula, 1997, Götmark et al., 1995). Other nest sites commonly used include Juniper, shrubs, hedges, broadleaved trees, fallen trees/brash and pines (Götmark et al., 1995, Edula, 1997, Schnack, 1991, Kelleher and O'Halloran, 2007). In a spruce forest, nest sites were found in exposed to completely concealed sites, most nests were partly concealed, but the degree of concealment had no relationship with nest predation rates (Götmark et al., 1995). Although in a more mixed habitat, including parks and gardens, Sitka spruce plantation and mixed coppice, well concealed nests had a lower probability of predation (Kelleher and O'Halloran, 2007). Nest locations are often located in areas of denser trees or shrubs (Götmark et al., 1995, Kelleher and O'Halloran, 2007).

2. Territory selection/attributes It appears that Song Thrushes select areas within the landscape which have a higher density of dense shrub or tree cover. Within farmed landscapes Song Thrush territories are located along hedgerows and gardens more frequently than expected based on availability, and within a mixed farmed landscape woodland was used more often than expected except where farming was arable and with little woodland (Peach et al., 2004). Within mixed forest there was a strong selection for areas with Norway spruce, 81% of territories were centred on spruce which only covered 16% of the study area (Spaar and Hegelbach, 1994).

During breeding in farmed landscapes males ranged over 1.5 ha and females 0.5 ha, with the foraging area increasing seasonally; birds from later nests had larger foraging ranges than early nests (Peach et al., 2004). Territory size depends on habitat, with smaller territories in gardens than farmland (Peach et al., 2004).

3. Main food items Nestlings are mostly fed invertebrates with the main prey items being earthworms, insect larvae including caterpillars, snails, beetles and occasionally spiders, flies and plant material (Schnack, 1991, Gruar et al., 2003). Proportions of these different groups fed to nestlings vary between studies (Gruar et al., 2003, Schnack, 1991). Studies in English farmland and garden habitats found earthworms the main food item early in the breeding season, replaced by snails latter in the breeding season (Gruar et al., 2003). A study in Austria found that populations in city parks fed a considerable amount of fruit to nestlings with 30-100% of diet from plant sources, while a nearby woodland population fed nestlings predominantly on caterpillars

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(Schnack, 1991). Adult diet in the breeding season is similar to nestlings (Gruar et al., 2003).

In autumn and winter Song Thrushes additionally feed on berries to a lesser extent compared to Blackbirds (Boddy, 1991, Hernandez, 2003).

4. Foraging methods During the breeding season Song Thrushes feed mainly on the ground, but sometimes feed in trees on caterpillars (Hogstad, 2005, Ulfstrand, 1976). When feeding in the open on grassy areas they mostly forage within 50 m of trees or other cover, coinciding with areas of highest prey density (Greenwood and Harvey, 1978). Three feeding techniques were observed during dry summer weather; snatching at flying insects or from edges of foliage (63% of observations); jabbing at insects and spiders on the surface of the ground (36%), and probing the ground for subsurface invertebrates such as earthworms (1%) (Greenwood and Harvey, 1978). The proportional use of these foraging methods was dependent on prey availability and dry conditions; probing dominates when soil moisture is higher which is when earthworms become an important prey item. Rate of ground feeding is highest in the early morning and snatching for flying insects during the middle part of the day when flying insects were most active (Greenwood and Harvey, 1978). In winter, Song Thrush will also feed on small fruit by plucking them from bushes (Hernandez, 2003).

5. Breeding habitat In the UK most breeding populations occur in gardens and farmland, however relative to availability woodland, scrub and human habitats are preferred habitats (Gregory and Baillie, 1998, Mason, 2000). At the landscape scale mixed landscapes including arable, pastoral gardens and woodland have higher densities compared to mainly arable landscapes (Peach et al., 2002). Within farmland landscapes, Song Thrushes are more likely to select hedgerows and woodland edges than would be expected from random habitat selection (Whittingham et al., 2009). This preference for habitats where woodland and farmland meet in lowland landscapes is also suggested by greater occupation than expected in small woods compared with large woods, with a higher occurrence in small woods related to the presence of hedgerows and nearby human habitats (Hinsley et al., 1995, Hinsley et al., 1996). Within the UK, Song Thrushes are more abundant in broadleaved woodland with low Bramble or a mixed species understory with more dead wood on the ground (Hewson et al., 2011, Carpenter et al., 2009, Smith et al., 1987). Another study found no detectable habitat selection in oak plantations (Donald et al., 1998). Song Thrushes occur at low densities in upland conifer plantations mainly, but not exclusively, in 5-15 year old replanted areas (Moss et al., 1979, Currie and Bamford, 1981, Bibby et al., 1985). Woodlands with an ericaceous understory have a low occurrence (Hewson et al., 2011). In contrast, in northern continental Europe Song Thrushes regularly breed in conifer forests and show a preference for Norway spruce (Gjerde and Saetersdal, 1997, Hansson, 1983, Haila et al., 1996). Within large size forests Song Thrushes show higher densities in forest edges adjacent to clearcuts (Hansson, 1983). Song Thrushes were frequent across a wide range of forest types in a heavily forested landscape, including mature Beech, oak and spruce as well as

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forest edges and young forest stands; they were only scarce in spruce plantations younger than 5 years old (Paquet et al., 2006, Baguette et al., 1994). In contrast to the UK, a study on continental Europe found that forest was better habitat, with higher densities and breeding success, than vegetated urban areas (Schnack, 1991).

6. Post fledging behaviour/requirements Juvenile Song Thrushes mostly disperse <20 km in first months after fledging (Milwright, 2006).

7. Seasonal differences in habitat for resident species Little is known of changes in habitat outside the breeding season, complicated by partial migration within the UK and an influx of continental birds. One year round study found they used similar habitats in winter to the breeding season, but also used game cover crops as has been found elsewhere (Parish and Sotheton, 2004, Peach et al., 2004, Hinsley et al., 2010).

8. Effects of forest management and deer on habitat Song Thrushes nest in, and territories are centred on, areas of dense shrub. However woodland edges appear very important for foraging, allowing access to both woodland and open habitats which provide different food resources under different weather and soil conditions. As primarily a ground forager, a dense field layer reduces habitat quality. Management that promotes a dense shrub layer is likely to be beneficial; this includes group felling, coppice, heavy thinning and clearfell. Clearfell and coppice can also provide woodland edges which may be beneficial soon after harvest before a dense field layer develops. Dense canopy woodlands can provide an open ground layer suitable for foraging, especially where damp soil conditions are prevalent.

Heavy deer browsing is likely to have a negative effect on Song Thrush habitats through a reduction in nesting cover and changing an herbaceous field layer to one dominated by coarse grasses making the woodland floor less suitable for foraging. At the UK scale Song Thrush abundance is negatively associated with higher densities of Reeves’ muntjac deer (Newson et al., 2012) and another study found Song Thrush populations declined at a greater rate with high deer populations (Perrins and Overall, 2001).

9. References Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Boddy, M. 1991. Some aspects of frugivory by bird populations using coastal dune scrub in Lincolnshire. Bird Study 38: 188-199. Carpenter, J., Charman, E., Smart, J., Amar, A., Gruar, D. & Grice, P. 2009. Habitat associations of woodland birds II. In RSPB Research Report. Vol. 36.

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Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Donald, P.F., Fuller, R.J., Evans, A.D. & Gough, S.J. 1998. Effects of forest management and grazing on breeding bird communities in plantations of broadleaved and coniferous trees in western England. Biological Conservation 85: 183-197. Edula, E. 1997. Anmeid Vainu- ja Laulurasta pesitsemisest Vijandi umbruses aastatel. Hirundo 2: 3-13. Gjerde, I. & Saetersdal, M. 1997. Effects on avian diversity of introducing spruce Picea spp. plantations in the native pine Pinus sylvestris forests of western Norway. Biological Conservation 79: 241-250. Götmark, F., Blömqvist, D., Johansson, O.C. & Bergkvist, J. 1995. Nest site selection: A trade-off between concealment and view of the surroundings? Journal of Avian Biology 26: 305-312. Greenwood, P.J. & Harvey, P.H. 1978. Foraging and territory utilization of blackbirds (Turdus merula) and song thrushes (Turdus philomelos). Animal Behaviour 26: 1222-1236. Gregory, R.D. & Baillie, S.R. 1998. Large-scale habitat use of some declining British birds. Journal of Applied Ecology 35: 785-799. Gruar, D., Peach, W. & Taylor, R. 2003. Summer diet and body condition of Song Thrushes Turdus philomelos in stable and declining farmland populations. Ibis 145: 637-649. Haila, Y., Nicholls, A.O., Hanski, I.K. & Raivio, S. 1996. Stochasticity in bird habitat selection: Year-to-year changes in territory locations in a boreal forest bird assemblage. Oikos 76: 536-552. Hansson, L. 1983. Bird numbers across edges between mature conifer forest and clearcuts in central Sweden. Ornis Scandinavica 14: 97-103. Hernandez, A. 2003. Variations in spindle Euonymus europaeus consumption by frugivorous birds during the fruiting season. Ardeola 50: 171-180. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1995. Habitat and landscape factors influencing the presence of individual breeding bird species in woodland fragments. Journal of Avian Biology 26: 94-104. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1996. Influences of population size and woodland area on bird species distributions in small woods. Oecologia 105: 100-106. Hinsley, S.A., Redhead, J.W., Bellamy, P.E., Broughton, R.K., Hill, R.A., Heard, M.S. & Pywell, R.F. 2010. Testing agri-environment delivery for farmland birds at the farm scale: the Hillesden experiment. Ibis 152: 500-514. Hogstad, O. 2005. Numerical and functional responses of breeding passerine species to mass occurrence of geometrid caterpillars in a subalpine birch forest: a 30-year study. Ibis 147: 77-91. Kelleher, K.M. & O'Halloran, J. 2007. Influence of nesting habitat on breeding Song Thrushes Turdus philomelos. Bird Study 54: 221-229.

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Mason, C.F. 2000. Thrushes now largely restricted to the built environment in eastern England. Diversity and Distributions 6: 189-194. Milwright, R.D.P. 2006. Post-breeding dispersal, breeding site fidelity and migration/wintering areas of migratory populations of Song Thrush Turdus philomelos in the Western Palearctic. Ringing and Migration 23: 21-32. Moss, D., Taylor, P.N. & Easterbee, N. 1979. The effects on song-bird populations of upland afforestation with spruce. Forestry 52: 129-150. Newson, S.E., Johnston, A., Renwick, A.R., Baillie, S.R. & Fuller, R.J. 2012. Modelling large-scale relationships between changes in woodland deer and bird populations. Journal of Applied Ecology 49: 278-286. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Parish, D.M.B. & Sotheton, N.W. 2004. Game crops and threatened farmland songbirds in Scotland: a step towards halting population declines? Bird Study 51: 107-112. Peach, W., Taylor, R., Cotton, P.A., Gruar, D., Hill, I.F. & Denny, M. 2002. Habitat utilisation by Song thrushes Turdus philomelos on lowland farmland during summer and winter. Aspects of Applied Biology 67: 11-20. Peach, W.J., Denny, M., Cotton, P.A., Hill, I.F., Gruar, D., Barritt, D., Impey, A. & Mallord, J. 2004. Habitat selection by song thrushes in stable and declining farmland populations. Journal of Applied Ecology 41: 275-293. Perrins, C.M. & Overall, R. 2001. Effect of increasing numbers of deer on bird populations in Wytham Woods, central England. Forestry 74: 299-309. Schnack, S. 1991. The breeding biology and nestling diet of the blackbird Turdus merula L. and the song thrush Turdus philomelos C.L. Brehm in Vienna and in an adjacent wood. Acta Ornithologica 26: 85-106. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Spaar, R. & Hegelbach, J. 1994. Neststandort und Brutbiologie der singdrossel Turdus philomelos im Zurichbergwald. der Ornithologische Beobachter 91: 31-41. Tomialojc, L. 1992. Colonization of dry habitats by the song thrush Turdus philomelos: is the type of nest material an important constraint? Bulletin of the British Ornithologists' Club 112: 27-34. Ulfstrand, S. 1976. Feeding niches of some passerine birds in a south Swedish coniferous plantation in winter and summer. Ornis Scandinavica 7: 21-27. Whittingham, M.J., Krebs, J.R., Swetnam, R.D., Thewlis, R.M., Wilson, J.D. & Freckleton, R.P. 2009. Habitat associations of British breeding farmland birds. Bird Study 56: 43-52.

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Final report WC0793/CR0485 (29.04.14)

Nightingale Luscinia megarhynchos

1. Nest site selection/requirements Nightingale nests are usually located on or close to the ground with few located higher than 30cm above ground level, and none above 240 cm, with an average nest height of 20 cm in the UK and 69 cm in Italy (Morgan, 1982, Caffi, 2004, Wilson et al., 2005). Dense low vegetation is required to conceal nests and nests are often situated within a dense field layer at edges of thickets (Wilson et al., 2005) and are frequently located in rose, Bramble, nettle, gorse although many species of small shrubs and trees are used (Sharipov, 1977, Morgan, 1982). Nests are typically constructed mainly of dead leaves, often oak, lined with fine grasses, feathers and hair (Collar, 2005, Fuller et al., 1999).

2. Territory selection/attributes Average breeding home range in one study in UK was 14,114m2 based on radio tracking, whereas in the same study area mean male singing territory area based on mapped singing locations was only 1,691 m2 (Holt et al., 2010). Mean areas of singing territories of 6,745 m2 were recorded in Austria (Gruell, 1981), 5,400 m2 in Germany (Naguib et al., 2001) and 1,690 m2 in England (Holt et al., 2010, Holt et al., 2012). Territory size is smaller where the proportion of dense cover is higher (Bayes and Henderson, 1988). Adult nightingales exhibit high site fidelity, with male site fidelity higher than for females (Boano et al., 2004, Gruell, 1981). In Austria 47.5% of adult males returned to previous breeding sites with 76% of these occupying the same or adjacent territory (Gruell, 1981).

Territory selection is very specific, requiring a variety of vegetation structures in close proximity to provide song posts, nesting habitat, feeding habitat and cover for fledged young. Territories contain habitat of a particular structure rather than containing particular tree and shrub species (Hewson et al., 2005, Bayes and Henderson, 1988, Wilson et al., 2005) although in England territories are often associated with higher oak standard density (Fuller et al., 1989, Stuttard and Williams.K, 1971). Territories are situated in woodland or scrub with a dense and continuous canopy, and dense shrub layer dominated by shrubs or trees 2-5 m high (Holt et al., 2012, Hudson, 1979, Fuller and Henderson, 1992, Wilson et al., 2005). Ideal vegetation structure in scrub habitats comprises a shell-like densely foliated scrub layer 1.5-2.5 m high that typically extends to the ground at its edges and is surrounded by a dense growth of field layer vegetation (Wilson et al., 2005). In woodlands containing Nightingales horizontal visibility is on average 4.1 m while in woodlands without Nightingales the average is 8.6 m (Bayes and Henderson, 1988). Nightingales additionally have a requirement within territories for areas with little or no ground vegetation, an average of 61% bare ground in the centres of territories (Wilson et al., 2005). In England proximity of scrub to patches of mature trees may also be important in territory selection (Holt et al., 2012, Hudson, 1979). Coppiced woodlands with coppice aged 3-9 years after cutting provide optimal structure for territories, with peak nightingale densities found around 6 years after cutting by several studies, while coppice less than 3 years and older than 10 years is avoided (Fuller et al., 1989, Fuller and Henderson, 1992, Holt et al., 2010, Stuttard and

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Williams.K, 1971). Territories in mature coppice are usually associated with edge of the coppice block (Fuller et al., 1989).

Territory establishment follows a despotic settlement pattern, related to adult age, with more marginal habitat types increasingly selected at higher densities (Nemethova, 2007, Gruell, 1981).

In England territories are situated, and occur at higher densities, in areas with soils associated with higher moisture content and higher organic and nutrient levels, especially earthy peat or humic sandy gley soils which may be associated with higher food availability (Wilson et al., 2005).

3. Main food items Diet comprises a wide variety of small invertebrates, but predominantly ants and beetles found in the leaf litter or on bare ground (Cramp, 1988). Other invertebrates taken include spiders, caterpillars, flies, earthworms, grasshoppers, millipedes and crustaceans (Collar, 2005). They also take berries and seeds in late summer and autumn including strawberry, cherry, currant, Alder buckthorn, Elder, service and Dogwood (Collar, 2005).

4. Foraging methods Nightingales forage for invertebrates mainly on bare ground and in leaf litter under dense cover, infrequently feeding in the open. They occasionally drop from a perch onto prey, glean on low branches and leaves and make aerial sallies after flying invertebrates (Collar, 2005).

5. Breeding habitat Breeding habitat is found at low elevations; all English territories are located below 244 m, with most below 60 m (Morgan, 1982, Wilson et al., 2002). Habitat selection is more based on vegetation structure than species composition, with dense shrub cover, bare ground and a deep litter layer as important attributes. Nightingale are associated primarily with scrub and dense woodland but are found in a variety of habitat types with suitable structure including coppiced woodland, mature broadleaf woodland particularly containing high proportions of oak, birch, Elder or willow or thickets of Hazel, Blackthorn, Hawthorn or Holly, young mixed or coniferous woodlands, Carr woodland, scrubby heathland and occasionally hedgerows and human influenced scrub habitats such as railway embankments and former gravel pits (Morgan, 1982, Wilson et al., 2002, Hudson, 1979, Wilson et al., 2005). In England they occur at a higher density in some scrub habitats with an apparent shift from woodland to scrub dominated habitats as range has contracted (Wilson et al., 2002), and less often in coniferous and mixed plantations as fewer young plantations are available due to the maturation of former habitat (Fuller et al., 1999). Nightingales prefer areas with wetter soils with high nutrient and organic content (Wilson et al., 2005).

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In fragmented habitats woodland size may affect habitat occupancy, with greater abundance in woodlands >10 ha (Mason, 2001), although in this study all woodland sizes were relatively small with few woods >10 ha.

6. Post fledging behaviour/requirements Young leave the nest 3-5 days before they can fly and it is thought that dense field vegetation provides refuge for recently fledged young (Cramp, 1988). Males may look after the fledged brood while females begin incubating a second clutch with young becoming independent after 15-30 days, by which time broods may have divided and dispersed beyond the territory (Collar, 2005).

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Key habitat requirement for Nightingale in England is a dense scrub layer, dense closed canopy trees or shrubs with high shrub layer foliage density, and areas of bare ground and leaf litter. Coppicing provides particularly suitable habitat when it has the greatest stem density, 3-10 years after harvesting and just prior to canopy closure. Coppice immediately post-harvest and once canopy has closed is unsuitable, and monoculture or closely spaced coppice, particularly of Sweet chestnut, is less suitable because it has little low undergrowth or dense scrub layer (Fuller et al., 1999). Thinning, clearfell, selective felling and cleaning operations to reduce weedy tree species from within crops are all likely to be detrimental to Nightingale populations as are any forestry operations that include removal of scrub or thickets.

Nightingale populations are not able to persist with high levels of deer browsing (Wilson et al., 2002), and even moderate levels of deer browsing are likely to detrimentally modify Nightingale habitat (Fuller, 2001). Deer browsing strongly alters vegetation structure by reducing canopy cover and shrub layer foliage density, although deer do not affect invertebrate density per unit of foliage (Holt et al., 2011). Development of grasses and sedges in deer browsed areas probably reduce the quality of habitat for feeding (Fuller and Henderson, 1992). An English study in coppiced woodland experimentally showed very strong preference by Nightingale for plots where deer were excluded in terms of both settlement patterns and habitat use (Holt et al., 2010). At the UK scale Nightingale abundance is negatively associated with higher densities of Roe deer (Newson et al., 2012).

9. References Bayes, K. & Henderson, A.C.B. 1988. Nightingales and Coppiced Woodland. In RSPB Conservation Review. Vol. 2, pp. 47-49. RSPB. Boano, G., Bonardi, A. & Silvano, F. 2004. Nightingale Luscinia megarhynchos survival rates in relation to Sahel rainfall. Avocetta 28: 77-85. Caffi, M. 2004. Breeding biology of nightingale, Luscinia megarhynchos, nesting in a river bank wood in the Po Plane in Lombardy (Italy). Rivista Italiana di Ornitologia 73: 125-132.

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Collar, N. 2005. Nightingale. In Handbook of Birds of the World. (Dds. J. del Hoyo, A. Elliott & D. Christie), Vol. 10, pp. 746. Barcelona: Lynx Edicions. Cramp, S. 1988. In Handbook of the Birds of Europe, the Middle East and North Africa. The Birds of the Western Palearctic. Volume 5: Tyrant flycatchers to thrushes. Vol. 5. Oxford: Oxford University Press. Fuller, R., Stuttard, P. & Ray, C. 1989. The distribution of breeding songbirds within mixed coppiced woodland in Kent, England, in relation to vegetation age and structure. Annales Zoologici Fennici 26: 265-275. Fuller, R.J. 2001. Responses of woodland birds to increasing numbers of deer: a review of evidence and mechanisms. Forestry 74: 289-298. Fuller, R.J. & Henderson, A.C.B. 1992. Distribution of breeding songbirds in Bradfield woods, Suffolk, in relation to vegetation and coppice management. Bird Study 39: 73-88. Fuller, R.J., Henderson, A.C.B. & Wilson, A. 1999. The Nightingale in England - problems and prospects. British Wildlife 10: 221-230. Gruell, A. 1981. Investigations on the territory of the Nightingale Luscinia megarhynchos. Journal fuer Ornithologie 122: 259-284. Hewson, C.M., Fuller, R.J. & Day, C. 2005. An investigation of habitat occupancy by the nightingale Luscinia megarhynchos with respect to population change at the edge of its range in England. Journal of Ornithology 146: 244-248. Holt, C.A., Fraser, K.H., Bull, A.J. & Dolman, P.M. 2012. Habitat use by Nightingales in a scrub–woodland mosaic in central England. Bird Study 59: 416-425. Holt, C.A., Fuller, R.J. & Dolman, P.M. 2010. Experimental evidence that deer browsing reduces habitat suitability for breeding Common Nightingales Luscinia megarhynchos. Ibis 152: 335-346. Holt, C.A., Fuller, R.J. & Dolman, P.M. 2011. Breeding and post-breeding responses of woodland birds to modification of habitat structure by deer. Biological Conservation 144: 2151-2162. Hudson, R. 1979. Nightingales in Britain in 1976. Bird Study 26: 204-212. Mason, C.F. 2001. Woodland area, species turnover and the conservation of bird assemblages in lowland England. Biodiversity and Conservation 10: 495-510. Morgan, R. 1982. The breeding biology of the Nightingale Luscinia megarhynchos in Britain. Bird Study 29: 67-72. Naguib, M., Altenkamp, R. & Griessmann, B. 2001. Nightingales in space: song and extra-territorial forays of radio tagged song birds. Journal Fur Ornithologie 142: 306-312. Nemethova, D. 2007. Density-dependent habitat selection in nightingale (Luscinia megarhynchos C.L. Brehm) in selected windbreaks of SW Slovakia. Ekologia (Bratislava) 26: 313-321. Newson, S.E., Johnston, A., Renwick, A. R., Baillie, S.R. & Fuller, R.J. 2012. Modelling large-scale relationships between changes in woodland deer and bird populations. Journal of Applied Ecology 49: 278-286. Sharipov, M.A. 1977. Nesting behavior of Luscinia megarhynchos hafizi in towns of the Ferghana Valley. Vestnik Zoologii: 24-28. Stuttard, P. & Williams.K. 1971. Habitat requirements of Nightingale. Bird Study 18: 9-&.

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Wilson, A.M., Fuller, R.J., Day, C. & Smith, G. 2005. Nightingales Luscinia megarhynchos in scrub habitats in the southern fens of East Anglia, England: associations with soil type and vegetation structure. Ibis 147: 498-511. Wilson, A.M., Henderson, A.C.B. & Fuller, R.J. 2002. Status of the Nightingale Luscinia megarhynchos in Britain at the end of the 20th Century with particular reference to climate change. Bird Study 49: 193-204.

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Final report WC0793/CR0485 (29.04.14)

Dunnock Prunella modularis

1. Nest site selection/requirements Dunnocks build an open cup nest of twigs, stems and roots lined with hair, moss and feathers well concealed in low woody vegetation, 0.5-3.5 m from ground (Cramp, 1988). In managed conifer forest, nests are mostly in young Spruce trees (63%) or brash piles (19%) (Tuomenpuro, 1991). Nest sites are selected where tree density and cover of trees is higher compared with random locations, with nest success higher for more concealed nests (Tuomenpuro, 1991).

2. Territory selection/attributes Territory sizes vary from 0.75 -2.75 ha, and are selected in areas with dense shrubby cover such as overgrown hedgerows or young spruce trees (Tuomenpuro, 1989, Bishton, 2001).

3. Main food items Nestlings are primarily fed invertebrates with seeds occasionally fed to nestlings, particularly in later broods (Bishton, 1985, Moreby and Stoate, 2001). The main invertebrate groups fed to nestlings are beetles, spiders, flies and caterpillars (Bishton, 1985, Moreby and Stoate, 2001).

During the breeding season adults feed primarily on invertebrates especially beetles and spiders along with smaller numbers of gastropods, worms, flies and springtails (Bishton, 1986). From August through to March seeds also form an important component of diet with up to 50% of winter food being seed (Bishton, 1986, Martinez-Cabello et al., 1991). In the UK, frequently taken seeds are nettle, Yorkshire fog, Elder and dock (Bishton, 1986).

4. Foraging methods The most common foraging technique used by Dunnocks is picking seeds and invertebrates from the ground (82% of observations throughout the year) (Bishton, 1986). During the breeding season and in autumn they will also take invertebrates from foliage or seeds direct from plants, and rarely in late winter from soft soil or leaf litter by digging (Bishton, 1986, Shaw, 1996).

5. Breeding habitat In the UK, Dunnocks breed in a wide variety of habitats including farmland, gardens, scrub, conifer woodland, broadleaf woodland and moorland (Currie and Bamford, 1981, Bibby et al., 1985, Bevington, 1991, Fuller et al., 2001, Gregory and Baillie, 1998, Hinsley et al., 1995, Lavers et al., 2005, Moss et al., 1979, Patterson et al., 1995, Smith et al., 1987). Highest densities are found in habitats with a dense shrub layer and Dunnock have been shown to prefer scrub, overgrown hedgerows and gardens over woodland and other habitats (Bevington, 1991, Fuller et al., 2001, Lavers et al., 2005). In broadleaved woodland Dunnock prefer smaller woods and are more likely to breed in woods with greater Bramble cover or containing mature scrub (Hinsley et al., 1995, Bellamy et al., 2000, Smart et al., 2007). In conifer woodland and broadleaved plantations Dunnock select growth stages with the

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densest cover in the shrub layer, including thicket stages of newly afforested areas with trees >5 m tall, and younger growth stages in replanted areas (Moss et al., 1979, Currie and Bamford, 1981, Bibby et al., 1985, Wilson et al., 2006, Hewson et al., 2011). Dunnock are scarce in the youngest establishment phase and after canopy closure (Moss et al., 1979, Smith et al., 1987, Patterson et al., 1995, Wilson et al., 2006). Within woodlands, Dunnock occur at higher densities where there are large gaps in the canopy allowing thick shrub growth, than under closed canopy (Fuller, 2000). In heavily wooded landscapes, Dunnock occurred in a wide range of forest types but are most frequent in young forest <15 years old and forest edges; in mature forest they were more frequent in conifer than broadleaf (Baguette et al., 1994, Paquet et al., 2006).

6. Post fledging behaviour/requirements No information was found.

7. Seasonal differences in habitat for resident species In winter most Dunnock leave upland habitats but otherwise are largely sedentary and remain in the same habitat (Davies, 1987, Bevington, 1991, Patterson et al., 1995). Within agricultural habitats Dunnocks use game cover crops and provided ‘wild bird seed’ food patches in winter but rarely conventional crops or stubbles (Parish and Sotheton, 2004, Hinsley et al., 2010).

8. Effects of forest management and deer on habitat The presence of dense low cover is important for nesting, territory selection and providing feeding cover. Any management that reduces canopy cover and promotes low shrub growth will benefit Dunnock populations. Such operations include clearfell, coppice, groupfell and thinning. Dunnocks also forage under rank herbaceous vegetation such as nettles, and disturbance due to forestry operations may provide areas of such feeding habitat.

A UK scale study found no relationships between deer density and Dunnock abundance (Newson et al., 2012). However heavy deer browsing is likely to have a negative effect on Dunnock habitats through a reduction in nesting cover and changing a herbaceous field layer to one dominated by coarse grasses making the woodland floor less suitable for foraging. In coppice woodland significantly higher numbers of Dunnock were found when deer were excluded (Holt et al., 2011) and in more varied woodland Dunnock decreased with an increase in deer (Perrins and Overall, 2001).

9. References Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Bellamy, P.E., Rothery, P., Hinsley, S.A. & Newton, I. 2000. Variation in the relationship between numbers of breeding pairs and woodland area for passerines in fragmented habitat. Ecography 23: 130-138.

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Bevington, A. 1991. Habitat selection in the Dunnock Prunella modularis in northern England. Bird Study 38: 87-91. Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Bishton, G. 1985. The diet of nestling Dunnocks Prunella modularis. Bird Study 32: 113-115. Bishton, G. 1986. The diet and foraging behaviour of the Dunnock Prunella-modularis in a hedgerow habitat. Ibis 128: 526-539. Bishton, G. 2001. Social structure, habitat use and breeding biology of hedgerow Dunnocks Prunella modularis. Bird Study 48: 188-193. Cramp, S. (1988) Handbook of the Birds of Europe, the Middle East and North Africa. The Birds of the Western Palearctic. Volume 5: Tyrant flycatchers to thrushes. Oxford: Oxford University Press. Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Davies, N.B. 1987. Studies of West Palearctic birds: 188. Dunnock. British Birds 80: 604-624. Fuller, R.J. 2000. Influence of treefall gaps on distributions of breeding birds within interior old-growth stands in Bialowieza forest, Poland. Condor 102: 267-274. Fuller, R.J., Chamberlain, D.E., Burton, N.H.K. & Gough, S.J. 2001. Distributions of birds in lowland agricultural landscapes of England and Wales: How distinctive are bird communities of hedgerows and woodland? Agriculture Ecosystems & Environment 84: 79-92. Gregory, R.D. & Baillie, S.R. 1998. Large-scale habitat use of some declining British birds. Journal of Applied Ecology 35: 785-799. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T H. 1995. Habitat and landscape factors influencing the presence of individual breeding bird species in woodland fragments. Journal of Avian Biology 26: 94-104. Hinsley, S.A., Redhead, J.W., Bellamy, P.E., Broughton, R.K., Hill, R.A., Heard, M.S. & Pywell, R.F. 2010. Testing agri-environment delivery for farmland birds at the farm scale: the Hillesden experiment. Ibis 152: 500-514. Holt, C.A., Fuller, R.J. & Dolman, P.M. 2011. Breeding and post-breeding responses of woodland birds to modification of habitat structure by deer. Biological Conservation 144: 2151-2162. Lavers, C., McCullagh, M.J. & Fuller, R.J. 2005. Combining archive territory mapping data and aerial photography to investigate bird–habitat relationships: a case study from the Lincolnshire coast. Bird Study 52: 314-322. Martinez-Cabello, A., Soler, M. & Soler, J.J. 1991. Alimentacion del acetor comun (Prunella modularis) durante su invernada en el sureste de la peninsula Iberica. Ardeola 38: 305-315. Moreby, S.J. & Stoate, C. 2001. Relative abundance of invertebrate taxa in the nestling diet of three farmland passerine species, Dunnock Prunella modularis, Whitethroat Sylvia communis and Yellowhammer Emberzia citrinella in Leicestershire, England. Agriculture Ecosystems & Environment 86: 125-134.

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Moss, D., Taylor, P. N. & Easterbee, N. 1979. The effects on song-bird populations of upland afforestation with spruce. Forestry, 52: 129-150. Newson, S. E., Johnston, A., Renwick, A. R., Baillie, S. R. & Fuller, R. J. 2012. Modelling large-scale relationships between changes in woodland deer and bird populations. Journal of Applied Ecology 49: 278-286. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Parish, D.M.B. & Sotheton, N.W. 2004. Game crops and threatened farmland songbirds in Scotland: a step towards halting population declines? Bird Study 51: 107-112. Patterson, I.J., Ollason, J.G. & Doyle, P. 1995. Bird populations in upland spruce plantations in northern Britain. Forest Ecology and Management 79: 107-131. Perrins, C.M. & Overall, R. 2001. Effect of increasing numbers of deer on bird populations in Wytham Woods, central England. Forestry 74: 299-309. Shaw, G. 1996. Dunnock feeding in spruce canopy. Scottish Birds 18: 185. Smart, J., Taylor, E., Amar, A., Smith, K., Bierman, S., Carpenter, J., Grice, P., Currie, F. & Hewson, C. 2007. Habitat associations of woodland birds: implications for woodland management for declining species. In RSPB Research Report No., Vol. 26. Sandy: RSPB. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Tuomenpuro, J. 1989. Habitat preferences and territory size of the Dunnock Prunella modularis in southern Finland. Ornis Fennica 66: 133-141. Tuomenpuro, J. 1991. Effect of nest site on nest survival in the Dunnock Prunella modularis. Ornis Fennica 68: 49-56. Wilson, M.W., Pithon, J., Gittings, T., Kelly, T.C., Giller, P.S. & O'Halloran, J. 2006. Effects of growth stage and tree species composition on breeding bird assemblages of plantation forests. Bird Study 53: 225-236.

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Final report WC0793/CR0485 (29.04.14)

Lesser Redpoll Carduelis cabaret

1. Nest site selection/requirements Lesser Redpoll build an open cup nest of twigs lined with plant material, hair or feathers in a tree or shrub (Cramp et al., 1994).

2. Territory selection/attributes No information was found.

3. Main food items During the breeding season diet is mainly a range of invertebrates taken from trees (Cramp et al., 1994). Winter diet consists principally of seeds from trees and herbaceous plants with the main tree species used being birch, larch, spruce and alder (Cramp et al., 1994).

4. Foraging methods Foraging mainly occurs in trees (Cramp et al., 1994), but very little information is available.

5. Breeding habitat Although Lesser Redpoll are recorded from a wide range of woodland types and habitats with dense scrub, its main breeding habitat is young spruce plantations (Smith et al., 1987, Patterson et al., 1995, Lavers et al., 2005, Smart et al., 2007). Lesser Redpolls are a scarce breeding species in broadleaved woodland (Smith et al., 1987, Smith et al., 1992, Smart et al., 2007). Within broadleaved woodland, occupied woods have more moss and lichen cover than unoccupied woods, fewer dead limbs and less Ivy (Smart et al., 2007). Highest densities of breeding redpolls are found in upland spruce plantations where they principally breed in young growth stages of crops of 5-15 years old with few, if any, in older plantations (Moss et al., 1979, Patterson et al., 1995, Sweeney et al., 2010). Within young conifer plantations breeding densities are higher in newly afforested areas than in replanted second rotation forests (Currie and Bamford, 1981, Sweeney et al., 2010) and in replanted areas plots with a higher cover of conifers and few other tree species had higher breeding densities (Bibby et al., 1985). Outbreaks of November moths on birch increase habitat quality with higher breeding densities and breeding success in outbreak years (Hogstad, 2005).

6. Post fledging behaviour/requirements No information was found.

7. Seasonal differences in habitat for resident species Little information was found on winter habitat use but Lesser Redpoll were scarce or absent from upland conifer plantations in winter (Patterson et al., 1995).

8. Effects of forest management and deer on habitat The principal habitat of Lesser Redpoll seems to be young spruce plantations, although there is little information on more detailed requirements. It seems likely

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that clearfell and replanting in upland spruce forests is beneficial for Lesser Redpoll populations. The planting or encouragement of tree species whose seed provide food in winter (e.g. birch, Larch, Alder) may also be beneficial.

It is unlikely that deer will have a large effect on Lesser Redpoll populations unless unfenced spruce plantation are subjected to browsing intensities high enough to cause crop failure.

9. References Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Cramp, S., Perrins, C.M. & Brooks, D.J. (1994) Handbook of the birds of Europe, the Middle East and North Africa. Volume 8: Crows to Finches., Oxford: Oxford University Press. Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Hogstad, O. 2005. Numerical and functional responses of breeding passerine species to mass occurrence of geometrid caterpillars in a subalpine birch forest: a 30- year study. Ibis 147: 77-91. Lavers, C., McCullagh, M.J. & Fuller, R.J. 2005. Combining archive territory mapping data and aerial photography to investigate bird–habitat relationships: a case study from the Lincolnshire coast. Bird Study 52: 314-322. Moss, D., Taylor, P.N. & Easterbee, N. 1979. The effects on song-bird populations of upland afforestation with spruce. Forestry 52: 129-150. Patterson, I.J., Ollason, J. G. & Doyle, P. 1995. Bird populations in upland spruce plantations in northern Britain. Forest Ecology and Management 79: 107-131. Smart, J., Taylor, E., Amar, A., Smith, K., Bierman, S., Carpenter, J., Grice, P., Currie, F. & Hewson, C. 2007. Habitat associations of woodland birds: implications for woodland management for declining species. In RSPB Research Report No., Vol. 26. Sandy: RSPB. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Smith, K.W., Burges, D.J. & Parks, R.A. 1992. Breeding bird communities of broadleaved plantation and ancient pasture woodlands of the New Forest. Bird Study 39: 132-141. Sweeney, O.F.M., Wilson, M.W., Irwin, S., Kelly, T.C. & O'Halloran, J. 2010. Breeding bird communities of second-rotation plantations at different stages of the forest cycle. Bird Study 57: 301-314.

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Final report WC0793/CR0485 (29.04.14)

Bullfinch Pyrrhula pyrrhula

1. Nest site selection/requirements Bullfinch build a nest of twigs, lined with dry grass and roots (Cramp et al., 1994). Nests are located in dense scrub, hedges or on conifer branches, often 1-2 m from the ground (Bijlsma, 1982, Greig-Smith and Wilson, 1984).

2. Territory selection/attributes Bullfinches do not defend territories and may move many kilometres between nesting attempts to track localised food supply (Newton, 2000).

3. Main food items Diet is predominantly seeds but when in short supply in late winter and early spring they feed on tree buds and catkins (Newton, 1967, Greig-Smith and Wilson, 1984, Marquiss, 2007). A small amount of invertebrate food is taken during summer (Newton, 1967). Seeds taken are from a wide variety of herbs, shrubs and trees depending on local availability (Newton, 1967, Marquiss, 2007). 40-50 plant species have been recorded in the diet throughout the year and 5-15 species in any one month (Newton, 1967, Marquiss, 2007). Important seed resources in winter are Ash in southern England, heathers in upland Scotland, and Rowan in continental Europe (Newton, 1967, Marquiss, 2007, Munilla and Guitian, 2012, Fox et al., 2009). In spring, elm seeds are eaten in addition to buds and flowers (Bryant, 2011). Preference for seeds and buds is related to palatability and varies between species, varieties and individual trees depending on chemical content (Greig-Smith, 1985, Greig-Smith and Wilson, 1985).

4. Foraging methods Bullfinches rarely feed on the ground, and usually forage from a perch on a tree, shrub or herbaceous seed head (Newton, 1967). When feeding in open habitats they mainly feed close to cover and only move more than 10 m from cover when closer food sources are depleted (Newton, 1967, Greig-Smith and Wilson, 1984, Marquiss, 2007).

5. Breeding habitat Bullfinches breed in a variety of wooded and non-wooded habitats including scrub, farmland, orchards, gardens, broadleaved woodland and conifer woodland (Moss et al., 1979, Greig-Smith and Wilson, 1984, Bibby et al., 1985, Hinsley et al., 1995, Wilson et al., 2006, Sweeney et al., 2010, Gregory and Baillie, 1998). At the UK scale, the two broad habitats most used are farmland and woodland, within these there was a strong preference for scrub, broadleaved and mixed woodland and a relative avoidance of heath, grass and arable farmland (Gregory and Baillie, 1998). Woodlands with an intermediate understory growth stage have a higher chance of occupation, particularly open thicket stage (Hewson et al., 2011). Bullfinch is a scarce breeding species in conifer forests, when found they mostly occupy younger age classes (Bibby et al., 1985, Sweeney et al., 2010, Wilson et al., 2006). However, one study in the UK found only older stands occupied and the Scandinavian race of bullfinch mostly occurs in older stands of conifers (Moss et al., 1979, Haila et al.,

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1996, Gjerde and Saetersdal, 1997, Jokimäki and Solonen, 2011). In temperate continental Europe, Bullfinches occur more frequently in younger stages of forest growth and within mature forest they occur more frequently in conifers than broadleaves (Baguette et al., 1994, Paquet et al., 2006). Within broadleaved woodland, Bullfinches occur in woodlands with denser shrub layers (Hinsley et al., 1995, Carpenter et al., 2009). They also have a higher abundance in woodland with birch canopy and a lower abundance in oak woodland (Carpenter et al., 2009). In farmland habitat the presence of dense scrub patches and hedgerows are important (Greig-Smith and Wilson, 1984, Hinsley et al., 1995).

6. Post fledging behaviour/requirements No information was found.

7. Seasonal differences in habitat for resident species Bullfinches move widely between habitats even within seasons, with movement of up to 30 km recorded within a breeding season (Greig-Smith and Wilson, 1984, Newton, 2000, Marquiss, 2007). Types of habitat used track food resources, and vary between regions depending on types of food utilised. In an area of mixed farmland, orchard and woodland in southeast England non-wooded habitats were used most frequently during October to January and during the breeding season (Greig-Smith and Wilson, 1984). In northern Scotland, moorland habitats were only used in winter while farmland and scrub habitats were used most during autumn (Marquiss, 2007).

8. Effects of forest management and deer on habitat Within woodland Bullfinches require dense shrub cover for nesting, and diverse tree, shrub and ground flora to provide a range of seed food throughout the year. Management which can provide the dense shrub layer needed for nesting includes clearfell and coppicing. Any management which reduces shrub cover, such as cleaning during thinning, will be detrimental. Planting a range of trees which provide different seed sources will also be beneficial for Bullfinches.

Deer browsing can reduce dense shrub cover making woodland less suitable for nesting and may also reduce diversity of the shrub and field layer affecting food supply. The change from a herbaceous field layer to one dominated by grasses and sedges may also have an impact on food supplies.

9. References Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Bijlsma, R.G. 1982. Breeding season, clutch size and breeding success in the Bullfinch Pyrrhula pyrrhula. Ardea 70: 25-30. Bryant, D. 2011. Multi-species groups of finches feeding on Wych Elm fruits in spring. Scottish Birds 31: 311-314.

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Carpenter, J., Charman, E., Smart, J., Amar, A., Gruar, D. & Grice, P. 2009. Habitat associations of woodland birds II. In RSPB Research Report. Vol. 36. Cramp, S., Perrins, C.M. & Brooks, D.J. (1994) Handbook of the birds of Europe, the Middle East and North Africa. Volume 8: Crows to Finches., Oxford: Oxford University Press. Fox, A.D., Kobro, S., Lehikoinen, A., Lyngs, P. & Vaisanen, R. A. 2009. Northern Bullfinch Pyrrhula p. pyrrhula irruptive behaviour linked to rowanberry Sorbus aucuparia abundance. Ornis Fennica 86: 51-60. Gjerde, I. & Saetersdal, M. 1997. Effects on avian diversity of introducing spruce Picea spp. plantations in the native pine Pinus sylvestris forests of western Norway. Biological Conservation 79: 241-250. Gregory, R.D. & Baillie, S.R. 1998. Large-scale habitat use of some declining British birds. Journal of Applied Ecology 35: 785-799. Greig-Smith, P.W. 1985. The importance of flavour in determining the feeding preferences of Bullfinches (Pyrrhula pyrrhula) for the buds of two pear cultivars. Journal of Applied Ecology 22: 29-37. Greig-Smith, P.W. & Wilson, G.M. 1984. Patterns of activity and habitat use by a population of Bullfinches (Pyrrhula pyrrhula) in relation to bud-feeding in orchards. Journal of Applied Ecology 21: 401-422. Greig-Smith, P.W. & Wilson, M.F. 1985. Influences of seed size, nutrient composition and phenolic content on the preferences of Bullfinches feeding in ash trees. Oikos 44: 47-54. Haila, Y., Nicholls, A.O., Hanski, I.K. & Raivio, S. 1996. Stochasticity in bird habitat selection: Year-to-year changes in territory locations in a boreal forest bird assemblage. Oikos 76: 536-552. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1995. Habitat and landscape factors influencing the presence of individual breeding bird species in woodland fragments. Journal of Avian Biology 26: 94-104. Jokimäki, J. & Solonen, T. 2011. Habitat associations of old forest bird species in managed boreal forests characterized by forest inventory data. Ornis Fennica 88: 57-70. Marquiss, M. 2007. Seasonal pattern in hawk predation on common Bullfinches Pyrrhula pyrrhula: evidence of an interaction with habitat affecting food availability. Bird Study 54: 1-11. Moss, D., Taylor, P.N. & Easterbee, N. 1979. The effects on song-bird populations of upland afforestation with spruce. Forestry 52: 129-150. Munilla, I. & Guitian, J. 2012. Numerical response of Bullfinches Pyrrhula pyrrhula to winter seed abundance. Ornis Fennica 89: 197-205. Newton, I. 1967. The Feeding Ecology of the Bullfinch (Pyrrhula pyrrhula L.) in Southern England. Journal of Animal Ecology 36: 721-744. Newton, I. 2000. Movements of Bullfinches Pyrrhula pyrrhula within the breeding season. Bird Study 47: 372-376. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture-

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dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Sweeney, O.F.M., Wilson, M.W., Irwin, S., Kelly, T.C. & O'Halloran, J. 2010. Breeding bird communities of second-rotation plantations at different stages of the forest cycle. Bird Study 57: 301-314. Wilson, M.W., Pithon, J., Gittings, T., Kelly, T.C., Giller, P.S. & O'Halloran, J. 2006. Effects of growth stage and tree species composition on breeding bird assemblages of plantation forests. Bird Study 53: 225-236.

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Final report WC0793/CR0485 (29.04.14)

Hole nesters and species nesting in the tree canopy.

Lesser Spotted Woodpecker Dendrocopos minor

1. Nest site selection/requirements Lesser Spotted Woodpeckers nest in cavities with a nest entrance diameter of 3-3.5 cm which are excavated by both parents, often in cavities with some previous excavation by other woodpeckers (Charman et al., 2012, Winkler and Christie, 2002). Cavities are most often located in a branch off the main trunk, or the trunk itself, mainly in dead trees, dead stumps, or in dead branches of live trees (Charman et al., 2012, Smith, 2007, Yamauchi et al., 1997, Kosiński and Kempa, 2007, Kawada, 1980, Glue and Boswell, 1994). Branch cavities are usually located on the underside, often against the main trunk, with branch diameter ranging from 12-50 cm with averages of 20 and 24 cm recorded (Charman et al., 2012, Glue and Boswell, 1994). Nest cavities are most often located in the upper half of selected trees (Charman et al., 2012, Kosiński and Kempa, 2007) at heights ranging from 2-22 m across studies with mean heights of 14.9 and 12.0 m recorded, nest height was related to tree species and height of tree selected (Kosiński and Kempa, 2007, Glue and Boswell, 1994, Charman et al., 2012). Mean height of nest tree was 14.9 m in England (Charman et al., 2012) and 20 m in Poland (Kosiński and Kempa, 2007) and tree DBH ranged from 15-73 cm, averaging 38 cm in England (Charman et al., 2012) and 48 cm in Poland (Kosiński and Kempa, 2007). A wide selection of tree species are used for locating nest sites with birch, Alder, willow, oak and fruit trees frequently selected species (Glue and Boswell, 1994, Charman et al., 2012) with oak being the most commonly selected in England (Charman et al., 2012). Nest substrate is dead material (Charman et al., 2012).

2. Territory selection/attributes Breeding territory size is up to 43 ha, with winter ranges of 742 ha gradually becoming smaller from early spring until much smaller breeding territory is established (Wiktander et al., 2001). In England territories are situated in oak dominated, mature woodlands, with an open understorey and low herbaceous field layer (Charman et al., 2010, Charman et al., 2012).

In Sweden, pre-breeding season territories are influenced by availability of two key prey species, Argyresthia goedarthella which occurs on birch in some years and Alder in other years, and Stenostola dubia which occurs on lime trees, especially where they are more abundant (Olsson et al., 2001).

3. Main food items For most of the year diet is predominantly invertebrate larvae under bark, but switches to surface-living invertebrates during the breeding season from the onset of spring tree bud burst. A wide variety of taxa are taken. In summer diet consists mostly of aphids, caterpillars, ants, beetles, and other surface-dwelling arthropods including Diptera and spiders (Török, 1990, Wiktander et al., 1994). Aphids and caterpillars are frequently fed to nestlings; in Germany 52% of food items were aphids and 25% caterpillars (Rossmanith et al., 2007), in Hungary 51% aphids and

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35% caterpillars with 85% dry weight of nestling food being caterpillars (Török, 1990). Craneflies and wood living larvae are also important in nestling diet (Rossmanith et al., 2007, Török, 1990). The proportion of caterpillars in chick diet declines seasonally and is replaced mainly by wood-boring larvae (Rossmanith et al., 2007). The larval stage of two invertebrates is particularly important in Lesser Spotted Woodpecker diet. The moth Argyresthia goedartella is available cyclically on birch or Alder and in Norway breeding is synchronised with their availability (Selas et al., 2008), and from April are a significant food source in Sweden (Olsson et al., 1999). Larvae of the beetle Stenostola dubia is an important prey item found on lime trees (Olsson et al., 2001).

In winter diet is mainly wood-boring invertebrate larvae (mainly Buprestidae, Cerambycidae and Curculionidae) and invertebrates living under bark (mainly Scolytidae and Ipidae) (Winkler and Christie, 2002). Females have a wider foraging niche than males in winter and pre-breeding (Hogstad, 2010).

4. Foraging methods In England 80% of foraging is recorded in oak (Charman et al., 2012) and in Norway foraging by both sexes was predominantly in birch and Grey alder, while Scots pine was avoided relative to availability (Hogstad, 2010). Two important prey items are host tree specific, and cyclical, therefore preferred tree species for foraging are likely to vary between years (Olsson et al., 2001).

Foraging activity by English Lesser Spotted Woodpeckers in the pre-breeding season is exclusively on tree branches rather than trunks, and predominantly in live trees (98%) (Charman et al., 2012). Continental studies also record foraging mostly on branches rather than trunks (Olsson et al., 1999). In Norway foraging differs seasonally and between sexes with females having a wider foraging niche outside the main breeding season. In winter females foraged more than males on dead wood and more on live parts of trees in the breeding season. Females tend to feed on smaller diameter and higher trees, although there is no difference between sexes in foraging height (Hogstad, 2010). Males and females usually forage alone (Hogstad, 2009, Hogstad, 2010) and in winter forage later rather than earlier in the day (Olsson et al., 2000).

In England foraging methods tend to be by gleaning from branch surfaces in the upper third of trees, and mainly in the outer part of the tree canopy, and also include pecking and bark-scaling (Charman et al., 2012). In the same study females probed more than males during the winter and pre-breeding periods and gleaned more outside the winter, while males used bark-scaling and pecking more than females year round.

5. Breeding habitat Distribution is affected by variables reflecting both relative positions of woodlands within the landscape and forest structure. Lesser Spotted Woodpeckers mostly occur at lower elevations (Miranda and Pasinelli, 2001), below 274 m in the UK (Glue and Boswell, 1994) and in larger blocks of woodland, particularly woodlands >38 ha

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(Wiktander et al., 1992). In the UK, Lesser Spotted Woodpeckers are associated with mature oak woodland with abundant dead wood (Charman et al., 2010). In continental studies, Lesser Spotted Woodpeckers occupy mature diverse stands with many snags, a high density of dead trees and avoid mixed softwood/deciduous stands (Olsson et al., 1992, Delahaye et al., 2002, Delahaye et al., 2010). In Belgium occurrence is more likely where cover of oak is high and in woodland with a selection of younger stands; average snag and dead wood volume where they occurred was 9m3/ha (Delahaye et al., 2010).

Occurrence in the UK is influenced by proximity to other woodland, being more common within more heavily wooded landscapes (Charman et al., 2010, Broughton et al., 2013). In Poland they are only found in landscapes types without intensively managed woodland (Angelstam et al., 2002) and in Sweden require larger areas in managed woodland compared to natural woodland (Wiktander et al., 1992).

6. Post fledging behaviour/requirements Female Lesser Spotted Woodpeckers frequently cease feeding older nestlings, but resume feeding after fledging (Wiktander et al., 2003). Eruptive autumn dispersal, especially by juveniles, is detected in continental populations (Gohli et al., 2011, Winkler and Christie, 2002).

7. Seasonal differences in habitat for resident species Lesser Spotted Woodpeckers show no seasonal differences in habitat although diet and foraging methods differ seasonally, see above.

8. Effects of forest management and deer on habitat Key habitat requirements for Lesser Spotted Woodpecker are mature woodland with abundant dead wood. Lesser Spotted Woodpeckers are largely absent from intensively managed woodlands, and landscapes which predominantly consist of managed woodland. Lesser Spotted Woodpeckers are more likely to be abundant in small scale harvesting systems such as continuous cover forestry, group fell and selective felling. Dead branches on live trees are important for nesting and so presence in active coppice and recently thinned stands is less likely.

Lesser Spotted Woodpeckers make little use of the understory and so are unlikely to be affected by high deer browsing. Lesser Spotted Woodpeckers forage above deer browsing height so foraging habitat is unlikely to be affected.

9. References Angelstam, P., Breuss, M., Mikusinski, G., Stenstrom, M., Stighall, K. & Thorell, D. 2002. Effects of forest structure on the presence of woodpeckers with different specialisation in a landscape history gradient in NE Poland. In Avian Landscape Ecology: Pure and Applied Issues in the Large-Scale Ecology of Birds. (Eds. D. Chamberlain & A. Wilson), pp. 25-38. Broughton, R.K., Hill, R.A. & Hinsley, S.A. 2013. Relationships between patterns of habitat cover and the historical distribution of the Marsh Tit, Willow Tit and Lesser Spotted Woodpecker in Britain. Ecological Informatics 14: 25-30.

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Charman, E.C., Smith, K.W., Dodd, S., Gruar, D.J. & Dillon, I.A. 2012. Pre-breeding foraging and nest site habitat selection by Lesser Spotted Woodpeckers Dendrocopos minor in mature woodland blocks in England. Ornis Fennica 89: 182- 196. Charman, E.C., Smith, K.W., Gruar, D.J., Dodd, S. & Grice, P.V. 2010. Characteristics of woods used recently and historically by Lesser Spotted Woodpeckers Dendrocopos minor in England. Ibis 152: 543-555. Delahaye, L., Derouaux, A. & Delvingt, W. 2002. Habitat modeling: a tool for forest ecology management. A case study of lesser spotted woodpecker (Dendrocopos minor) in Belgian Ardenne. Aves (Liege) 39: 129-143. Delahaye, L., Monticelli, D., Lehaire, F., Rondeux, J. & Claessens, H. 2010. Fine-scale habitat selection by two specialist woodpeckers occurring in beech and oak- dominated forests in southern Belgium. Ardeola 57. Glue, D.E. & Boswell, T. 1994. Comparative nesting ecology of the three British breeding woodpeckers. British Birds 87: 253-269. Gohli, J., Røer, J.E., Selås, V., Stenberg, I. & Lislevand, T. 2011. Migrating Lesser Spotted Woodpeckers Dendrocopos minor along the coast of southern Norway: where do they come from? Ornis Fennica 88: 00-00. Hogstad, O. 2009. Sexual differences of labour during breeding activities and territory use in the lesser spotted woodpecker Dendrocopos minor. Ornis Norvegica 32: 42- 48. Hogstad, O. 2010. Sexual differences in foraging behaviour in the Lesser Spotted Woodpecker Dendrocopos minor. Ornis Norvegica 33: 135-146. Kawada, M. 1980. Breeding biology of Dendrocopos major japonicus and Dendrocopos minor in Obihiro, Hokkaido, Japan. Journal of the Yamashina Institute for Ornithology 12: 48-70. Kosiński, Z. & Kempa, M. 2007. Density, distribution and nest-sites of woodpeckers picidae, in a managed forest of Western Poland. Polish Journal of Ecology 55: 519- 533. Miranda, B. & Pasinelli, G. 2001. Habitat requirements of the Lesser Spotted Woodpecker (Dendrocopos minor) in forests of northeastern Switzerland. Journal Fur Ornithologie 142: 295-305. Olsson, O., Nilsson, I.N., Nilsson, S.G., Pettersson, B., Stagen, A. & Wiktander, U. 1992. Habitat preferences of the Lesser spotted woodpecker Dendrocopos minor. Ornis Fennica 69: 119-125. Olsson, O., Wiktander, U., Holmgren, N.M.A. & Nilsson, S.G. 1999. Gaining ecological information about Bayesian foragers through their behaviour. II. A field test with woodpeckers. Oikos 87: 264-276. Olsson, O., Wiktander, U., Malmqvist, A. & Nilsson, S. G. 2001. Variability of patch type preferences in relation to resource availability and breeding success in a bird. Oecologia 127: 435-443. Olsson, O., Wiktander, U. & Nilsson, S. 2000. Daily foraging routines and feeding effort of a small bird feeding on a predictable resource. Proceedings of the Royal Society of London. Series B: Biological Sciences 267: 1457-1461. Rossmanith, E., Höntsch, K., Blaum, N. & Jeltsch, F. 2007. Reproductive success and nestling diet in the Lesser Spotted Woodpecker (Picoides minor): the early bird gets the caterpillar. Journal of Ornithology 148: 323-332.

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Selas, V., Steen, R., Kobro, S., Lislevand, T. & Stenberg, I. 2008. Direct and indirect weather impacts on spring populations of lesser spotted woodpecker (Dendrocopos minor) in Norway. Scandinavian Journal of Forest Research 23: 148- 153. Smith, K.W. 2007. The utilization of dead wood resources by woodpeckers in Britain. Ibis 149: 183-192. Török, J. 1990. Resource partitioning amoung three woodpecker species Dendrocopos spp. during the breeding season. Holarctic Ecology 13: 257-264. Wiktander, U., Nilsson, I.N., Nilsson, S.G., Olsson, O., Pettersson, B. & Stagen, A. 1992. Occurance of the Lesser spotted woodpecker Dendrocopos minor in relation to area of deciduous forest. Ornis Fennica 69: 113-118. Wiktander, U., Nilsson, S.G., Olsson, O. & Stagen, A. 1994. Breeding success of a Lesser Spotted Woodpecker Dendrocopos minor population. Ibis 136: 318-322. Wiktander, U., Olsson, O. & Nilsson, S.G. 2001. Seasonal variation in home-range size, and habitat area requirement of the lesser spotted woodpecker (Dendrocopos minor) in southern Sweden. Biological Conservation 100: 387-395. Wiktander, U., Olsson, O. & Nilsson, S.G. 2003. Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor. Journal of Avian Biology 31: 447-456. Winkler, H. & Christie, A. 2002. Family Picidae (Woodpeckers). In Handbook of Birds of the World Vol. 7 Jacamars to Woodpeckers. (Eds. J. del Hoyo, A. Elliott & J. Sargatal), Vol. 7, pp. 477. Barcelona: Lynx Edicions. Yamauchi, K., Yamazaki, S. & Fujimaki, Y. 1997. Breeding habitats of Dendrocopos major and D. minor in urban and rural areas. Japanese Journal of Ornithology 46: 121-131.

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Final report WC0793/CR0485 (29.04.14)

Willow Tit Poecile montanus

1. Nest site selection/requirements Willow Tits excavate nest holes in dead wood, often narrow diameter trunks of dead trees (Lewis et al., 2009a). Nest holes are mostly <1.5 m from the ground (range 0.15 – 10 m) in willow, birch and elder trees located in scrub dominated by willow, birch and hawthorn (Stewart, 2010). Nests are lined with shredded bark, plant fibres, fur and feathers (Lewis et al., 2009a).

2. Territory selection/attributes Willow Tits are sedentary, remaining on territories year round (Lewis et al., 2009a, Cramp and Perrins, 1993). Little is known of territory size or requirements in the UK. However, Boreal Willow Tits have a median winter home range of 12.6 ha (2.3-34.3 ha), which contain 7.1 ha (2.3-14.3 ha) of forest (Siffczyk et al., 2003). These winter ranges are not exclusive and neighbouring home ranges overlap (Haftorn, 1999).

3. Main food items A review of 13 studies of nestling diet studies identified the main food items as caterpillars and spiders, taken from both broadleaved and conifer trees, with flies, beetles and bugs also of importance in nestling diet in some studies (Gibb and Betts, 1963, Cholewa and Wesolowski, 2011). Proportions of different groups of invertebrates in Willow Tit diet vary seasonally, with early broods feeding more flies and late broods feeding more spiders (Mattes et al., 1996).

Adults also largely feed on invertebrates but also consume seeds during autumn and early winter (Cramp and Perrins, 1993, Brodin, 1994). Seeds of pine, spruce, Juniper, hemp nettles and woundworts are frequently recorded Willow Tit diet (Brodin, 1994, Hughes and Dougharty, 1978).

4. Foraging methods All information on foraging comes from the Boreal and continental subspecies foraging in conifer forests during winter. Here Willow Tits forage almost entirely in the canopy of trees, rarely feeding in shrubs or on the ground (Ulfstrand, 1976, Hogstad, 1978). They forage in all tree species present, with a preference for spruce, pine or birch dependant on study (Ulfstrand, 1976, Hogstad, 1978). Within the canopy Willow Tits prefer foraging on branches near to the trunk and lower in the canopy, with feeding in needles and twigs of outer branches less preferred (Ulfstrand, 1976, Hogstad, 1978, Krams, 1996, Krams et al., 2001, Lens, 1996). Location of foraging is affected more by predation risk and competition than food abundance (Lens, 1996, Krams, 1996, Krams et al., 2001).

Within the UK, Willow Tits mainly inhabit scrub habitats and are scarce in conifer forests, therefore detail of where in the canopy they forage is not relevant. However the principles of foraging in areas with low predation risk and with a lack of competitors are relevant and they are likely to forage in denser areas of scrub.

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5. Breeding habitat In the UK, Willow Tits are most abundant in wet habitats compared with woodland or farmland (Siriwardena, 2004). They are primarily found in scrub habitats, often former industrial sites which have high soil moisture content, and are scarce in woodland habitats with a mature tree canopy (Lewis et al., 2009a, Lewis et al., 2007, Lewis et al., 2009b, Stewart, 2010, Probert, 2011). Such former industrial sites are prone to habitat change through human development or succession of scrub habitats to woodland. Within woodland, Willow Tits are more likely to occur in areas with a dense understory (2-4 m high), low tree cover and greater amounts of dead wood (Probert, 2011, Stewart, 2010, Lewis et al., 2009b). Unlike continental races of Willow Tit, UK Willow Tits are scarce in conifer forest, with occupied woodlands containing fewer conifers than unoccupied woodlands (Gibb and Betts, 1963, Currie and Bamford, 1981, Currie and Bamford, 1982, Bibby et al., 1985, Stewart, 2010). Willow Tits are more likely to occur and are more stable in landscapes with greater woodland cover (Broughton et al., 2013).

Continental races of Willow Tit are primarily a species of conifer forests, where they mainly occur in mixed spruce, pine and birch forest. They are most abundant in mature forest >80 years old and younger clearcuts 5-30 years old, with few in younger clearcuts or in dense mid-age stands (Hansson, 1983, Baguette et al., 1994, Gjerde and Saetersdal, 1997, Mortberg, 2001, Rodriguez et al., 2001, Siffczyk et al., 2003, Paquet et al., 2006). Young forests are more likely to be used when they have a higher broadleaved component (Rodriguez et al., 2001).

6. Post fledging behaviour/requirements Young are dependent on their parents for 7-11 days after fledging, and remain in a loose family group on the natal territory for a further 1-2 weeks (Haftorn, 1997, Lewis et al., 2009a). After dispersal from their natal territory juvenile Willow Tits wander for several weeks before most settle on winter territories within six weeks of fledging, during late July to early September (Haftorn, 1997, Hogstad, 1990, Haftorn, 1999). During winter, juveniles continue to range more widely than adults visiting neighbouring territories looking for opportunities for breeding territories (Hogstad, 1999).

7. Seasonal differences in habitat for resident species Willow Tits are resident and remain on the same territory throughout the year (Cramp and Perrins, 1993). In winter home ranges are large and overlapping (Siffczyk et al., 2003).

8. Effects of forest management and deer on habitat Habitat used by Willow Tits in the UK are not managed forest habitats, therefore management within existing forest is unlikely to affect Willow Tits. However, they require wet scrubby secondary woodland and management of these habitats for forestry is likely to be detrimental. Drainage, planting with timber trees and clearance of scrub or dead wood are all likely to be detrimental to Willow Tit habitat. The main beneficial management would be to allow development of natural scrub on wet areas of woodland, retain dead wood for nesting and remove larger trees.

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Willow Tit habitat has a dense understory or shrub layer above 2 m, and less dense below 2 m, therefore deer browsing which mostly affects vegetation below 2 m, may only have a small short term negative effect on Willow Tit habitat. However, longer term heavy browsing can reduce regeneration and alter woodland structure leading to a reduction in availability of scrub habitats required by Willow Tits. A UK scale study found Willow Tit abundance and populations negatively affected by deer density, particularly by Roe deer (Newson et al., 2012).

9. References Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Bibby, C.J., Phillips, B.N. & Seddon, A.J.E. 1985. Birds of restocked conifer plantations in Wales. Journal of Applied Ecology 22: 619-633. Brodin, A. 1994. The role of naturally stored food supplies in the winter diet of the boreal willow tit Parus montanus. Ornis Svecica 4: 31-40. Broughton, R.K., Hill, R.A. & Hinsley, S.A. 2013. Relationships between patterns of habitat cover and the historical distribution of the Marsh Tit, Willow Tit and Lesser Spotted Woodpecker in Britain. Ecological Informatics 14: 25-30. Cholewa, M. & Wesolowski, T. 2011. Nestling food of European hole-nesting passerines: do we know enough to test the adaptive hypotheses on breeding seasons? Acta Ornithologica 46: 105-116. Cramp, S. & Perrins, C. M. (1993) Handbook of the birds of Europe, the Middle East and North Africa: The birds of the Western Palearctic. Vol. VII Flycatchers to Shrikes, Oxford: Oxford University Press. Currie, F.A. & Bamford, R. 1981. Bird populations of sample pre-thicket forest plantations. Quarterly Journal of Forestry 75: 75-82. Currie, F.A. & Bamford, R. 1982. The value to birdlife of retaining small conifer stands beyond normal felling age within forests. Quarterly Journal of Forestry 76: 153- 159. Gibb, J.A. & Betts, M.A. 1963. Food and food supply of nestling Tits (Paridae) in Breckland Pine. Journal of Animal Ecology 32: 489-533. Gjerde, I. & Saetersdal, M. 1997. Effects on avian diversity of introducing spruce Picea spp. plantations in the native pine Pinus sylvestris forests of western Norway. Biological Conservation 79: 241-250. Haftorn, S. 1997. Natal dispersal and winter flock formation in the willow tit Parus montanus. Fauna Norvegica Series C Cinclus 20: 17-35. Haftorn, S. 1999. Flock formation, flock size and flock persistence in the Willow Tit Parus montanus. Ornis Fennica 76: 49-63. Hansson, L. 1983. Bird numbers across edges between mature conifer forest and clearcuts in central Sweden. Ornis Scandinavica 14: 97-103. Hogstad, O. 1978. Differentiation of foraging niche among tits, Parus spp., in Norway during winter. Ibis 120: 139-146. Hogstad, O. 1990. Dispersal date and settlement of juvenile willow tits Parus montanus in winter flocks. Fauna Norvegica Series C Cinclus 13: 49-55. Hogstad, O. 1999. Territory acquisition during winter by juvenile Willow Tits Parus montanus. Ibis 141: 615-620.

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Hughes, S.W.M. & Dougharty, F.W. 1978. Vegetable food of willow tit. British Birds 71: 465. Krams, I.A. 1996. Predation risk and shifts of foraging sites in mixed willow and crested tit flocks. Journal of Avian Biology 27: 153-156. Krams, I.A., Krams, T. & Cernihovics, J. 2001. Selection of foraging sites in mixed Willow and Crested Tit flocks: rank-dependent survival strategies. Ornis Fennica 78: 1-11. Lens, L. 1996. Wind stress affects foraging site competition between crested tits and willow tits. Journal of Avian Biology: 41-46. Lewis, A.J.G., Amar, A., Charman, E.C. & Stewart, F.R.P. 2009a. The decline of the willow tit in Britain. British Birds 102: 386-393. Lewis, A.J.G., Amar, A., Cordi-Piec, D. & Thewlis, R.M. 2007. Factors influencing Willow Tit Poecile montanus site occupancy: a comparison of abandoned and occupied woods. Ibis 149: 205-213. Lewis, A.J.G., Amar, A., Daniells, L., Charman, E.C., Grice, P. & Smith, K. 2009b. Factors influencing patch occupancy and within-patch habitat use in an apparently stable population of Willow Tits Poecile montanus kleinschmidti in Britain. Bird Study 56: 326-337. Mattes, H., Tumbrinck, J. & Fischbacher, M. 1996. Nestling food of tits (Parus major, P. ater, P. montanus, P. cristatus) in a subalpine larici-cembretum of the Upper Engadine, Switzerland. Ornithologische Beobachter 93: 293-314. Mortberg, U.M. 2001. Resident bird species in urban forest remnants; landscape and habitat perspectives. Landscape Ecology 16: 193-203. Newson, S.E., Johnston, A., Renwick, A.R., Baillie, S.R. & Fuller, R.J. 2012. Modelling large-scale relationships between changes in woodland deer and bird populations. Journal of Applied Ecology 49: 278-286. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management, 227: 59-70. Probert, E. 2011. Habitat selection of Willow tits Poecile montanus kleinschmidt in a post-industrial landscape. Vol. MSc., pp. 13. Manchester: Manchester Metropolitan University. Rodriguez, A., Andren, H. & Jansson, G. 2001. Habitat-mediated predation risk and decision making of small birds at forest edges. Oikos 95: 383-396. Siffczyk, C., Brotons, L., Kangas, K. & Orell, M. 2003. Home range size of willow tits: a response to winter habitat loss. Oecologia 136: 635-642. Siriwardena, G.M. 2004. Possible roles of habitat, competition and avian nest predation in the decline of the Willow Tit Parus montanus in Britain. Bird Study, 51: 193-202. Stewart, F.R.P. 2010. Ecology and conservation of the Willow tit Poecile montanus in Britain. Vol. PhD., pp. 130. Nottingham: Nottingham University. Ulfstrand, S. 1976. Feeding niches of some passerine birds in a south Swedish coniferous plantation in winter and summer. Ornis Scandinavica 7: 21-27.

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Final report WC0793/CR0485 (29.04.14)

Marsh Tit Poecile palustris

1. Nest site selection/requirements Marsh Tits tend to select nest holes in the lower half of trees, mean heights recorded are 3.0 m and 5.0 m although height can be very variable with nests recorded from ground level to >10 m (Wesolowski, 2002, Broughton et al., 2011). The most common location for nests is within natural cavities in live trees, although a wide range of locations are used including nestboxes, holes in woody shrubs, cavities in tree bases, dead trees and old woodpecker holes (Wesolowski, 2002, Broughton et al., 2011). Tree species used for nesting depends on that available, although nesting in Ash is greater than expected and nesting in oak of lower frequency than expected based on availability (Wesolowski, 2002, Broughton et al., 2011). In Poland, mean entrance hole diameter was 31 mm with a mean cavity depth of 90 mm (Wesolowski, 2006). The size of entrance hole has no effect on predation rates of nests, however deeper cavities are less likely to be predated (Wesolowski, 2002). Nests in upward facing knot holes are more prone to failure due to flooding (Wesolowski et al., 2002), but location and nest cavity dimensions had no effect on infestation rates of blow-fly nest parasites (Wesolowski, 2001). Nest sites are located randomly within the territory for first year birds and within core of territories in older birds. Within the territory, nest sites are located within areas of taller Ash and Field maple with a sparse understory, while areas of oak are avoided (Broughton et al., 2012)

2. Territory selection/attributes Breeding territories are 2-8 ha in extent with a mean area of 4.1 ha (Broughton et al., 2006). Marsh Tits occupy territories in areas within woods that have the highest canopy height, and canopy cover, leaving the areas of lowest canopy cover unoccupied (Broughton et al., 2006, Hinsley et al., 2009). Where understory is sparse breeding territories are centred on areas of highest shrub density (Hinsley et al., 2007). There is no apparent selection for particular tree or shrub species, although in some studies there is a selection for greater shrub diversity but not all studies (Broughton et al., 2006, Hinsley et al., 2009, Hinsley et al., 2007). Marsh Tits have home ranges centred on the breeding territory all year, but range over wider areas during winter (Cramp and Perrins, 1993); P Bellamy & R Broughton unpublished data).

3. Main food items Diet is mainly small invertebrates of a wide range of taxa and seeds (Cramp and Perrins, 1993, Amann, 2007).

A review of 8 published studies of nestling diet identified caterpillars and spiders as the predominant food items, with bugs also important in 3 studies (Cholewa and Wesolowski, 2011).

During the breeding season diet is predominantly invertebrates, with the number of seeds eaten increasing in autumn and winter (Amann, 2007). Seeds of both herbaceous plants and shrubs are eaten (Amann, 2007, Morris, 1992).

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4. Foraging methods Foraging for invertebrates is mainly in the upper two thirds of trees, finding food by gleaning from bark and twigs and visual searching, although they also forage to a lesser extent on the ground and in shrubs, and by gleaning from leaves or by hanging and feeding (Hinsley et al., 2007, Amann, 2007). Fruit seeds are removed from the fruit pulp and taken to cover to remove seed coat before eating (Morris, 1992). When a source of abundant seed is found seeds are often cached, and are mostly retrieved within a few days (Stevens and Krebs, 1986).

5. Breeding habitat Marsh Tits are most likely to breed in larger woods, with a low probability of breeding in woodlands <10 ha, and are more likely to occur in landscapes with greater woodland cover (Hinsley et al., 1995, Hinsley et al., 1996, Broughton et al., 2013). Marsh Tits prefer broadleaved woodland but will breed in pine woodland, although are not recorded from woodland with >25 % spruce in Norway (Cramp and Perrins, 1993, Gjerde and Saetersdal, 1997). In a comparison across different forest types, Marsh Tits were more frequently encountered in mature oak and Beech forest than in mature conifers, young forests or forest edges (Paquet et al., 2006). However in a study of succession in Beech and spruce forests Marsh Tits were most frequent in younger Beech forest 1-30 years old and not present in stands >60years old (Baguette et al., 1994). A UK scale study found Marsh Tit occurrence more likely in woodlands at an older growth stage (Hewson et al., 2011). Woodlands occupied by Marsh Tits have higher tree cover or taller trees, a greater understory cover and higher shrub diversity compared to unoccupied woods (Smith et al., 1987, Carpenter et al., 2010). However another study found no apparent habitat selection (Smith et al., 1992).

6. Post fledging behaviour/requirements After fledging young remain in a family group and are dependent on parents for 11- 14 days (Nilsson and Smith, 1985, Broughton et al., 2010). Mortality during the dependent stage is 0.11-0.33% per day (Nilsson and Smith, 1985). Dispersal from natal territories occurs soon after independence and all dispersed within 14 days (Nilsson and Smith, 1985, Broughton et al., 2010). Most dispersal occurs within the two weeks of gaining independence with little further dispersal occurring until first breeding (Nilsson and Smith, 1988, Broughton et al., 2010). Dispersal distances are 0.5-5 km, representing 0-6 breeding territories crossed before first breeding (Nilsson and Smith, 1988, Broughton et al., 2010). Females disperse further than males (Nilsson and Smith, 1988).

7. Seasonal differences in habitat for resident species There is no difference in habitat between seasons, and individuals remain resident within home ranges centred on the breeding territory year round (Cramp and Perrins, 1993); P Bellamy & R Broughton unpublished data).

8. Effects of forest management and deer on habitat Key habitat requirements for Marsh Tit include large areas of mature woodland with a diverse shrub layer and potential nest holes. These requirements suggest

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unmanaged woodland, or woodland not managed for timber production, is likely to provide better quality habitat as trees with rot holes and understory shrubs are likely to be removed during thinning and timber harvesting through clearfell, while coppicing reduces quantities of mature woodland and fragments remaining areas. Harvesting systems such as group fell and selective felling, which removes timber at a smaller spatial scale and do not lead to large areas of even age plantation, are more likely to provide suitable habitat for Marsh Tits. Small scale disturbance from forestry operations along internal woodland edges and tracks can provide tall ruderal vegetation, including thistles and Burdock which provide a seed food source in winter.

High deer browsing has not been shown to affect Marsh Tit populations or habitat. However, long term impacts of high deer browsing could affect winter food supply though reducing diversity of the shrub layer and seed availability from shrubs and the field layer. Marsh Tits are known to feed on the seeds of several shrub species which have shown to be negatively affected by deer browsing (e.g. Bramble, Honeysuckle, Dogwood).

9. References Amann, F. 2007. Balzfuttern, Nahrung und Samenversteeken bei der Sumpfmeise Parus palustris. Ornithologishe Beobachter 104: 91-100. Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Broughton, R.K., Hill, R.A., Bellamy, P.E. & Hinsley, S. A. 2010. Dispersal, ranging and settling behaviour of Marsh Tits Poecile palustris in a fragmented landscape in lowland England. Bird Study 57: 458-472. Broughton, R.K., Hill, R.A., Bellamy, P.E. & Hinsley, S.A. 2011. Nest-sites, breeding failure, and causes of non-breeding in a population of British Marsh Tits Poecile palustris. Bird Study 58: 229-237. Broughton, R.K., Hill, R.A., Henderson, L.J., Bellamy, P.E. & Hinsley, S.A. 2012. Patterns of nest placement in a population of Marsh Tits Poecile palustris. Journal of Ornithology 153: 735-746. Broughton, R.K., Hill, R.A. & Hinsley, S.A. 2013. Relationships between patterns of habitat cover and the historical distribution of the Marsh Tit, Willow Tit and Lesser Spotted Woodpecker in Britain. Ecological Informatics 14: 25-30. Broughton, R.K., Hinsley, S.A., Bellamy, P.E., Hill, R.A. & Rothery, P. 2006. Marsh Tit Poecile palustris territories in a British broad-leaved wood. Ibis 148: 744-752. Carpenter, J., Smart, J., Amar, A., Gosler, A., Hinsley, S. & Charman, E. 2010. National- scale analyses of habitat associations of Marsh Tits Poecile palustris and Blue Tits Cyanistes caeruleus: two species with opposing population trends in Britain. Bird Study 57: 31-43. Cholewa, M. & Wesolowski, T. 2011. Nestling food of European hole-nesting passerines: do we know enough to test the adaptive hypotheses on breeding seasons? Acta Ornithologica 46: 105-116. Cramp, S. & Perrins, C.M. (1993) Handbook of the birds of Europe, the Middle East and North Africa: The birds of the Western Palearctic. Vol. VII Flycatchers to

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Shrikes, Oxford: Oxford University Press. Gjerde, I. & Saetersdal, M. 1997. Effects on avian diversity of introducing spruce Picea spp. plantations in the native pine Pinus sylvestris forests of western Norway. Biological Conservation 79: 241-250. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1995. Habitat and landscape factors influencing the presence of individual breeding bird species in woodland fragments. Journal of Avian Biology 26: 94-104. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1996. Influences of population size and woodland area on bird species distributions in small woods. Oecologia 105: 100-106. Hinsley, S.A., Carpenter, J.E., Broughton, R.K., Bellamy, P.E., Rothery, P., Amar, A., Hewson, C.M. & Gosler, A.G. 2007. Habitat selection by Marsh Tits Poecile palustris in the UK. Ibis 149: 224-233. Hinsley, S.A., Hill, R.A., Fuller, R.J., Bellamy, P.E. & Rothery, P. 2009. Bird species distributions across woodland canopy structure gradients. Community Ecology 10: 99-110. Morris, P.I. 1992. Feeding behaviour of Marsh tit. British Birds 85: 313-314. Nilsson, J.A. & Smith, H.G. 1985. Early Fledgling Mortality and the Timing of Juvenile Dispersal in the Marsh Tit Parus palustris. Ornis Scandinavica 16: 293-298. Nilsson, J.A. & Smith, H.G. 1988. Effects of Dispersal Date on Winter Flock Establishment and Social Dominance in Marsh Tits Parus palustris. Journal of Animal Ecology 57: 917-928. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Smith, K.W., Burges, D.J. & Parks, R.A. 1992. Breeding bird communities of broadleaved plantation and ancient pasture woodlands of the New Forest. Bird Study 39: 132-141. Stevens, T.A. & Krebs, J.R. 1986. Retrieval of stored seeds by marsh tits Parus palustris in the field. Ibis 128: 513-525. Wesolowski, T. 2001. Host-parasite interactions in natural holes: marsh tits (Parus palustris) and blow flies (Protocalliphora falcozi). Journal of Zoology 255: 495-503. Wesolowski, T. 2002. Anti-predator adaptations in nesting Marsh Tits Parus palustris: the role of nest-site security. Ibis 144: 593-601. Wesolowski, T. 2006. Nest-site re-use: Marsh Tit Poecile palustris decisions in a primeval forest. Bird Study 53: 199-204. Wesolowski, T., Czeszczewik, D., Rowinski, P. & Walankiewicz, W. 2002. Nest soaking in natural holes - a serious cause of breeding failure? Ornis Fennica 79: 132-138.

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Final report WC0793/CR0485 (29.04.14)

Spotted Flycatcher Muscicapa striata

1. Nest site selection/requirements Spotted Flycatcher nests are mainly on trees or buildings, frequently concealed in cavities or creepers (Kirby et al., 2005, Stoate and Szczur, 2006). Average height of nests is 3-3.6 m, but ranging from 1.3-10 m (Kirby et al., 2005, Stoate and Szczur, 2006). Nest success is greater for higher nests although this relationship is not necessarily linear (Kirby et al., 2005, Stoate and Szczur, 2006, Stevens et al., 2007). Nest success is lower in woodland and farmland than in gardens due to higher nest predation (Stoate and Szczur, 2006, Stevens et al., 2007).

2. Territory selection/attributes Occupation of habitat is greater in churchyards containing more large trees (Kirby et al., 2005).Within occupied woodland, Spotted Flycatchers will select areas dominated by Ash, with a maximum tree height of 20-25 m and low shrub diversity (Smart et al., 2007).

3. Main food items During the breeding season diet is almost entirely composed of arthropods. The type of insects taken depends on foraging method, foliar gleaning or aerial sallying. When foliar gleaning adults mainly take small (<5 mm) aphids and flies, however nestlings are preferentially fed large flies (54% of items) (Davies, 1977). When aerial sallying adults mostly eat both large and small flies (40% of diet is Diptera >5 mm, 26% of Diptera is <5 mm, other prey items taken were beetles, Ichneumids and aphids). Nestlings are preferentially fed large flies (75% of diet) (Davies, 1977). Large flies were actively selected from both feeding methods. In other, quantitative, Mediterranean studies flies were less important in diet (5-14%) with Hymenoptera (41-54%) and beetles (14-35%) more important (these studies are compared in (Boukhemza-Zemmouri et al., 2011)).

During both spring and autumn migration insects make up the majority of diet (Hernandez, 2009, Marchetti et al., 1998). On Sardinia, diet of migrant Spotted Flycatchers was predominantly large flying insects and medium sized beetles, with large flying insects selected for relative to insect abundance (Marchetti et al., 1998). In Spain 90% of diet was insects, although they also took Dogwood berries, particularly in early mornings when insect activity was low (Hernandez, 2009). The diet of migrant Spotted Flycatchers in Britain is likely to be similar to these Mediterranean studies.

4. Foraging methods Aerial sallying from a perch to catch insects in flight is the main foraging method used by Spotted Flycatchers (Davies, 1977, Alatalo and Alatalo, 1979, Randler, 2010). Perches used in a large garden were 0.5-2 m from the ground, and the distance between perch and prey was greater when catching larger prey (median 10 m) than small prey (4 m) (Davies, 1977). The height of perches used is likely to be affected by the structure of the habitat occupied. When insect activity is low, foraging method switches to gleaning insects from canopy leaves either in flight or from perches

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(Davies, 1977). Catching prey from the ground and tree bark has also been recorded in Finland (Alatalo and Alatalo, 1979). In this study pine forest was favoured for foraging over spruce and birch due to a more open forest structure.

5. Breeding habitat In England nesting habitat includes farmland, gardens and woodland (Kirby et al., 2005, Stoate and Szczur, 2006, Stevens et al., 2007) with woodland preferred over farmland (Fuller et al., 2001). Both woodland and farmland have lower breeding success than gardens (Stoate and Szczur, 2006, Stevens et al., 2007). Types of woodland used in the UK include mature broadleaved and pine woods (Smart et al., 2007, Taylor and Summers, 2010). Structural attributes of Spotted Flycatcher habitat include presence of large mature trees, gaps in the canopy, a low density of understory and shrub layers and large dead limbs in the canopy (Smith et al., 1987, Hinsley et al., 1995, Fuller, 1995, Cramp and Perrins, 1993). In Swedish conifer forests, pure spruce forests were avoided and Spotted Flycatchers were more frequent in woods with increasing proportion of pine (Gjerde and Saetersdal, 1997). Spotted Flycatchers were less likely to occur in Beech woodland and are more abundant in woods with more Bracken and dead wood on the ground (Smart et al., 2007).

6. Post fledging behaviour/requirements No information was found.

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Within woodland, Spotted Flycatchers require sheltered open space for foraging and are often associated with large mature trees with multi-layered canopies allowing feeding in a range of weather conditions. Therefore even-aged plantations will provide poor habitat for Spotted Flycatchers, particularly in younger age classes. Management that provides openings in the canopy and a varied crown structure are likely to benefit Spotted Flycatchers. Such management may include thinning, selective or group felling, and retention of mature trees or stands beyond an economically mature stage. Short term increases in breeding densities occurred in thinning oak woodland in France (Lovaty, 2004).

Spotted Flycatchers nest and feed primarily in the canopy so deer browsing is unlikely to have a short term effect on their preferred habitat. However, Ivy and Honeysuckle on trees provide good cover for nest sites and both plants are palatable and negatively affected by deer browsing. Prolonged browsing could reduce the amount of these climbing plants within the canopy.

9. References Alatalo, R.V. & Alatalo, R.H. 1979. Resource partitioning among a flycatcher guild in Finland. Oikos 33: 46-54. Boukhemza-Zemmouri, N., Belhamra, M., Boukhemza, M., Doumandji, S. & Voisin, J.

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F. 2011. Trophic ecology of the Spotted flycatcher Muscicapa striata during the breeding period in Algeria. Revue D Ecologie-La Terre Et La Vie 66: 183-194. Cramp, S. & Perrins, C.M. (1993) Handbook of the birds of Europe, the Middle East and North Africa: The birds of the Western Palearctic. Vol. VII Flycatchers to Shrikes, Oxford: Oxford University Press. Davies, N.B. 1977. Prey selection and the search strategy of the spotted flycatcher (Muscicapa striata): A field study on optimal foraging. Animal Behaviour 25: 1016- 1033. Fuller, R.J. (1995) Bird life of woodland and forest. Cambridge: Cambridge University Press. Fuller, R.J., Chamberlain, D.E., Burton, N.H.K. & Gough, S.J. 2001. Distributions of birds in lowland agricultural landscapes of England and Wales: How distinctive are bird communities of hedgerows and woodland? Agriculture Ecosystems & Environment 84: 79-92. Gjerde, I. & Saetersdal, M. 1997. Effects on avian diversity of introducing spruce Picea spp. plantations in the native pine Pinus sylvestris forests of western Norway. Biological Conservation 79: 241-250. Hernandez, A. 2009. Summer-autumn feeding ecology of Pied Flycatchers Ficedula hypolueca and Spotted Flycatchers Muscicapa striata: the importance of frugivory in a stopover area in north-west Iberia. Bird Conservation International 19: 224- 238. Hinsley, S.A., Bellamy, P.E., Newton, I. & Sparks, T.H. 1995. Habitat and landscape factors influencing the presence of individual breeding bird species in woodland fragments. Journal of Avian Biology 26: 94-104. Kirby, W., Black, K., Pratt, S. & Bradbury, R. 2005. Territory and nest-site habitat associations of Spotted Flycatchers Muscicapa striata breeding in central England. Ibis 147: 420-424. Lovaty, F. 2004. Common Redstart Phoenicurus phoenicurus and Spotted Flycatcher Muscicapa striata population size variations in mature Oak timber forests of the Allier (France). Alauda 72: 81-86. Marchetti, C., Locatelli, D.P., Van Noordwijk, A.J. & Baldaccini, N.E. 1998. The effects of prey size on diet differentiation of seven passerine species at two spring stopover sites. Ibis 140: 25-34. Randler, C. 2010. Resource partitioning between the breeding migrant Cyprus Wheatear, Oenanthe cypriaca, and the passage migrant Spotted Flycatcher, Muscicapa striata, in Cyprus (Aves: Passeriformes). Zoology in the Middle East 49: 33-38. Smart, J., Taylor, E., Amar, A., Smith, K., Bierman, S., Carpenter, J., Grice, P., Currie, F. & Hewson, C. 2007. Habitat associations of woodland birds: implications for woodland management for declining species. In RSPB Research Report No., Vol. 26. Sandy: RSPB. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica 8: 209-217. Stevens, D.K., Anderson, G.Q.A., Grice, P.V. & Norris, K. 2007. Breeding success of Spotted Flycatchers Muscicapa striata in southern England - is woodland a good habitat for this species? Ibis 149: 214-223.

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Stoate, C. & Szczur, J. 2006. Potential influence of habitat and predation on local breeding success and population in Spotted Flycatchers Muscicapa striata. Bird Study 53: 328-330. Taylor, S. & Summers, R.W. 2010. Numbers of breeding birds in old Scots Pine wood at Abernethy Forest, Badenoch & Strathspey, from 1977 to 1987. Scottish Birds 30: 302-311.

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Redstart Phoenicurus phoenicurus

1. Nest site selection/requirements Redstart nests in cavities, most typically in trees but also in man-made cavities such as nestboxes and in man-made structures such as stone walls (Lesinski, 2000, Järvinen, 1978, Collar, 2005). The Redstart is a secondary hole-nester, frequently using holes excavated by woodpeckers (Summers, 2004) but do not exclusively use cavities and are also recorded nesting on the ground between boulders and under shrubs (Edington and Edington, 1972). Nest height is variable but tends to be less than 3 m from the ground (Edington and Edington, 1972), with 1.9 m and 1.75 m average nest heights recorded in two studies (Van Balen et al., 1982, Edington and Edington, 1972) and 10.1 m in mature semi-natural pine woodland, in trees with an average DBH of 40 cm (Summers, 2004). Nest entrance diameter was on average 10.8 cm in one small study, (Van Balen et al., 1982) and very variable in a Welsh study, up to 5.5 x 13 cm (Edington and Edington, 1972). These studies found cavity entrances of varying shapes and would often have a large initial cavity entrance which became smaller before opening into the nest chamber. The average bottom area of natural cavities was 93 cm3 (Van Balen et al., 1982), while in a Finnish study they preferred large nestboxes to small ones although no nestbox size information was available (Järvinen, 1978). In areas with nestboxes, natural cavities are preferred (Van Balen et al., 1982).

Nest is constructed of grasses, roots and moss and is lined with hair and feathers (Collar, 2005).

2. Territory selection/attributes Mean breeding territory size is 0.76 ha, of which a mean of 0.19 ha was used for feeding during chick rearing (Edington and Edington, 1972). In a continental study adult Redstart exhibit moderate between-year breeding patch fidelity with 46.3% of males and 31.1% of females returning to previous breeding sites in subsequent years with 56.1% of males and 47.4% of females shifting territory within the study area between years (Sedláček and Fuchs, 2008).

A number of studies find vegetation structure of key importance in territory selection, territories were associated with areas of bare ground or sparse ground vegetation, with this preference found in several different habitat types (Schaub et al., 2010, Martinez, 2012, Martinez et al., 2010, Sedláček et al., 2004). A greater quantity of bare ground and sparse vegetation within territories also increases clutch size (Martinez, 2012). A study in an agricultural landscape in Switzerland found that occupancy was most likely where there was 60% bare ground and declined thereafter (Schaub et al., 2010), while in a semi-urban landscape territories contained at least 27% woodland and no more than 23% buildings (Sedláček et al., 2004).

3. Main food items Nestling diet comprised a wide variety of invertebrates. There isn’t a strong selection for particular invertebrate groups, two studies from Turkey & Czech Republic found

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the most frequent groups (Coleoptera, Lepidoptera and Hymenoptera) made up only 15-35% of the diet (Erdogan et al., 2005, Sedláček et al., 2007). A review of 17 nestling diet studies, found that only 35% listed caterpillars as either the primary or secondary prey item and a wide variety of invertebrate taxa were taken, with Araneae, Isopoda, Diptera, Coleoptera, moths, Gastropoda, Hymenoptera and Formicidae being frequently recorded (Cholewa and Wesolowski, 2011)). Bilberries have also been recorded fed to young (Porkert and Spinka, 2005)

On migration they also are mainly insectivorous, taking small and medium sized beetles and flies (Marchetti et al., 1998). But, many fruits and berries have been recorded as food items outside of the breeding season (Collar, 2005).

4. Foraging methods Foraging by Redstart occurs most frequently within areas of bare ground and sparse vegetation, with one study showing this foraging behaviour occurs even when invertebrate biomass was lower in sparse areas than in adjacent taller vegetation (Martinez et al., 2010). This study also found that small and patchily distributed sparse areas are preferred, probably because of increased length of edge microhabitat.

Foraging method is mainly by watching and hunting from a perch (Sedláček et al., 2004, Krystofkova et al., 2006, Edington and Edington, 1972), which constituted 82.9% of foraging behaviour in one study(Sedláček et al., 2004). Foraging by gleaning also occurs from tree trunks, foliage and stonewalls/buildings and by catching prey in the air from short sallies (Sedláček et al., 2004, Edington and Edington, 1972). The majority of prey is taken from the ground, about 50% in two studies (Sedláček et al., 2004, Alatalo and Alatalo, 1979) and a little less in Wales (Edington and Edington, 1972). Two comprehensive studies describe foraging behaviour in detail (Alatalo and Alatalo, 1979, Edington and Edington, 1972). In Finland, foraging behaviour depended on dominant tree species; in pine woodlands most prey was taken from the trunk, in spruce woodlands most was taken from needled branches, and in deciduous woodlands most was taken from the foliage (Alatalo and Alatalo, 1979). In all woodland types foraging was mostly low to the ground with only 27% of time spent hunting in the upper half of trees in pine, 10% in spruce and 20% in deciduous woodland. Aerial foraging was infrequent and also low to the ground, mostly below 3 m. In Wales foraging was mostly from the air, ground or from branches (Edington and Edington, 1972). Early studies also find very similar results for foraging height and method, although differences between preferences are less strong (Buxton, 1950).

Foraging technique preferences can vary between sexes with males foraging more from trees and less from the ground compared to females (Edington and Edington, 1972, Alatalo and Alatalo, 1979). Sex differences in foraging technique can also be seasonal, with females increasingly foraging by foliage gleaning when feeding young nestlings in a study within pine woodland (Krystofkova et al., 2006).

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5. Breeding habitat Distribution is positively influenced by the proximity of other woodlands (Opdam et al., 1985), and availability of habitats containing sparse ground vegetation (Schaub et al., 2010). Densities can be influenced by woodland management with higher densities found in association with clearfells and stands thinned up to five years previously (Lovaty, 2004). A UK scale study found a minimum of 2.4 ha of deciduous woodland was required for occupation (Moore and Hooper, 1975). In the New Forest densities were higher, although not significantly, in wood pasture habitats compared with plantations (Smith et al., 1992) and across several UK studies densities are higher in oak woodland compared to Scots pine (Taylor and Summers, 2009). Smaller areas of suitable habitat, and woodlands smaller than 5 ha, are less suitable as they are first occupied later in the breeding season and have a higher proportion of unpaired males (Huhta and Jokimaki, 2001). In the UK, Redstart are associated with mature woodland containing large trees and a sparse shrub layer (Smith et al., 1992, Hewson et al., 2011), avoiding thickets and dense, closely spaced Scots pine woodlands (Summers, 2007, Taylor and Summers, 2009). In continental studies Redstart are commonly found in mature orchards (Birrer et al., 2007), in subalpine woodlands where they preferred meadow birch woodland compared to heath woodlands (Järvinen, 1978), in moorland fringe habitat with scattered trees (Edington and Edington, 1972) and in some semi-urban habitats (Sedláček et al., 2004, Lesinski, 2000).

One study suggests that breeding population density is influenced by vole density, with Redstart density lower two years after a peak in vole density (Veistola et al., 1996).

6. Post fledging behaviour/requirements Fledglings in a study in Germany started foraging independently from 9 days after fledging although were not independent until 14 days after fledging, with adults remaining with young for up to 31 days, when they still were vocally alerting fledglings to predators (Loehrl, 1976). Fledglings start southern migration about 7-8 weeks post-fledging with a suggestion from ringing recoveries that fledglings first explore in a northward direction before a southward movement (Baker, 1993).

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Key habitat requirement for Redstart are mature open woodlands with a sparse shrub layer and sparse ground vegetation. Redstart are most likely to be abundant in harvesting systems that create open space and that have a sparse ground cover such as clearfell, group fell, selective felling and in recently thinned (<5 years) stands (Lovaty, 2004). Tree cavities are important for nesting, therefore presence in managed woodlands may be lower where nest sites are in short supply.

Redstart makes extensive use of the understory and woodland floor for foraging and may benefit from high deer browsing, although no study shows this.

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9. References Alatalo, R.V. & Alatalo, R.H. 1979. Resource partitioning among a flycatcher guild in Finland. Oikos 33: 46-54. Baker, R.R. 1993. The function of post-fledging exploration: a pilot study of three species of passerines ringed in Britain. Ornis Scandinavica: 71-79. Birrer, S., Spiess, M., Herzog, F., Jenny, M., Kohli, L. & Lugrin, B. 2007. The Swiss agri- environment scheme promotes farmland birds: but only moderately. Journal of Ornithology 148: 295-303. Buxton, J. (1950) The redstart. London: Collins. Cholewa, M. & Wesolowski, T. 2011. Nestling food of European hole-nesting passerines: do we know enough to test the adaptive hypotheses on breeding seasons? Acta Ornithologica 46: 105-116. Collar, N. 2005. Common redstart. In Handbook of Birds of the World. (Eds. J. del Hoyo, A. Elliott & D. Christie), Vol. 10, pp. 771-772. Barcelona: Lynx Edicions. Edington, J. & Edington, M. 1972. Spatial patterns and habitat partition in the breeding birds of an upland wood. Journal of Animal Ecology 41: 331-357. Erdogan, A., Kacar, M.S., Turan, L.S., Kiziroglu, I. & Oz, M. 2005. Forest pest control by common redstart (Phoenicurus phoenicurus L.) near Antalya, Turkey. Journal of Pest Science 78: 243-245. Hewson, C.M., Austin, G.E., Gough, S.J. & Fuller, R.J. 2011. Species-specific responses of woodland birds to stand-level habitat characteristics: The dual importance of forest structure and floristics. Forest Ecology and Management 261: 1224-1240. Huhta, E. & Jokimaki, J. 2001. Breeding occupancy and success of two hole-nesting passerines: the impact of fragmentation caused by forestry. Ecography 24: 431- 440. Järvinen, A. 1978. Population dynamics of the Redstart Phoenicurus phoenicurus in a subarctic area. Ornis Fennica 55: 69-76. Krystofkova, M., Exnerova, A. & Porkert, J. 2006. Parental foraging strategies and feeding of nestlings in Common Redstart Phoenicurus phoenicurus. Ornis Fennica 83: 49-58. Lesinski, G. 2000. Location of bird nests in vertical metal pipes in suburban built-up area of Warsaw. Acta Ornithologica (Warsaw) 35: 211-214. Loehrl, H. 1976. Observations on a pair of redstarts Phoenicurus phoenicurus. Vogelwelt 97: 132-139. Lovaty, F. 2004. Common Redstart Phoenicurus phoenicurus and Spotted Flycatcher Muscicapa striata population size variations in mature Oak timber forests of the Allier (France). Alauda 72: 81-86. Martinez, N. 2012. Sparse vegetation predicts clutch size in Common Redstarts Phoenicurus phoenicurus. Bird Study. Martinez, N., Jenni, L., Wyss, E. & Zbinden, N. 2010. Habitat structure versus food abundance: the importance of sparse vegetation for the common redstart Phoenicurus phoenicurus. Journal of Ornithology 151: 297-307. Moore, N.W. & Hooper, M.D. 1975. On the number of bird species in British woods. Biological Conservation 8: 239-250. Opdam, P., Rijsdijk, G. & Hustings, F. 1985. Bird communities in small woods in an agricultural landscape: effects of area and isolation. Biological Conservation 34: 333-352.

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Porkert, J. & Spinka, M. 2005. Feeding of bilberries to common redstart nestlings: an emergency strategy? British Birds 98: 265-266. Schaub, M., Martinez, N., Tagmann-Ioset, A., Weisshaupt, N., Maurer, M. L., Reichlin, T.S., Abadi, F., Zbinden, N., Jenni, L. & Arlettaz, R. 2010. Patches of bare ground as a staple commodity for declining ground-foraging insectivorous farmland birds. PloS one 5: e13115-e13115. Sedláček, O. & Fuchs, R. 2008. Breeding site fidelity in urban Common Redstarts Phoenicurus phoenicurus. Ardea 96: 261-269. Sedláček, O., Fuchs, R. & Exnerová, A. 2004. Redstart Phoenicurus phoenicurus and black redstart P. ochruros in a mosaic urban environment: neighbours or rivals? Journal of Avian Biology 35: 336-343. Sedláček, O., Fuchs, R. & Exnerová, A. 2007. Differences in the nestling diets of sympatric Redstarts Phoenicurus phoenicurus and Black Redstarts P ochruros: species-specific preferences or responses to food supply? Acta Ornithologica 42: 99-106. Smith, K.W., Burges, D.J. & Parks, R.A. 1992. Breeding bird communities of broadleaved plantation and ancient pasture woodlands of the New Forest. Bird Study 39: 132-141. Summers, R.W. 2004. Use of pine snags by birds in different stand types of Scots Pine Pinus sylvestris. Bird Study 51: 212-221. Summers, R.W. 2007. Stand selection by birds in Scots pinewoods in Scotland: the need for more old-growth pinewood. Ibis 149 (suppl 2): 175-182. Taylor, S.D. & Summers, R.W. 2009. Breeding numbers and stand type preferences of Redstarts Phoenicurus phoenicurus and Tree Pipits Anthus trivialis in a Scots Pine Pinus sylvestris wood. Bird Study 56: 120-126. Van Balen, J., Booy, C., Van Franeker, J. & Osieck, E. 1982. Studies on hole nesting birds in natural nest sites. Ardea 70: 1-24. Veistola, S., Lehikoinen, E., Eeva, T. & IsoIivari, L. 1996. The breeding biology of the Redstart Phoenicurus phoenicurus in a marginal area of Finland. Bird Study 43: 351-355.

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Pied Flycatcher Ficedula hypoleuca

1. Nest site selection/requirements The Pied Flycatcher is a secondary hole-nester, nesting in cavities. Nest cavities are predominantly in trees but nestboxes very readily occupied, and if provided in sufficient quantity in suitable habitat then are preferred over natural cavities to the extent that nesting in natural cavities becomes rare (Van Balen et al., 1982). Predation is reported to be lower in nestboxes compared with natural cavities by one study (Nilsson, 1984) but is contradicted in another (Czeszczewik, 2004). Occasionally they will nest in cavities in stone walls (Campbell, 1955). Natural cavities used are frequently woodpecker excavated; 34% of nests in Sweden (Alatalo et al., 1988) and 75% in Poland (Czeszczewik and Walankiewicz, 2003). Average height of natural nests was 8.2 m in Poland where they were up to 23 m high (Czeszczewik and Walankiewicz, 2003), 2.4 m in England (Edington and Edington, 1972), and 3.3 m and 5 m in two Swedish studies (Nilsson, 1984). Most nestbox studies situate boxes 1-4 m high, although height is rarely reported in such studies (Alatalo et al., 1986, Taylor, 2006).

Hornbeam and riverine Alder and Ash trees were preferred tree species for locating natural nests although nests were also situated in coniferous trees, particularly if these were dead (Alatalo et al., 1986). Overall 84% were located in trunks and 16% in limbs, with 76% in live trees (Alatalo et al., 1986). Mean DBH of selected nest trees from two studies were was 46 cm (Czeszczewik and Walankiewicz, 2003) and 39 cm (Alatalo et al., 1986).

Size of natural nest cavities selected was very variable with the bottom area in Sweden ranging between 33-735 cm2 (median 123cm2) (Alatalo et al., 1986) and in Poland 28-2826 cm2 (average 102c m2) (Czeszczewik and Walankiewicz, 2003). Median volume was 2035 cm3 (range 210-12725 cm3) in the Swedish study (Alatalo et al., 1986). Some studies report that larger nestboxes are preferred to smaller ones (Slagsvold, 1987). Nest entrances of natural holes are usually circular or slightly oval and have a diameter of 3.5-3.8 cm (Czeszczewik and Walankiewicz, 2003) and 3.9-5.4 cm (Edington and Edington, 1972). Nestbox entrance diameters are typically 2.8-3.5 cm. Predation rates are influenced by depth of cavity with deeper cavities having lower rates of predation; the average depth of natural nests in one study was 19.3 cm (Czeszczewik and Walankiewicz, 2003). Both natural and nestbox nests with entrances on a level plane are preferred to entrances sloping up or down (Slagsvold, 1987, Czeszczewik and Walankiewicz, 2003)

Nestbox studies show that nests containing nest material from previous years are preferentially selected, possibly as these are an indication of a good territory compared to a clean cavity (Olsson and Allander, 1995). However one study contradicted this, finding clean boxes preferred (Alatalo et al., 1986).

Although nest sites are situated within woodlands, and so are largely sheltered from effects of weather, the facing direction of nest entrances was found to be important in two studies; in Germany east facing entrances (sheltered from prevailing weather)

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were preferred (Gaedecke and Winkel, 2005) and in England nest success was lower in nests facing the prevailing weather, south-southwest (Goodenough et al., 2008).

Nests are constructed of dead leaves, plant stems, moss and grasses (Taylor, 2006).

2. Territory selection/attributes Mean breeding territory size is 0.44 ha, of which 0.17 ha was used while feeding chicks (Edington and Edington, 1972). Adult Pied Flycatchers often exhibit strong between year breeding patch fidelity with up to 46% of males and 32% of females returning to previous breeding sites in Russia (Chernetsov et al., 2009), 23% of females in England (Campbell, 1959), 35-44% in Spain and ranging between 7-49% from 24 studies from across Europe (Sanz, 2001).

Territories are centred away from nest sites of potential raptor predators, 330-430 m away from Sparrowhawk nests in Finland (Thomson et al., 2006) and away from Pygmy owl territories (Morosinotto et al., 2010).

Vegetation structure is important in territory selection; woodland with a lack of scrub cover is most important (Campbell, 1955), preferring mature woodland that is open and light (Taylor, 2006). In coniferous woodland nests were located in areas containing 45% pine within 50 m (Eeva et al., 1997).

There is some evidence that Pied Flycatcher territory settlement in spring is despotic, with the best territories (in deciduous woodland, away from the woodland edge and in larger woodlands) occupied first (Huhta and Jokimaki, 2001, Huhta et al., 1999, Huhta et al., 1998). Density dependence in breeding success is detected in some studies (Stenning et al., 1988).

3. Main food items Main food items taken depend on foraging habitat and weather (Cramp and Perrins, 1993). Of 62 studies of nestling diet reviewed, 42% listed caterpillars as either the primary or secondary prey item, with 66% of prey being Lepidoptera at any life stage (Cholewa and Wesolowski, 2011). However, a wide variety of invertebrate taxa are taken with Araneae, Hymenoptera, Coleoptera, Diptera, Homoptera, Diplopoda, adult invertebrates, Hemiptera, Gastropod, Formicidae and Ephemeroptera all frequently recorded (Cholewa and Wesolowski, 2011, Belskii and Belskaya, 2013).

Prey items taken vary seasonally, with the proportion of caterpillars in nestling diet declining seasonally in deciduous woodlands (Burger et al., 2012). There was between year variation in the quantity of caterpillars fed to nestlings with more spiders fed in years when less caterpillars were fed (Siikamaki et al., 1998, Bel'skii and Bel'skaya, 2009). Another study in subarctic Finland found more ground dwelling species of spider were taken during the early part of the breeding season (Eeva et al., 2000).

Prey items taken also varied between woodland types with the average proportions of caterpillars taken lower in coniferous or birch woodland (23.7%) compared with

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oak (37.5%)(Burger et al., 2012). A review of 17 studies found the same, but also that the proportion of Coleoptera in nestling diet was higher in coniferous woodland (Sanz, 1998). Snails as a source of calcium are an important part of nestling diet (Eeva et al., 2010), constituting up to 7% of nestling diet in Germany (Wimmer and Winkel, 2001).

On migration, they are also mainly insectivorous taking beetles, ants and some flying insects (Marchetti et al., 1998) but will take berries, with up to 30% of diet being fruits including dogwood, Elder and Bramble on migration in Spain (Hernandez, 2009).

4. Foraging methods Foraging by Pied Flycatcher is either by taking prey from the ground, from trees and foliage, or from the air in aerial sallies but preferred foraging method varies between studies and woodland types, and was influenced by seasonality and weather. In several studies aerial foraging is most frequent in favourable weather conditions but this shifts to ground foraging when activity of flying insects is reduced by low temperatures (Alatalo and Alatalo, 1979, Lennerstedt, 1983, Veistola et al., 1997). In Wales, foraging was equally from the air, and from trees (41%) with 17.6% of foraging from the ground (Edington and Edington, 1972).

In Finland, foraging behaviour depended on dominant tree species; in pine woodlands most prey was taken from needled branches and in deciduous woodlands most was taken from foliage (Alatalo and Alatalo, 1979). Within coniferous woodlands pine trees were preferred over spruce and birch (39% of foraging during nesting and 80% thereafter) (Eeva et al., 1997).

Foraging method varies seasonally with predominantly ground feeding and some aerial feeding during the nesting period then progressively switching to feeding in trees after young have fledged (Alatalo and Alatalo, 1979, Eeva et al., 1997); this may be related to abundance of larvae on birch trees (Veistola et al., 1997).

In birch woodland, foraging height was mainly 2-6 m (58%) with 42% between 0-2 m and none above 6 m(Veistola et al., 1997). In this study, foraging is primarily on outer parts of trees and in foliage (Veistola et al., 1997) although in another study foraging was mainly from trunks (43%) but also branches (18%) and foliage (12%) (Eeva et al., 1997). In a captive experiment, foraging was primarily on the ground and in lower parts of trees (Adamik and Bures, 2007).

No differences in foraging methods or prey choice have been reported between sexes when examined (Siikamaki et al., 1998, Eeva et al., 2005).

5. Breeding habitat In the UK Pied Flycatchers are associated with mature oakwoods (Campbell, 1955) but also occur in mixed woodlands (Wright et al., 2010). There is a relationship between woodland size and occupation with larger woodlands having higher densities and woodlands <1 ha being largely unoccupied with a high proportion of

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unpaired males (Huhta and Jokimaki, 2001, Huhta et al., 1998, Loman, 2006). Provision of nestboxes increases density in large woodlands but not small (Loman, 2006). The density of breeding pairs is higher in deciduous woodlands compared to coniferous woodlands, with timing of spring arrival and breeding being earlier and breeding success higher in deciduous woodland (Silverin, 1998, Lundberg et al., 1981, Huhta et al., 1998). Within coniferous woodlands clutch size and reproductive success tended to be higher in Scots pine compared to Lodgepole pine (Sjoberg et al., 1993). For habitat near rivers regulation of river flow can reduce breeding success through alteration of emergent aquatic insects communities which can be an important component of nestling diet (Strasevicius et al., 2013).

6. Post fledging behaviour/requirements Fledglings stay in their natal area for about 27 days after fledging before dispersal into surrounding areas (Van Balen, 1979), which includes non-wooded habitats (M. Burgess unpublished). Females disperse further than males.

7. Seasonal differences in habitat for resident species N/A

8. Effects of forest management and deer on habitat Key habitat requirements for Pied Flycatcher are mature open woodlands with a sparse shrub layer and sparse ground vegetation. Pied Flycatcher are largely absent from intensively managed woodlands and are more likely to be abundant in harvesting systems that retain mature trees and continuous canopy cover, create open space and that have a sparse ground cover such as group and selective felling. Tree cavities are vital for nesting and so presence in young, thinned or felled stands is much less likely unless nestboxes are provided. A few studies suggest that foliage feeders are negatively affected by thinning management, which may affect Pied Flycatcher (Virkkala, 1987, de la Montaña et al., 2006). In a UK oak wood light thinning of the canopy resulted in higher territory occupation rates but not in higher clutch size or nest success (Burgess, in review).

Pied Flycatchers make extensive use of the understory and ground for foraging and may benefit from high deer browsing, although no study shows this.

9. References Adamik, P. & Bures, S. 2007. Experimental evidence for species-specific habitat preferences in two flycatcher species in their hybrid zone. Naturwissenschaften 94: 859-863. Alatalo, R.V. & Alatalo, R.H. 1979. Resource partitioning among a flycatcher guild in Finland. Oikos 33: 46-54. Alatalo, R.V., Carlson, A. & Lundberg, A. 1988. Nest cavity size and clutch size of pied flycatchers Ficedula hypoleuca breeding in natural tree-holes. Ornis Scandinavica 19: 317-319. Alatalo, R.V., Lundberg, A. & Glynn, C. 1986. Female pied flycatchers choose territory quality and not male characteristics. Nature 323: 152-153.

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Bel'skii, E.A. & Bel'skaya, E.A. 2009. Composition of pied flycatcher (Ficedula hypoleuca Pall.) nestling diet in industrially polluted area. Russian Journal of Ecology 40: 342-350. Belskii, E. & Belskaya, E. 2013. Diet composition as a cause of different contaminant exposure in two sympatric passerines in the Middle Urals, Russia. Ecotoxicology and Environmental Safety 97: 67-72. Burger, C., Belskii, E., Eeva, T., Laaksonen, T., Mägi, M., Mänd, R., Qvarnström, A., Slagsvold, T., Veen, T., Visser, M. E., Wiebe, K. L., Wiley, C., Wright, J. & Both, C. 2012. Climate change, breeding date and nestling diet: how temperature differentially affects seasonal changes in pied flycatcher diet depending on habitat variation. Journal of Animal Ecology 81: 926-936. Burgess, M.D. in review. Weather is a stronger influence than woodland thinning on occupancy, fecundity and productivity of hole nesting birds in an upland Atlantic oakwood. Forest Ecology and Management. Campbell, B. 1955. The breeding distribution and habitats of the Pied Flycatcher (Muscicapa hypoleuca) in Britain. Bird Study 2: 24-32. Campbell, B. 1959. Attachment of Pied flycatchers Muscicapa hypoleuca to nest- sites. Ibis 101: 445-448. Chernetsov, N., Sokolov, L.V. & Kosarev, V. 2009. Local survival rates of pied flycatchers Ficedula hypoleuca depend on their immigration status. Avian Ecol. Behav 16: 11-20. Cholewa, M. & Wesolowski, T. 2011. Nestling food of European hole-nesting passerines: do we know enough to test the adaptive hypotheses on breeding seasons? Acta Ornithologica 46: 105-116. Cramp, S. & Perrins, C.M. (1993) Handbook of the birds of Europe, the Middle East and North Africa: The birds of the Western Palearctic. Vol. VII Flycatchers to Shrikes, Oxford: Oxford University Press. Czeszczewik, D. 2004. Breeding success and timing of the Pied Flycatcher Ficedula hypoleuca nesting in natural holes and nest-boxes in the Bialowieza Forest, Poland. Acta Ornithologica 39: 15-20. Czeszczewik, D. & Walankiewicz, W. 2003. Natural nest sites of the pied flycatcher Ficedula hypoleuca in a Primeval Forest. Ardea 91: 221-229. de la Montaña, E., Rey-Benayas, J.M. & Carrascal, L.M. 2006. Response of bird communities to silvicultural thinning of Mediterranean maquis. Journal of Applied Ecology 43: 651-659. Edington, J. & Edington, M. 1972. Spatial patterns and habitat partition in the breeding birds of an upland wood. Journal of Animal Ecology 41: 331-357. Eeva, T., Lehikoinen, E. & Pohjalainen, T. 1997. Pollution-related variation in food supply and breeding success in two hole-nesting passerines. Ecology 78: 1120- 1131. Eeva, T., Rainio, K. & Suominen, O. 2010. Effects of pollution on land snail abundance, size and diversity as resources for pied flycatcher, Ficedula hypoleuca. Science of the Total Environment 408: 4165-4169. Eeva, T., Ryoma, M. & Riihimaki, J. 2005. Pollution-related changes in diets of two insectivorous passerines. Oecologia 145: 629-639. Eeva, T., Veistola, S. & Lehikoinen, E. 2000. Timing of breeding in subarctic passerines in relation to food availability. Canadian Journal of Zoology 78: 67-78.

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Gaedecke, N. & Winkel, W. 2005. Are tits Parus spp. and other hole-nesting passerines preferring at the choice of their breeding holes the weather-opposing side? Vogelwarte 43: 15-18. Goodenough, A.E., Maitland, D.P., Hart, A.G. & Elliot, S.L. 2008. Nestbox orientation: a species-specific influence on occupation and breeding success in woodland passerines. Bird Study 55: 222-232. Hernandez, A. 2009. Summer-autumn feeding ecology of Pied Flycatchers Ficedula hypolueca and Spotted Flycatchers Muscicapa striata: the importance of frugivory in a stopover area in north-west Iberia. Bird Conservation International 19: 224-238. Huhta, E. & Jokimaki, J. 2001. Breeding occupancy and success of two hole-nesting passerines: the impact of fragmentation caused by forestry. Ecography 24: 431- 440. Huhta, E., Jokimaki, J. & Rahko, P. 1999. Breeding success of pied flycatchers in artificial forest edges: The effect of a suboptimally shaped foraging area. Auk 116: 528-535. Huhta, E.S.A., Jokimakp, J. & Rahko, P. 1998. Distribution and reproductive success of the Pied Flycatcher Ficedula hypoleuca in relation to forest patch size and vegetation characteristics; the effect of scale. Ibis 140: 214-222. Lennerstedt, I. 1983. Feeding ranges of willow warbler and pied flycatcher in subalpine birch forest. Var Fagelvarld 42: 11-20. Loman, J. 2006. Does nest site availability limit the density of hole nesting birds in small woodland patches? Web Ecology 6: 37-43. Lundberg, A., Alatalo, R.V., Carlson, A. & Ulfstrand, S. 1981. Biometry, habitat distribution and breeding success in the Pied flycatcher Ficedula hypoleuca. Ornis Scandinavica 12: 68-79. Morosinotto, C., Thomson, R.L. & Korpimaki, E. 2010. Habitat selection as an antipredator behaviour in a multi-predator landscape: all enemies are not equal. Journal of Animal Ecology 79: 327-333. Nilsson, S.G. 1984. The evolution of nest-site selection among hole-nesting birds: The importance of nest predation and competition. Ornis Scandinavica 15: 167-175. Olsson, K. & Allander, K. 1995. Do fleas, and/or old nest material, influence nest-site preferences in hole-nesting passerines. Ethology 101: 160-170. Sanz, J.J. 1998. Effect of habitat and latitude on nestling diet of Pied Flycatchers Ficedula hypoleuca. Ardea 86: 81-88. Sanz, J.J. 2001. Latitudinal variation in female local return rate in the philopatric pied flycatcher (Ficedula hypoleuca). Auk 118: 539-543. Siikamaki, P., Haimi, J., Hovi, M. & Ratti, O. 1998. Properties of food loads delivered to nestlings in the pied flycatcher: effects of clutch size manipulation, year, and sex. Oecologia 115: 579-585. Silverin, B. 1998. Territorial behaviour and hormones of pied flycatchers in optimal and suboptimal habitats. Animal Behaviour 56: 811-818. Sjoberg, K., Pettersson, R. & Atlegrim, O. 1993. Differences in bird habitat quality between plantations of Scots and Lodgepole pine measured in terms of Pied flycatcher Ficedula hypoleuca breeding success. Ornis Svecica 3: 59-68. Slagsvold, T. 1987. Nest site preference and clutch size in the Pied flycatcher Ficedula hypoleuca. Ornis Scandinavica 18: 189-197.

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Stenning, M., Harvey, P.H. & Campbell, B. 1988. Searching for density-dependent regulation in a population of Pied flycatchers Ficedula hypoleuca Pallas. Journal of Animal Ecology 57: 307-317. Strasevicius, D., Jonsson, M., Nyholm, N.E.I. & Malmqvist, B. 2013. Reduced breeding success of Pied Flycatchers Ficedula hypoleuca along regulated rivers. Ibis 155: 348-356. Taylor, P.B. 2006. Pied flycatcher. In Handbook of Birds of the World (Eds. J. del Hoyo, A. Elliott & D. Christie), Vol. 11, pp. 129-130. Barcelona: Lynx Edicions. Thomson, R.L., Forsman, J.T., Sarda-Palomera, F. & Monkkonen, M. 2006. Fear factor: prey habitat selection and its consequences in a predation risk landscape. Ecography 29: 507-514. Van Balen, J., Booy, C., Van Franeker, J. & Osieck, E. 1982. Studies on hole nesting birds in natural nest sites. Ardea 70: 1-24. Van Balen, J.H. 1979. Observations on the post-fledging dispersal of the pied flycatcher, Ficedula hypoleuca. Ardea 67: 134-137. Veistola, S., Lehikoinen, E. & Eeva, T. 1997. Foraging niches of two passerines at their subarctic limit of distribution: The Siberian tit Parus cinctus and the pied flycatcher Ficedula hypoleuca. Ornis Fennica 74: 79-87. Virkkala, R. 1987. Effects of forest management on birds breeding in northern Finland. Annales Zoologici Fennici 24: 281-294. Wimmer, W. & Winkel, W. 2001. Snails (Gastropoda) in the diet of Pied Flycatcher (Ficedula hypoleuca) nestlings. Vogelwarte 41: 70-80. Wright, J., Karasov, W.H., Kazem, A.J.N., Braga Goncalves, I. & McSwan, E. 2010. Begging and digestive responses to differences in long-term and short-term need in nestling pied flycatchers. Animal Behaviour 80: 517-525.

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Final report WC0793/CR0485 (29.04.14)

Hawfinch Coccothraustes coccothraustes

1. Nest site selection/requirements Hawfinch nests are constructed from a base of sticks, with a cup built of roots and grass and decorated with lichens (Roberts and Lewis, 1988, Westwood and Watson, 2010, Mountford, 1957). A mixture of fine birch twigs and stouter oak twigs made up most nests in Britain (Mountford, 1957). Nests are located in a wide range of tree species, 10-20 tree species in individual study areas (Bijlsma, 1998, Tomialojc, 2005). Broadleaved trees are used more often than conifers with Ash, oak, maple, Larch, birch and Beech preferred nesting trees in more than one study (von Haartman, 1978, Roberts and Lewis, 1988, Bijlsma, 1998, Smith, 2004, Tomialojc, 2005, Westwood and Watson, 2010). Location of nest sites is very variable although always in trees, often nesting in upper and mid crowns of broadleaved trees but lower branches of conifers, at a height of 5-25m (von Haartman, 1978, Roberts and Lewis, 1988, Bijlsma, 1998, Smith, 2004, Tomialojc, 2005). Hawfinches are also found using Ivy, Honeysuckle and Mistletoe as cover for nest sites (Roberts and Lewis, 1988, Smith, 2004, Tomialojc, 2005).

2. Territory selection/attributes Hawfinches are not strongly territorial and often nest in loose colonies (Roberts and Lewis, 1988). Display activity and defence is often focused on the female rather than nest location and territorial activity can often take place in stands used for feeding, which can be up to 200 m from nest locations (Bijlsma, 1998). Hawfinches can range over large areas using more than one wood during the breeding season (Bijlsma, 1998, Calladine and Morrison, 2010).

3. Main food items Nestlings are fed mostly invertebrates, particularly caterpillars (Cramp et al., 1994, Bijlsma, 1998). Adult birds feed on seeds and buds from a wide variety of tree and shrub species and have the ability to eat hard stoned fruits such as cherry not available to other bird species, seeds most regularly recorded are a variety of prunus species, yew, beech and hornbeam (Cramp et al., 1994, Mountford, 1957). In spring, Elm seeds and buds (poplars, ash, maples & beech) and flowers (oak, maples, beech) from a range of tree species are important (von Haartman, 1978, Bijlsma, 1998, Bryant, 2011, Tomialojc, 2012, Mountford, 1957). In spring they will also feed on invertebrates, in particular geometrid moth caterpillars during outbreak years (Tomialojc, 2012, Mountford, 1957). During the rest of the year seeds of Hornbeam, Prunus species, Ash and maple are important food sources, but locally can make use of other abundant seed sources such as pines or from bird tables (von Haartman, 1978, Batt, 1993, Bijlsma, 1998).

4. Foraging methods During the breeding season foraging mostly occurs in the canopy, often in loose flocks, but in late winter after most seed has fallen they forage in flocks on the ground (Cramp et al., 1994).

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5. Breeding habitat Breeding habitat always includes mature trees, either plantations, parkland/wood pasture or semi natural forests (Roberts and Lewis, 1988, Smith et al., 1992, Bijlsma, 1998, Hubalek, 2001, Müller, 2005, Tomialojc, 2005, Smart et al., 2007). Within the UK, Hawfinches may be associated with non-native plantations with high tree diversity (Smith, 2004). Hawfinches are also associated with more wooded landscapes and woods that have more trees with Ivy (Smith et al., 1987, Smart et al., 2007).

On continental Europe they occur in flood plain forest, dense mature oak forest, young plantations and semi-natural oak/lime/hornbeam forest (Bijlsma, 1998, Hubalek, 2001, Müller, 2005, Tomialojc, 2005) but are more frequent in broadleaf compared to coniferous forest (Baguette et al., 1994, Paquet et al., 2006).

6. Post fledging behaviour/requirements Fledglings can leave the nesting area within a few days of fledging, but post-fledging ranging is largely unknown (Tomialojc, 2004).

7. Seasonal differences in habitat for resident species In winter Hawfinches use similar types of woodland as during the breeding season but utilise dense conifers such as Norway spruce, Douglas fir and Western hemlock for roosting (Smith, 2004, Calladine and Morrison, 2010).

8. Effects of forest management and deer on habitat There is little information on the impacts of forest management on Hawfinch, however, some inferences can be drawn based on their habitat requirements. As Hawfinch is primarily a species of mature woodland year-round management such as coppicing or clearfell that results in large areas of young stands will reduce the amount of habitat available. In contrast, harvesting systems aimed at maintaining mature forest cover are likely to be beneficial to Hawfinch populations if the stands have high tree diversity (de Warnaffe and Deconchat, 2008). Restoration of ancient woodland sites by the removal of non-native tree species has the potential to reduce the amount of Hawfinch habitat available, as they feed on a wide range of non- native tree seed and are frequently found in mixed broadleaf/conifer woodland. Hawfinches also utilise conifers, which may be removed during ancient woodland restoration, for winter roost sites.

The presence of high deer densities in woodland is likely to have only a minor impact on Hawfinch numbers. Deer have little effect on canopy trees where Hawfinches breed and forage, however deer browsing can reduce the frequency of climbers present and these are often used as cover for early nests. A significant reduction in Ivy and Honeysuckle could lead to increased nest predation and lower productivity. There is also the possibility for direct competition for food resources in winter when deer and Hawfinches may both feed on fallen tree seeds.

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9. References Baguette, M., Deceuninck, B. & Muller, Y. 1994. Effects of spruce afforestation on bird community dynamics in a native broad-leaved forest area. Acta Oecologica 15: 275-288. Batt, L. 1993. Hawfinches feeding like crossbills on Scots pine. British Birds 86: 133. Bijlsma, R.G. 1998. Breeding biology and population trend of hawfinches Coccothraustes coccothraustes in Flevoland. Limosa 71: 137-148. Bryant, D. 2011. Multi-species groups of finches feeding on Wych Elm fruits in spring. Scottish Birds 31: 311-314. Calladine, J. & Morrison, N. 2010. The ranging behaviour and habitat selection by three Hawfinches Coccothraustes coccothraustes in late winter in Scotland. Ornis Fennica 87: 119-123. Cramp, S., Perrins, C.M. & Brooks, D. J. (1994) Handbook of the birds of Europe, the Middle East and North Africa. Volume 8: Crows to Finches., Oxford: Oxford University Press. de Warnaffe, G.D. & Deconchat, M. 2008. Impact of four silvicultural systems on birds in the Belgian Ardenne: implications for biodiversity in plantation forests. Biodiversity and Conservation 17: 1041-1055. Hubalek, Z. 2001. Seasonal variation of forest habitat preferences by birds in a lowland riverine ecosystem. Folia Zoologica 50: 281-289. Mountford, G. (1957) The Hawfinch, London: Collins. Müller, J. 2005. Bird communities as indicators for woodland structures in oak woods. Ornithologische Beobachter 102: 15-32. Paquet, J.Y., Vandevyvre, X., Delahaye, L. & Rondeux, J. 2006. Bird assemblages in a mixed woodland-farmland landscape: The conservation value of silviculture- dependant open areas in plantation forest. Forest Ecology and Management 227: 59-70. Roberts, S. & Lewis, J. 1988. Observations on the sensitivity of nesting hawfinch. Gwent Bird Report: 7-10. Smart, J., Taylor, E., Amar, A., Smith, K., Bierman, S., Carpenter, J., Grice, P., Currie, F. & Hewson, C. 2007. Habitat associations of woodland birds: implications for woodland management for declining species. In RSPB Research Report No., Vol. 26. Sandy: RSPB. Smith, D. 2004. Breeding and roosting hawfinches (Coccothraustes coccothraustes) in Merioneth. Welsh Birds 4: 11-19. Smith, K.W., Averis, B. & Martin, J. 1987. The breeding bird community of oak plantations in the Forest of Dean, southern England. Acta Oecologica, 8: 209-217. Smith, K.W., Burges, D.J. & Parks, R.A. 1992. Breeding bird communities of broadleaved plantation and ancient pasture woodlands of the New Forest. Bird Study 39: 132-141. Tomialojc, L. 2004. Accuracy of the mapping technique for a dense breeding population of the Hawfinch Coccothraustes coccothraustes in a deciduous forest. Acta Ornithologica 39: 67-74. Tomialojc, L. 2005. Distribution, breeding density and nest sites of Hawfinches Coccothraustes coccothraustes in the primeval forest of Bialowieza National Park. Acta Ornithologica 40: 127-138.

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Tomialojc, L. 2012. Reproduction and population dynamics of Hawfinches Coccothraustes coccothraustes in the primeval forest of Bialowieia National Park (NE Poland). Acta Ornithologica 47: 63-78. von Haartman, L. 1978. An account of a small population of hawfinches. Ornis Fennica 55: 132-133. Westwood, N.J. & Watson, M. 2010. Hazards of man-made material to nesting hawfinches. British Birds 103: 247.

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