Journal of Applied Ecology 2014, 51, 204–213 doi: 10.1111/1365-2664.

12173

Modelling edge effects of mature forest plantations on

peatland waders informs landscape-scale
conservation
Jeremy D. Wilson1*, Russell Anderson2, Sallie Bailey3, Jordan Chetcuti2, Neil R. Cowie1,
Mark H. Hancock4, Christopher P. Quine2, Norrie Russell5, Leigh Stephen1 and
Des B. A. Thompson6
1
RSPB Scotland, 2 Lochside View, Edinburgh Park, Edinburgh EH12 9DH, UK; 2Forest Research, Northern Research
Station, Roslin, Midlothian EH25 9SY, UK; 3Forestry Commission Scotland, Silvan House, 231 Corstorphine Road,
Edinburgh EH12 7AT, UK; 4RSPB Scotland, Etive House, Beechwood Park, Inverness IV2 3BW, UK; 5RSPB Peatland
Reserves Office, Forsinard, Sutherland KW13 6YT, UK; and 6Scottish Natural Heritage, Silvan House, 3rd Floor East,
231 Corstorphine Road, Edinburgh EH12 7AT, UK

Summary
1. Edge effects of native forest fragmentation have been well studied, but there are few stud-
ies of open-ground habitats fragmented by plantation forests. We measure forestry edge
effects on open-ground breeding birds, following one of Europe’s biggest and most controver-
sial land-use transformations.
2. The ‘Flow Country’ of northern Scotland is one of the world’s greatest expanses of blan-
ket bog. It became fragmented by conifer forests planted in the late 20th century, and these
now adjoin open peatlands protected under European conservation legislation. Detrimental
edge effects on breeding birds were anticipated, but not apparent shortly after planting.
3. Using survey data collected in 2003–2006, and logistic regression modelling, we tested
whether breeding distributions of three wader species of international conservation concern,
dunlin, European golden plover and common greenshank, were influenced by distance to
forest edge, controlling for habitat and topography.
4. All three species were more likely to occupy flatter, more exposed ground close to bog
pools and were influenced by peatland vegetation structure. There was an additive and
adverse effect of proximity to forest edge for dunlin and European golden plover, but not
common greenshank. This effect was strongest within 700 m of forest edges. We used these
results to predict which areas should benefit most from removal of adjacent forestry and so
guide maintenance and restoration of the bird interests of the protected areas.
5. Synthesis and applications. Edge effects of mature forestry on dunlin and golden plover
are apparent over several hundred metres and are now being used to guide forest planning in
northern Scotland. The scale of edge effect is broadly consistent with other avian studies in
open-ground habitats across Eurasia and North America, so buffer zones of this order are
consistent with possible impacts of plantation forestry on open-ground habitats of bird con-
servation interest. Given renewed interest in conifer afforestation as a climate change mitiga-
tion measure, an improved understanding of edge effects and the mechanisms through which
they operate is vital to managing plantation forestry in ways that maintain open-ground land-
scapes of high conservation value.
Key-words: blanket bog, climate change adaptation, dunlin, forestry, fragmentation, golden
plover, greenshank, nest predation, open habitat, protected areas

*Correspondence author. E-mail: jeremy.wilson@rspb.org.uk

© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society
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Forest edge effects on peatland waders 205

Introduction interests greater (Avery & Leslie 1990). The ‘Flow Coun-
try’ of northern Scotland is a globally important exam-
Understanding how habitat edges influence distribution,
ple. It is one of the world’s most extensive, contiguous
abundance and behaviour of wildlife is critical to large-
areas of blanket bog, a landscape dominated by deep
scale conservation management (Murcia 1995; Ries et al.
peat, and vegetation cover rich in peat-forming Sphagnum
2004; Fuller 2012). The pivotal role of predation and repro-
mosses, along with grasses, sedges and ericaceous dwarf
ductive failure in bird demography (Martin 1995) underlies
shrubs. This landscape is now fragmented by plantations
strong interest in edge effects in avian conservation.
of non-native lodgepole pine Pinus contorta and Sitka
Reviews show that effects are variable and context-specific
spruce Picea sitchensis, in areas that had been treeless for
(Batary & Baldi 2004) because land-use composition and
millennia (Stroud et al. 1987). This was one of the most
history determine the nest predator fauna, the hard or soft
controversial European land-use transformations of the
nature of habitat edges, and whether some species are
late 20th century (Avery & Leslie 1990; Oosthoek 2013).
adapted to young-growth habitat at ecotones between
Much remaining open peatland (146 000 ha) is now des-
forest and open habitats (Stephens et al. 2003; Thompson
ignated as special protection areas (SPAs) and special
2007; Dolman 2012; Fuller 2012).
areas of conservation (SACs) under European Union
Most evidence of edge effects comes from studies of
(EU) Birds and Habitats Directives. This recognizes a
forest habitats fragmented by agricultural land uses. These
globally unique assemblage of habitats and breeding
have found reduced occupancy and nest success close to
birds that is unusually diverse amongst northern temper-
edges in highly fragmented forest landscapes in both trop-
ate peatland systems. This included 66% of common
ical and temperate systems across a broad spectrum,
greenshank Tringa nebularia (Gunnerus, 1767), 35% of
including gamebirds (e.g. capercaillie Tetrao urogallus –
dunlin Calidris alpina (L.) and 17% of European
Kurki et al. 2000), tree-nesting seabirds (e.g. marbled
golden plover Pluvialis apricaria (L.) in the European
murrelet Brachyramphus marmoratus – Malt & Lank
Community at the time of designation (Stroud et al.
2007) and songbirds (Deng & Gao 2005; Poulin & Villard
1987; Lindsay et al. 1988).
2011; Zurita et al. 2012). These effects are usually driven
Forest planting resulted in an estimated loss of 17–19%
by nest predators (e.g. corvids and foxes) and brood para-
of the Flow Country populations of each of these species
sites (e.g. brown-headed cowbird Molothrus ater) which
(Stroud et al. 1987; Lindsay et al. 1988), but with little
persist at higher abundances in fragmented landscapes
initial evidence that forest edge effects were causing fur-
and may show behavioural associations with habitat edges
ther losses (Avery 1989; Stroud, Reed & Harding 1990).
(Andren 1995; Howell, Dijak & Thompson 2007;
However, such effects were predicted as plantations grew
Cervinka et al. 2011). Conversely, edge effects of planta-
and became more suitable habitat for generalist predators
tion forestry on bird populations in open habitats of high
such as hooded crows Corvus cornix and red foxes Vulpes
conservation interest remain little studied (Reino et al.
vulpes which hunt over surrounding areas (Stroud et al.
2009). This is despite plantation forestry now occupying
1987; Lindsay et al. 1988). Since then, as forestry has
7% of global forest cover (140 million hectares), and
matured, there have been declines of European golden
increasing rates of afforestation or reforestation, driven
plover and dunlin in some areas (e.g. Sim et al. 2005) and
by production, climate change adaptation, ecosystem
weak evidence of edge effects from studies at coarse
service protection and biodiversity conservation goals
spatial grain (Hancock, Grant & Wilson 2009).
(Bauhaus, van der Meer & Kanninen 2010).
We used annual bird survey data from the Forsinard
In the UK, increasing forest cover was established
Flows Nature Reserve to measure associations between
policy through the 20th century, responding to an histori-
habitat, land cover and topography, and occurrence of
cal reduction in woodland cover and lack of a strategic
dunlin, European golden plover and greenshank in the
timber reserve (Foot 2003; Oosthoek 2013). Available
Caithness and Sutherland Peatland SPA (Fig. 1) in order
land has often been at higher latitudes and altitudes where
to test whether there is now stronger evidence of edge
land values were lower, but landscape and conservation

Fig. 1. Grid squares (200 m) making up

study area (dots). Map covers 49 9 33 km
and is centred on Forsinard Flows Nature
Reserve (58°21′N 3°53′W). Forestry cover
(1998) shaded pale grey. Caithness and
Sutherland Peatland Special Protection
Area (SPA) shaded dark grey. Specific
locations are: 1 = Blar nam Faoileag,
2 = Cross Lochs, 3 = Imriche, 4 = Sletill,
5 = Talaheel.

© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society, Journal of Applied Ecology, 51, 204–213

206 J. D. Wilson et al.
effects. We controlled for both habitat (vegetation and
soil reflectance and proximity to bog pool systems) and
topographical variation (slope and elevation) because
these are known to influence distribution of breeding wad-
ers on peatlands (Lavers & Haines-Young 1996; Lavers,
Haines-Young & Avery 1996), and more widely across
upland landscapes (Whittingham, Percival & Brown 2002;
Pearce-Higgins & Grant 2006). In this sense, we followed
the recommendations of Burnham and Anderson (2002)
in defining a biologically informed model set as our start-
ing point. Where an additive association between distance
to forest edge and grid square occupancy was found, we
then used a simple scenario of forest removal to predict
areas where benefit of forestry removal through mitigation
of edge effects is likely to be greatest. This can then be
used to help determine whether forestry areas should be
replanted or restored as blanket bog, adding to other con-
siderations such as local timber and wood fuel markets,
and Government policies on woodland expansion and
greenhouse gas (GHG) consequences of growing trees on
deep peat soils (e.g. Reynolds 2007; Morison et al. 2010;
Lohila et al. 2011).
Materials and methods
BREEDING BIRD SURVEYS
2
Survey data were based on two visits to over 250 km of peatland
(6288 grid squares of 200 9 200 m; Fig. 1) in one (24% of
squares), two (2%), three (36%) or four (38%) years between
2003 and 2006. Territorial individuals of all bird species were
mapped by walking transects that covered the whole square to
within 100 m (Brown & Shepherd 1993). These data resolved a
species as either present (1) or absent (0) in each square, pooled
over the four survey years. This measure was the response vari-
able in subsequent occupancy modelling.
EXPLANATORY VARIABLES – HABITAT
Peatland vegetation cover was first characterized from four collin-
ear reflectance bands (Landsat 7; March 2000) covering the visi-
ble red to far infra-red range and with little effect from
atmospheric interference (Avery & Haines-Young 1990; Campbell
2002). The value for each square was derived as the mean of all
Table 1. Descriptive statistics for explanatory variables measured in each 200-m square in the Flow Country study area
Variable
First principal component of reflectance bands b3, b4,
b5 and b7 – an index of vegetation cover
Photosynthetic intensity (NDVI) = (b4 b3)/(b3 + b4)
Slope
Elevation
Topographical exposure (inverse)
Loge distance to forest edge
Loge distance to pool system
Effort
© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society, Journal of Applied Ecology, 51, 204–213
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30-m pixels within or overlapping the square. The first principal
component of these bands explained 86% of variation and was
used as a measure of vegetation cover. We also included the
Normalised Difference Vegetation Index (NDVI), values of which
range from 1 to +1, as a measure of density of photosyntheti-
cally active vegetation (Campbell 2002).
EXPLANATORY VARIABLES – TOPOGRAPHY
We measured distance from each grid cell centre to the nearest
bog pool and nearest point of forestry cover (see Appendix S1,
Supporting information), using Ordnance Survey digital 1 : 10
000 maps and the National Inventory of Woodland and Trees
(NIWT), respectively. We used the UK national digital terrain
model (DTM) to measure mean slope, altitude and topographi-
cal exposure of each grid square, derived as the mean of all
points on a 10-m grid within each 200-m square. Topographical
exposure was defined as the vertical angle to the horizon
summed across the eight prime compass points (Quine & White
1993).
Finally, we multiplied the number of visits by the proportion
of squares surveyed to create a nuisance variable to control for
variation in survey effort and the fact that some squares at
forestry, lake or study area edges could only be partially
surveyed. Descriptive statistics for all explanatory variables are
summarized in Table 1.
DATA ANALYSIS
Preliminary analyses
Collinearity of explanatory variables was low. Variance inflation
factors ranged from 1
08 to 2
70, and Pearson correlation coeffi-
cients for all pairs of explanatory variables were <0
7, except for
that between slope and topographical exposure. For each species,
we therefore conducted preliminary univariate analyses including
either slope or topographical exposure (correcting for survey
effort) in order to select one of these two variables (Wald chi-
square tests). The variables selected were topographical exposure
(dunlin, European golden plover) and slope (common green-
shank).
In squares where a species was detected, we tested whether
probability of detection on both (1) or just one survey visit (0) in
a year was related to any explanatory variable, using univariate
logistic regression models. In total, we fit 77 models (three
species 9 four years 9 seven explanatory variables; too few
Abbreviation
(Table 2) Units, range and mean
R Principal component( 8
92 to 8
77; mean = 0)
n Index ( 0
53 to +0
53; mean = 0
15)
s Degrees (0 to 32°, mean = 2°)
e km (0
043 to 0
525, mean = 0
167)
t Degrees (1
3 to 94
9°, mean = 11
2)
f Metres (0 to 5684, mean = 1100)
p Metres (0 to 2202, mean = 421)
(Number of years surveyed 9 proportion of square
surveyed)
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Forest edge effects on peatland waders 207

common greenshanks were recorded in one year to allow models
to be fit). Only five models showed a significant relationship at
P < 0
05, and none at P < 0
01 (Bonferroni-adjusted P = 0
0006),
suggesting that modelling correlates of occupancy is unlikely to
be biased by associations between explanatory variables and bird
detectability.
concerned. Finally, we compared these values between squares
that had experienced an increase in distance to forest between
1998 and 2003 as a result of early forest removals (see Appen-
dix S1, Supporting information) and squares whose distance to
forest was unaffected, using a t-test and assuming unequal
variances.

OCCUPANCY MODELLING Results

We used generalized linear models (PROC GENMOD; SAS 9.2, SAS
Institute Inc., Cary, NC, USA), specifying a binomial error distri-
bution. The response variable was presence (1) or absence (0) of a
species in each 200-m square. For each species, all combinations
of the six explanatory variables were fitted, also including the
effort variable in all cases, to give a candidate set of 63 models.
Models were compared using an information theoretic approach,
with lower values of Akaike’s information criterion (AIC) indicat-
ing better models, and we considered any model with an Akaike
weight (wi) of at least 10% of that of the best approximating (low-
est AIC) model to be plausible (Burnham & Anderson 2002).
Where several models were plausible, we used multimodel infer-
ence, with parameters estimated as the mean of those models,
weighted by the Akaike weight of each model (Burnham & Ander-
son 2002). The weight of evidence for each explanatory variable
over all models was estimated by summing Akaike weights of all
models including that variable.
The data come from an array of grid squares; therefore, infer-
ence may be influenced by spatial covariance in the data, but
formal modelling of spatial structure in model residuals risks
under-emphasizing the importance of explanatory variables that
are themselves spatially autocorrelated (Diniz-Filho, Bini &
Hawkins 2003). We therefore calculated Moran’s I statistics for
the residuals of these final models to quantify any residual spatial
covariance that may affect model interpretation.
ASSESSING GOODNESS-OF-FIT
Dunlin were recorded in 8
1%, European golden plover in
9
6% and common greenshank in 3
7% of squares over
the four years of bird surveys.
For dunlin (wi = 0
666) and European golden plover
(wi = 0
804), the global model was 3
6 and 4
3 times better
supported, respectively, than any other. For common
greenshank, there was no clear best model, with ten hav-
ing at least 10% of the maximum wi of 0
186 (Table 2).
Model-averaged parameter estimates and variable weights
(Table 3) show that all three species were strongly associ-
ated with flat ground and pool systems, but showed
weaker associations with elevation and photosynthetic
intensity of vegetation (positive), and vegetation cover
(negative). In addition, there was a strong positive associ-
ation with distance to forest edge for dunlin and
European golden plover, but not common greenshank,
indicating avoidance of locations closer to forest edges for
the first two species. The best approximating models for
each species were well-fitting. Concordance indices were
0
817 (dunlin), 0
737 (European golden plover) and 0
813
(common greenshank), indicating ‘very good’ (dunlin,
greenshank) and ‘good’ (golden plover) discriminatory
ability (Vaughan & Ormerod 2005), especially given low
prevalence (Liu et al. 2005). The best approximating
models were also well calibrated; neither the slope nor the

The SAS LOGISTIC procedure was used to calculate the concor-

dance index (c) for the best approximating model for each Table 2. Measures of performance of all plausible models (those
species. This index indicates the probability that the fitted values within 10% of best model, for which ΔAIC = 0). Abbreviations
in first column from Table 1. ‘All’ = global model containing all
from the model will place all pairs of squares (one occupied, the
explanatory variables. Sixty-three models were considered for
other unoccupied in each pair) in the correct rank order of likeli-
each species
hood of predicted occupancy. Thus, chance performance is
clearly defined (c = 0
5) and perfect discrimination is indicated by Variables in model AIC ΔAIC Akaike weight
c = 1 (Vaughan & Ormerod 2005). Model calibration (accuracy
of predicted probabilities) for best approximating models was Dunlin
tested by logistic regression of observed presences and absences All 2904
2 0 0
666
in grid squares against the logit-transformed probabilities of r,n,t,f,p 2906
8 2
6 0
186
occurrence from the model (Pearce & Ferrier 2000). Model fits European golden plover
are well calibrated if intercepts are not significantly different from All 3568
6 0 0
804
n,e,t,f,p 3571
6 2
9 0
186
zero, and slopes not significantly different from one.
Common greenshank
n,e,s,p 1656
7 0 0
186
n,s,p 1656
8 0
1 0
178
PREDICTING BENEFITS OF FOREST REMOVAL
r,n,e,s,p 1657
7 1
0 0
112
For any species showing an additive effect of distance to forest r,n,s,p 1657
7 1
1 0
110
edge, we calculated fitted probabilities of occupancy first from n,e,s,f,p 1658
5 1
8 0
076
n,s,f,p 1658
6 1
9 0
073
model-averaged parameter estimates (Pocc), and then having set
All 1659
4 2
7 0
049
distance to forest for each square to the maximum observed
r,n,s,f,p 1659
4 2
7 0
048
value of 5
7 km (Pmax). Values of Pmax Pocc were then e,s,p 1660
0 3
4 0
035
mapped to show those parts of the study area where probabil- s,p 1660
9 4
2 0
022
ity of occupancy is predicted to increase most for the species

© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society, Journal of Applied Ecology, 51, 204–213
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208 J. D. Wilson et al.

Table 3. Model-averaged parameter estimates (b) and standard errors (SE) for each explanatory variable. The last column gives the
probability that each variable is present in the best model (pbest)

Dunlin European golden plover Common greenshank

Variable b SE pbest b SE pbest b SE pbest

Intercept 4
887 0
580 6
495 0
516 2
977 0
551

Effort 0
504 0
059 1 0
532 0
051 1 0
513 0
086 1
Reflectance 0
095 0
037 0
893 0
071 0
032 0
808 0
042 0
049 0
379
NDVI 2
226 0
843 0
916 2
644 0
798 0
990 2
171 0
926 0
881
Elevation 2
324 1
059 0
793 5
171 0
912 1 2
522 1
743 0
509
Exposure 0
157 0
016 1 0
085 0
012 1 – – –
Slope – – – – – – 0
233 0
091 0
946
Forest distance 0
508 0
055 1 0
448 0
049 1 0
032 0
065 0
295
Pool distance 0
342 0
035 1 0
154 0
034 1 0
446 0
046 1

close to forest edges, but at greater distances from

Fig. 2. Mean predicted probability of occupation of 200-m
squares by dunlin (filled bars) and European golden plover (open
bars) (Pocc) in increasing distance bands from forest edge. Each
distance band encompasses 10% of all squares. Predictions
assume survey visits in all 4 years to remove effects of variation
in survey effort.
intercept of the relationship between observed presences
and absences (y) and logit-transformed predicted occu-
pancy likelihoods were significantly different from 1 or
zero, respectively (P-values from 0
27 to 1 for Wald chi-
square test of slope and intercept for all three species).
Moran’s I values were 0
136 (dunlin), 0
106 (golden
plover) and 0
030 (greenshank) over lag distances of one
grid square, declining to 0
009, 0
015 and 0
002, respec-
tively, over lag distances of 10 grid squares. All these
values were statistically significant due to the very large
sample sizes, but values of the order of 0
1 or below sug-
gest that spatial autocorrelation effects in model residuals
are so small as not to compromise inference (Ryan et al.
2004; Kraan et al. 2009).
Based on model-averaged parameter estimates, Fig. 2
shows mean fitted probability of occupation of 200-m
squares (Pocc) by dunlin and European golden plover in
bands of increasing distance to forest edge. Incremental
changes in distance to forest edge are highly influential
forested areas, they have less impact that is additive to
the combined influence of other variables. The peak in
predicted probability of occurrence in the most distant
band arises because this 10% of squares is concentrated
in one area of high-quality blanket bog habitat and is one
of very few large contiguous areas far from forestry (Blar
nam Faoileag – area 1, Fig. 1). In other distance bands,
grid squares were widely distributed across the study area.
Variants on the final models in which distance to forest
edge is reduced to a binary variable (‘close’ vs. ‘far’), with
the threshold set at 100 m increments from 100 m to
1500 m, all have much poorer goodness-of-fit
(DAIC ≥ 16
2 for dunlin, and ≥23
1 for European golden
plover) than the continuous effect. However, the mini-
mum DAIC summed over both species is achieved when
the threshold is set at 700 m. This may be a useful ‘rule-
of-thumb’ minimum distance to set forest edges from
areas of otherwise high-quality habitat for these species in
the Flow Country.
Figure 3 plots values of Pmax Pocc to illustrate where
forest removal is predicted to result in greatest increases
in probability of occupancy by dunlin and European
golden plover. Squares for which distance to forest
increased between 1998 and 2003 following early forest
removals showed greater predicted increases in probability
of occupancy than others (dunlin 0
14  0
10SE vs.
0
08  0
08, t = 22
3, P < 0
0001; European golden plover
0
14  0
08 vs. 0
08  0
06, t = 26
8, P < 0
0001).
Discussion
Twenty-five years after breeding habitat associations of
peatland waders in the Flow Country were first studied
(Stroud et al. 1987; Stroud, Reed & Harding 1990;
Lindsay et al. 1988; Lavers & Haines-Young 1996;
Lavers, Haines-Young & Avery 1996); this study found
similar associations with topography, vegetation cover
and proximity to pool systems. However, with maturation
of plantation forests, there is now an additive effect of
distance to forest edge for dunlin and European golden

© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society, Journal of Applied Ecology, 51, 204–213

(a)
(b)
Fig. 3. Predicted absolute increase in probability of occupancy of
200-m grid squares by (a) dunlin, and (b) European golden plo-
ver if distance to forest edge is set for all squares to the maxi-
mum value in the data set (5
7 km). Colour scales continuously
from minimum value of zero at blue end of spectrum to maxi-
mum of 0
53 (dunlin) and 0
46 (European golden plover) at red
end of spectrum. Forestry cover (1998) shaded pale grey. Bound-
ary of SPA shown as black line.
plover, with reduced occupancy within several hundred
metres from forest edges.
We cannot be certain that reduced habitat occupation
close to mature forests is a demographic or behavioural
response to perceived or real predation risk. However,
nest predators such as hooded crows, red foxes and
pine martens Martes martes occupy plantation conifer
forests across Scotland (Avery & Leslie 1990; Chadwick,
Hodge & Ratcliffe 1997; Ratcliffe 2007). Studies else-
where have shown increased nest predation on breeding
waders close to forest edges, and a preference for nest-
ing on flat ground may be associated with the need for
clear views of approaching predators (e.g. Valkama,
Currie & Korpimaki 1999; Whittingham, Percival &
Brown 2002; Finney, Pearce-Higgins & Yalden 2005).
Equally, the lack of edge effect on common greenshank
habitat occupancy is consistent with the fact that this
species (like other Tringa species, but unlike dunlin and
European golden plover) vigorously mobs predators and
occupies bog habitats within native forests (Nethersole-
Thompson & Nethersole-Thompson 1979). However,
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Forest edge effects on peatland waders 209
explanations based on forest influences on bog habitat
condition (e.g. through alteration of hydrology, grazing
patterns, disease or fire risk) also need to be considered.
For example, hydrological and habitat effects of forestry
on adjacent blanket bog in the Flow Country extend up
to 40 m from plantation boundaries, at least in the first
few decades (Shotbolt, Anderson & Townend 1998),
and could affect prey availability for breeding birds.
Forestry also holds high densities of ticks Ixodes ricinus
relative to blanket bog (Gilbert 2013), and although the
form of the relationship between tick densities and distance
to forest edges remains unknown, ticks can infest chicks
of moorland-breeding waders (Newborn et al. 2009).
CONSERVATION AND MANAGEMENT IMPLICATIONS
AND WIDER POLICY RELEVANCE
Edge effects of forests on occupancy or nest success of
open-ground breeding birds are detected consistently over
distances of up to hundreds of metres (and occasionally
beyond) across a wide range of upland grassland, lowland
farmland and shrubland habitats in Eurasia and North
America (Table 4). This suggests that landscape planning
should consider buffer zones hundreds of metres wide in
addition to direct habitat loss when assessing impacts of
plantation forestry on open-ground habitats of bird con-
servation interest, or when prioritizing plantations for
removal and restoration of open-ground habitat. None-
theless, such consideration should be cautious and
species-dependent. For example, this study found evidence
of edge effects for two breeding wader species, but not a
third, whilst evidence of absence of forest edge effects on
habitat occupancy by corncrakes Crex crex (Berg &
Hiron 2012) may be explained by this species’ exception-
ally low susceptibility to nest predation (Green et al.
1997). Equally, some species, amongst which black grouse
Tetrao tetrix and common cuckoo Cuculus canorus are
European examples, may benefit directly from landscapes
comprising both forest and open-ground or the specific
resources offered by ecotone habitats (e.g. young-growth)
at edges (Fuller 2012; Grant & Pearce-Higgins 2012). In
such cases, the mix of habitats might be described posi-
tively as a ‘mosaic’, rather than negatively as a ‘fragmen-
tation’.
Forestry advisers and managers in our study area have
begun to use maps of the kind illustrated by Fig. 3, in
conjunction with recent advice on the negative GHG
implications of forestry planting on deep peat (Morison
et al. 2010), to identify areas that, once harvested, should
not be brought into a second tree-planting rotation. This
will add to the 2200 ha of forestry previously removed in
support of blanket bog restoration under EU LIFE
Nature programmes (Wilkie & Mayhew 2003). Our analy-
ses also show that these early forest removals were well
located with respect to predicted long-term benefits from
mitigation of edge effects (areas 2–5 in Fig. 1 compared
with Fig. 3). These initiatives are predicted to improve
 Table 4. Summary of studies of relationships between occupancy or breeding success of open-ground bird species and distance to forest edge. Previous studies in the Flow Country are discussed
in main text and not detailed again here. Studies dependent entirely on artificial nest experiments are not included due to the difficulties in interpreting results (Dolman 2012)

Open-ground
Study Focal open-ground species Location habitat affected Key findings

Parr (1992) European golden plover NE Scotland Upland heathland Poor breeding success associated with severe egg predation and
coincident with afforestation of study area, bringing all
territories within 2 km of forestry
210 J. D. Wilson et al.

Valkama, Currie & Eurasian curlew W Finland Tilled farmland 20% of nests hatched young in a landscape where random points
Korpimaki (1999) on average 180 m from forestry, but 91% where random points
on average 600 m from forestry
Finney, Pearce- European golden plover N England Blanket bog Probability of use of 100 m squares by adult birds and
Higgins & Yalden probability of nests fledging young increased with distance from
(2005) conifer forestry in landscape with forestry at 0–5 km from focal
squares and nests
Pearce-Higgins & Red grouse Lagopus l. scoticus, European golden S Scotland & Upland grassland, No correlations between bird abundance and forestry cover after
Grant (2006) plover, Eurasian curlew, common snipe Gallinago N England heathland and controlling for other habitat and landscape variables, but in a
gallinago, skylark Alauda arvensis, meadow pipit blanket bog study where survey plots had been chosen to have no forestry
Anthus pratensis, northern wheatear Oenanthe within 200 m of their boundaries
oenanthe, whinchat Saxicola rubetra, European
stonechat S.rubicola
Amar et al. (2011) European golden plover, northern lapwing, dunlin, UK Upland grassland, 50% forest cover within 1 km of survey plot associated with
Eurasian curlew, common snipe heathland and 85-95% decline of common snipe, dunlin and European golden
blanket bog plover. Absence of forest cover associated with population
stability
Douglas et al. Eurasian curlew S Scotland & Upland grassland, Population changes and fledging success negatively associated
(2013) N England heathland and with forestry cover within 1 km of edge of study plots
blanket bog
Berg & Hiron (2012) Corncrake S Sweden Mixed farmland No avoidance of forest edges, with 42% of territories within
100 m
Shochat, Abramsky Scrub specialists including desert lark Ammomanes Central Israel Semi-desert scrub Scrubland specialists lost from afforested landscapes where scrub
& Pinshow (2001) deserti, long-billed pipit Anthus similis, spectacled patch size falls below 50 ha (equivalent to a circle of radius
warbler Sylvia conspicillata 400 m)
Buchanan et al. Ring ouzel Turdus torquatus Scotland Upland grassland 2-km squares with up to 30% forest cover more likely to
(2003) experience declines over a decade than squares with no forest
cover, despite almost no change in forest cover, and after
accounting for other habitat variables
Rodewald & Vitz Shrubland specialists including blue-winged warbler S Ohio, United Shrubland Twice as many individuals of 7/8 shrubland specialist species
(2005) Vermivora cyanoptera, prairie warbler Dendroica States mist-netted 80 m from forest edges than 20 m from edges
discolor, yellow-breasted chat Icteria virens, indigo
bunting Passerina cyanea and field sparrow
Spizella pusilla
Reino et al. (2009) Calandra lark Melanocorypha calandra and S Portugal Tilled farmland Point count abundances increased 8- to 14-fold up to 300 m
short-toed lark Calandrella brachydactyla from forest edges for these two open-ground species

© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society, Journal of Applied Ecology, 51, 204–213
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habitat quality for dunlin and European golden plover,
and, in the long term, there may be additional benefits as
the cleared areas become suitable for re-occupation by
breeding waders. In a recent study of effects of removal
of tree and shrub rows on breeding grassland birds in
Wisconsin, increases in nesting density of eastern mead-
owlark Sturnella magna, Henslow’s Sparrow Ammodramus
henslowii and bobolink Dolichonyx oryzivorus were
recorded within just three years, alongside reductions in
activity of woodland-associated predators such as raccoon
Procyon lotor (Ellison et al. 2013). In our peatland study
areas, changes may be more gradual because deep plough-
ing at planting and persistence of felling brash make bog
vegetation recovery a slow process, and felled areas may
favour predators for several years until the brash breaks
down and is incorporated by re-expanding bog vegetation.
This may explain why removal of some forest blocks
between 1998 and 2003–2006 (Fig. 1) had little impact on
model goodness-of-fit (see Appendix S1, Supporting infor-
mation). Recent developments in harvesting machinery
and markets for whole-tree harvesting for wood fuel are
likely to mean that much less brash remains after future
fellings. We propose to continue monitoring of bird popu-
lations and vegetation change as restoration of formerly
forested areas continues, to test whether edge effects
reduce over time, as predicted, and whether this varies
with harvesting treatment.
It is important to improve the evidence on impacts of
plantation forest expansion on birds of high conservation
importance, and the mechanisms underlying these effects.
Forest species may benefit from the creation of wooded
habitats and increased connectivity of forest cover, whilst
open-ground species may suffer from habitat loss and
fragmentation. It is also timely to consider how forests
should be sited in the landscape, given the general support
for woodland expansion in support of climate change mit-
igation and other policy goals (Read et al. 2009; Quine
et al. 2011), set alongside increasing concern for declining
wader populations (e.g. Newton 2004). In Scotland, gov-
ernment policy aspires to increase woodland cover from
17% to 25% of the land, an addition of some 650 000 ha
over the 21st century (Scottish Government 2009) –
10 000 ha per annum of new afforestation over the next
ten years (WEAG 2012). Because of the high agricultural
value of lowland farmland (WEAG 2012), marginal
upland areas are likely to continue to be favoured for
such expansion. Many such areas are also of conservation
value (Thompson et al. 1995), so where expansion does
take place, the emerging edge effects on breeding waders
should be considered. Studies of predator populations and
behaviour, and bird distribution and demography on
adjacent open-ground habitats are clearly important.
Acknowledgements
Simon Hodge, Gordon Patterson, Ian Bainbridge and Stuart Housden
supported initiation of this research. Mark Brewer, Paul Britten, Graeme
© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society, Journal of Applied Ecology, 51, 204–213
13652664, 2014, 1, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1111/1365-2664.12173 by CochraneAruba, Wiley Online Library on [30/04/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Forest edge effects on peatland waders 211
Buchanan, David Fouracre, Cynthia Moore, Emma Teuten and Ellen
Wilson provided GIS and statistical support. Andrew Coupar, Pete May-
hew, John Risby and Lesley Cranna advised on conservation and manage-
ment of peatlands. Comments from Chris Elphick, Nils Warnock and an
anonymous reviewer greatly improved earlier drafts. We thank staff and
volunteers at Forsinard Flows Nature Reserve for bird survey data collec-
tion, especially Alasdair Boulton, Ian Dillon, Paul Jacques, Ewen Munro
and James Plowman.
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Forest edge effects on peatland waders 213

Wilkie, N.M. & Mayhew, P.W. (2003) The management and restoration Supporting Information
of damaged blanket bog in the north of Scotland. Botanical Journal of
Scotland, 55, 125–133. Additional Supporting Information may be found in the online version
Zurita, G., Pe’er, G., Bellocq, M.I. & Hansbauer, M.M. (2012) Edge
of this article.
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Received 25 June 2013; accepted 9 September 2013
Handling Editor: Chris Elphick

© 2013 The Authors. Journal of Applied Ecology © 2013 British Ecological Society, Journal of Applied Ecology, 51, 204–213