BIRD STUDY
2019, VOL. 66, NO. 4, 461–470
https://doi.org/10.1080/00063657.2020.1725420
Anthropogenic pressures within the breeding range of the Hen Harrier
Circus cyaneus in Ireland
Anthony Caravaggi a,b, Sandra Irwina, John Lusbyc, Marc Ruddockd, Allan Meee∗ , Tony Naglee∗∗ ,
Lorcán O’Toolee, Shane O’Neilld and John O’Halloran a
a
School of Biological, Earth and Environmental Sciences, University College Cork, Cork, Ireland; bSchool of Applied Sciences, University of South
Wales, Pontypridd, UK; cBirdWatch Ireland, Kilcoole, Co. Wicklow, Ireland; dGolden Eagle Trust Ltd, Dublin 2, Ireland; eIrish Raptor Study Group
(IRSG) Stramore, Glendowan, Churchill, Co. Donegal, Ireland
ABSTRACT
ARTICLE HISTORY
Capsule: Patterns in the frequency and co-occurrence of anthropogenic pressures associated with
suitable breeding habitat for Hen Harriers Circus cyaneus demonstrates the need for specific,
focussed management and policy options aimed at mitigating impacts on this threatened
population.
Aims: To describe anthropogenic pressures and threats in the upland breeding range of Hen
Harriers and to explore their potential impacts on the declining Hen Harrier population.
Methods: We used text mining, mixed-effects models, principal component analysis and clustering
methods to explore anthropogenic pressures on suitable breeding and foraging habitats for Hen
Harriers in Ireland, based on the 2015 national breeding Hen Harrier survey data.
Results: Mixed-effects models described a strong influence of agriculture, forestry, predator activity,
and recreational activities on survey areas that contained Hen Harrier territories. Cluster analyses
described three discrete pressure clusters and showed consistent co-occurrence of independent
pressures.
Conclusions: Areas of suitable habitat for Hen Harriers in the uplands overlap with areas that
experience anthropogenic pressures known to negatively impact on this vulnerable bird species.
Combined with clear evidence for the co-occurrence of multiple pressures at a regional scale,
this demonstrates a clear need for statutory agencies to consider the potential cumulative
impacts of individual pressures when developing conservation strategies for Hen Harriers.
Received 4 July 2019
Accepted 20 December 2019
Many species, worldwide, are threatened by anthropogenic
pressures that require intervention to mitigate or eliminate
their negative impacts (Wilcove et al.1998, Carroll et al.
2015, Di Minin et al. 2016). Such pressures can result in
stress responses or reduced fitness in wildlife that, in
some cases, have severe impacts on individuals or
populations (Wilcove et al.1998, Taylor & Knight 2003,
Johnson et al. 2005, Ciuti et al. 2012, Coetzee & Chown
2016). Conservation processes typically aim to prevent
species population declines and extinctions (Soule 1985).
However, conservation policy must also be cognizant of
the sustainable management of environmental resources
and other activities of economic and social importance
including commercial forestry, agriculture, and recreation
(Young et al. 2005, Kareiva & Marvier 2012, Kennedy
et al. 2016, Vangansbeke et al. 2017).
Human activities in the vicinity of breeding birds
can lead to increased rates of nest desertion (White &
Thurow 1985), and reduced rates of site occupancy
(Webber et al. 2013), territory establishment (Bötsch
et al. 2017), breeding success (Balotari-Chiebao et al.
2016), and survival (Ruhlen et al. 2003, including
illegal killing, e.g. Smart et al. 2010). Quantifying the
extent and ecological relevance of each of these
impacts informs our understanding of human–wildlife
interactions and underpins conservation and resource
management processes. It is essential, therefore, that
human activities that have the potential to affect
wildlife, particularly vulnerable species of conservation
concern, are properly assessed and understood, so that
appropriate measures can be developed to facilitate
conservation and sustainable land and resource use.
Hen Harriers Circus cyaneus are medium-sized
raptors that nest largely in upland areas, preferentially
in heather moorland (Redpath et al.1998, Barton et al.
2006, Amar et al. 2008, Ruddock et al. 2012, 2016,
CONTACT John O’Halloran
j.ohalloran@ucc.ie
School of Biological, Earth and Environmental Sciences, University College Cork, Distillery Field, North Mall,
Cork, T23 XA50, Ireland
∗
Present address: RaptorLIFE, IRD Duhallow, Newmarket, Co. Cork, Ireland.
∗∗
Present address: The Rookery, Ballyfeard, Minane Bridge, Co. Cork, Ireland.
Supplemental data for this article can be accessed at https://doi.org/10.1080/00063657.2020.1725420.
© 2020 British Trust for Ornithology
462
A. CARAVAGGI ET AL.
Watson 2017), during the breeding season. Upland
habitats in Ireland have been subjected to degradation
and land-use change. As a result of a large-scale
afforestation programme in the Republic of Ireland
from the 1950s and the conversion of ‘traditional’ open
habitats to forest, Hen Harriers in Ireland are also
frequently associated with young (i.e. pre-thicket)
conifer plantations that provide them with areas for
nesting and foraging (Wilson et al. 2009, Irwin et al.
2012, Wilson et al. 2012, Ruddock et al. 2016).
Anthropogenic impacts such as afforestation and forest
management (NPWS 2015), landscape degradation and
land-use change (Wilson et al. 2009, 2012), livestock
grazing (O’Rourke & Kramm 2009), illegal burning
(Renou-Wilson et al. 2011), peat extraction (O’Riordan
et al. 2015), recreation (Hynes & Buckley 2007), and
wind energy development (Wilson et al. 2017) could
have important implications for breeding Hen Harriers
in Ireland. While some mortality (via poisoning/
shooting) has been recorded, the level of persecution
observed in Britain (Redpath et al. 2010, Murgatroyd
et al. 2019) has not been frequently observed in
Ireland, perhaps as there are no areas that are managed
solely for driven grouse shooting. Moreover, red grouse
(Lagopus lagopus scotica) are red-listed species of
conservation concern in Ireland (National Red Grouse
Steering Committee 2013) and there is a widespread,
self-imposed moratorium on grouse shooting by
shooting fraternity. However, raptors are known to
migrate within the British Isles (Mead, 1973) and
persecution of Hen Harriers in Britain could have
hitherto undescribed impacts on the Irish population.
The Hen Harrier population in Ireland is of national
conservation concern (Colhoun & Cummins 2013),
with a population of between 108 and 157 breeding
pairs recorded in the most recent national survey
(Ruddock et al. 2016). The species is listed under
Annex I of the European Commission Birds Directive
(2009/147/EC) that requires Member States to
designate Special Protection Areas (SPAs) for their
survival and reproduction. Six Hen Harrier SPAs
containing important breeding areas for the species
were designated in Ireland in 2007.
Hen Harrier monitoring and conservation research in
Ireland to date has focussed on national population
estimates (Barton et al. 2005, Ruddock et al. 2012,
2016), the impacts of afforestation (Irwin et al. 2012,
Wilson et al. 2009, 2012) and wind farm development
(Fernández-Bellon et al. 2015, Wilson et al. 2017) on
their populations, as required to inform conservation
management. Due to the targeted nature of previous
research, very little information is available in the
published literature regarding the broader range of
anthropogenic pressures that might impact breeding
Hen Harriers and associated foraging and breeding
habitat. Furthermore, previous research has considered
how individual pressures impact separately and in
specific contexts while consideration of the synergies
between pressures is lacking. To address these gaps, we
explored data on anthropogenic pressures affecting
Hen Harriers within their breeding range in Ireland,
with the aim of deriving information that would
inform conservation and management processes for
this threatened species.
Methods
The 2015 National Survey of Breeding Hen Harriers in
Ireland was conducted between March and September
2015 in suitable Hen Harrier habitat in upland areas,
largely, but not exclusively, between 200 and 600 m
above sea level and within the Hen Harrier breeding
range (Ruddock et al. 2016). Survey squares of 10 km2
(n = 268) were defined using the Irish National Grid
(Figure 1(a)). Anthropogenic activities that could
potentially impact on breeding Hen Harriers
(‘pressures’ from hereon) were recorded from vantage
points within each survey square during each of 4–6
dedicated watches per square, during the breeding
season. Where sites were occupied, vantage points were
a minimum of 500 m from nests sites. Vantage points
were identified a-priori based on habitat suitability,
topographical constraints, and the potential for
observers to cause disturbance to breeding birds
(Ruddock & Whitfield 2007, Whitfield et al. 2008).
Hen Harrier territories (n = 100, across 54 survey
squares) were recorded where identified; occupancy
was based on observations of Hen Harrier breeding
behaviour and the repeated presence of birds (Ruddock
et al. 2016). Data were collected by staff, members, and
volunteers from the National Parks & Wildlife Service
(NPWS), BirdWatch Ireland (BWI), Irish Raptor Study
Group (IRSG), Golden Eagle Trust (GET), university
researchers, and independent commercial and
voluntary ornithological surveyors.
Pressures were divided into 47 discrete categories
(online Table S1) aligned with the European
Commission Birds Directive (2009/147/EC) reporting
matrix. The frequency of occurrence of each pressure
within 2 km of vantage point locations was recorded
within each survey square. Initial exploration of the
data revealed extreme outliers, therefore we adopted a
precautionary approach and applied consistent
thresholds throughout. Values for individual pressures
that occurred beyond two standard deviations (SD)
from the mean were replaced with the maximum value
BIRD STUDY
463
Figure 1. Maps of Ireland showing: (a) survey squares and SPAs (as indicated by grey polygons), (b) the total number of confirmed Hen
Harrier territories per square, and (c) pressure indices derived from cumulative observations of pressures within each survey square. See
online Table S2 for square-specific data.
as defined by the aforementioned threshold, rounded to
the nearest whole integer. This allowed us to capture the
prevalence of each pressure at each location while
mitigating over-inflation. The sum frequency of each
pressure was calculated: (i) across all survey squares
where the total number of recorded pressures was
greater than zero (n = 146; online Table S2); (ii) across
squares located within SPA boundaries only (n = 24);
and (iii) across squares where confirmed Hen Harrier
territories were present (n = 54). It was necessary to
account for variation in survey effort as the number of
visits made to vantage points varied between observers.
Therefore, a Pressure Index (PI) was created, where the
total number of pressures was divided by the total
number of visits (Ruddock et al. 2016). PI scores were
normalized between 0 and 1 to facilitate comparisons
between sites. General linear models were used to
investigate differences between PI scores – with zero
counts removed and remaining data log transformed to
meet model assumptions – where PI was the
dependent variable and the location of vantage points
relative to SPA boundaries (inside/outside) and
confirmed Hen Harrier territories (present/absent)
were explanatory variables. Models explored each
category (SPA boundaries and Hen Harrier territories)
independently as well as part of a fully-factorial model
that included an interaction term.
Principal component analysis (Jolliffe & Cadima
2016) and linear mixed-effects models were used to
investigate relationships between the presence/absence
of Hen Harrier territories and pressure categories. Data
were Box Cox transformed to remove skewness,
centred, and standardized to have mean = 0 and
standard deviation = 1 prior to analysis. Principal
components (PCs) that cumulatively accounted for
over 50% of the variance were retained for inclusion in
models. The presence/absence of Hen Harrier
territories (Figure 1(b)) was entered as a binary
dependent variable, retained PCs were included as
explanatory variables and surveyor identity was
included as a random variable. Model permutations
were ranked using the Akaike Information Criterion
(AIC); the top subset of models was found within
ΔAIC ≤ 2 units (Burnham & Anderson 2002).
Cluster analysis was used to quantify associations
between individual pressure categories across all survey
squares. The various methods that comprise cluster
analyses provide a means of classifying multivariate
data into subgroups according to the similarity of their
attributes, thus revealing the underlying structure
(Everitt et al. 2009). We calculated the distance of each
recorded pressure from the cluster’s mean using a
Euclidean distance index and applied the Ward error
sum of squares hierarchical clustering method (Ward
1963) to the resultant data. The optimal number of
clusters (kt) was identified using average silhouettes
(Kaufman & Rousseeuw 1990) and Approximately
Unbiased (AU) P-values with multiscale bootstrap
resampling (B = 10 000) where clusters with P ≥ 0.95
were strongly supported (Suzuki & Shimodaira 2006).
All data analyses and plotting were carried out using
the statistical programme R (R Core Team 2017),
specifically the packages cluster (Maechler et al. 2018)
and pvclust (Suzuki & Shimodaira 2015), dendextend
(Galili 2015), nlme (Pinheiro et al. 2017), and caret
(Kuhn 2017). Data are subject to data-sharing
agreements and, therefore, cannot be redistributed.
However, R code used for data exploration and
analyses are available at http://doi.org/10.5281/zenodo.
3549584.
464
A. CARAVAGGI ET AL.
Results
A total of 2873 individual pressure occurrences were
recorded during this study. There were no
anthropogenic pressures recorded in 45% of survey
squares. The most frequently recorded pressures across
all survey squares were forest management and use
(13% of occurrences), paths, tracks, forest roads (11%),
uncontrolled burning (6%), and wind energy production
(6%). Similar pressures were recorded inside and
outside of SPA boundaries: forest management and use
(14% and 11% of occurrences, respectively), paths,
tracks, forest roads (10%, 0%), forest planting on open
ground (0%, 8%), uncontrolled burning (6.6%, 7%), and
wind energy production (9.5%, 7%). The most
frequently recorded pressures associated with
confirmed Hen Harrier territories were loss of habitat
features (13.7%), dispersed habitation (10.5%), paths,
tracks, forest roads (9.2%), and forest management
(8.1%). In contrast, pressures at vantage points where
there were no Hen Harrier territories were forest
management and use (16%), off-road motorized driving
(12%), forest planting on open ground (11%), and
mechanical removal of peat (11%).
Pressure indices varied between survey squares
(Figure 1(c)) and only one survey square had a PI >
0.5. Survey squares where vantage points occurred
within SPAs had a maximum PI of 0.22 (mean ± SD =
0.08 ± 0.07), which was significantly higher than those
outside SPAs (t = 0.028; β ± SD = −0.44 ± 0.20; P =
0.03). Survey squares where vantage points were
associated with Hen Harrier territories had a
maximum PI of 0.42 (mean ± SD = 0.10 ± 0.09; online
Table S1), which was significantly higher than those
that were not associated with territories (t = 0.038; β ±
SD = −0.39 ± 0.19; P = 0.04; Table 1).
Both silhouette and AU clustering methods supported
three discrete clusters (P ≥ 0.05). The largest cluster
(ii) consisted of 25 pressure categories while the smallest
(iii) was the most distinct and consisted of 5 pressure
categories. One sub-cluster was statistically supported (iv)
and was comprised of 17 pressure categories (Figure 2).
Table 1. General linear model results for regional differences in
pressures on Hen Harrier breeding habitat – expressed as a
Pressure Index (PI; log transformed). s = Special Protection
Areas (SPA; inside/outside); r = confirmed territories (present/
absent). Regression coefficients (β ± SE) and significance of
contributory variables are given, where ∗ P < 0.05.
Model
PI ∼ r
PI ∼ s
PI ∼ r + s + r∗ s
Variable
T
r
s
s
r
0.038
0.028
−0.515
−0.683
−1.128
s∗ r
β (±SE)
−0.39
−0.44
−0.14
−0.16
−0.46
±
±
±
±
±
0.19∗
0.20∗
0.28
0.23
0.40
A total of seven principal component axes, accounting
for >50% of the total variance, were retained for
inclusion in mixed-effects models investigating the
relationship between the presence/absence of Hen
Harrier territories and associated pressures. The top
subset of models (ΔAIC ≤ 2) included PC1, PC2, and
PC3. PC1 accounted for the greatest proportion of
total variance (20%); loadings were most strongly
weighted towards aspects of agricultural and forestry
activity and predators; PC2 (8%) was weighted towards
forest management and site access; and PC3 was
weighted towards forest clearance and recreational
activities (Table 2). The best approximating model was
positively influenced by PC1 and PC3, and negatively
influenced by PC2 (Table 2). It should be noted that
PC1 includes nest destruction, predation by birds, and
predation by mammals. These pressures can only occur
where Hen Harriers nest, hence the observed positive
association is to be expected.
Discussion
The results show that suitable Hen Harrier breeding
habitats in Ireland are subjected to a wide range of
anthropogenic pressures that could have significant
implications for this vulnerable species. The number
and variety of pressures recorded demonstrates the
potential for direct disturbance of Hen Harriers
throughout the breeding season. The issue is
potentially exacerbated by the fact that and Hen
Harriers are frequently recorded to over-winter in
these same upland areas (O’Donoghue, 2010).
Furthermore, the co-occurrence of pressures as
described by cluster analyses demonstrates the
considerable potential for cumulative effects.
Anthropogenic impacts are not homogenous in their
severity or extent. This is certainly true in the current
study, where some pressures will have more severe
consequences for Hen Harriers or will act at different
spatial scales. However, there is a dearth of quantitative
data on the impacts of described pressures on Hen
Harriers. Our results highlight the importance of
managing pressures in an integrated manner rather
than on an individual basis. This provides support for
the effective management of suitable breeding areas to
minimize the potential impact of anthropogenic
pressures on vulnerable Hen Harrier populations.
Planted forests and the presence of tracks or roads
were recorded at high frequencies in all survey squares
across Ireland. Large areas of Irish upland habitat have
been afforested in recent decades and total forest cover
is expected to continue to increase from the current
11% to as much as 18% in the next 30 years (NPWS
BIRD STUDY
465
Figure 2. Relationships between pressures associated with potentially suitable breeding habitat for Hen Harriers. Pressure codes are
taken and descriptions abbreviated from those given in Ruddock et al. (2016). Dashed grey rectangles indicate outermost clusters
identified via the silhouette method and multiscale bootstrapping (10 000 iterations; Approximately Unbiased P ≤ 0.05). ○ =
clusters supported at AU P ≤ 0.05. For detailed pressure definitions, see online Table S1.
2015). In the absence of their traditional open heath and
blanket bog habitat, Hen Harriers in Ireland are
frequently associated with young (i.e. pre-thicket)
conifer plantations that provide areas that Hen
Harriers use for nesting and foraging (Wilson et al.
2009, Irwin et al. 2012, Wilson et al. 2012, Ruddock
et al. 2016). Hen Harriers cannot use closed-canopy
forests for breeding or foraging, therefore the
maturation of the existing forest estate threatens to
deprive Hen Harriers of already scarce breeding
habitat, while further increases in forest cover could
also lead to increased habitat fragmentation and
subsequently reduce the capacity of the landscape to
support breeding pairs. Recreational activities were also
strongly associated with survey squares containing Hen
Harrier
territories.
Systematic
reviews
have
demonstrated that recreational activities can negatively
impact breeding birds (Steven et al. 2011, Larson et al.
2016) including above-ground foragers (Bötsch et al.
2017). Thus, there exists the potential for the direct
disturbance of Hen Harriers throughout the breeding
season.
466
A. CARAVAGGI ET AL.
Table 2. Linear mixed-effects model results for pressures – expressed as PC – associated with confirmed Hen Harrier territories
(present/absent). Models were evaluated according to their AIC value. Factors retained in the top subset of n models (<Δ2 AIC) are
highlighted. Constituent pressures along with pressure codes and associated loadings (coefficients in parentheses) are given.
Pressure codes are taken and descriptions are abbreviated from those given in Ruddock et al. (2016; see online Table S1).
Regression coefficients (β ± SE) and significance of contributory PCs are given, where ∗ P < 0.05, ∗∗ P < 0.01, and ∗∗∗ P < 0.001. For
constituent pressures in PC4-7, see online Table S3.
Principal Component (% variance explained)
PC1
(21%)
PC2
(10%)
PC3
(7%)
Pressure
β
Abandoned pastoral systems (A7; 0.29)
Removal of hedges, copse and scrub (A9; 0.29)
Forest replanting (B3; 0.27)
Nest destruction (F1; 0.29)
Controlled burning (J2; 0.27)
Predation by birds (K1; 0.31)
Predation by mammals (K2; 0.26)
Forest management and use (B2; −0.28)
Forest clearance (B4; −0.30)
Thinning of tree layer (B5; −0.26)
Paths, tracks, forest roads (D1; −0.35)
Roads, motorways (D2; −0.33)
Natural Fires (J1; −0.30)
Dispersed habitation (E2; −0.32)
Outdoor sport, leisure and recreation (G2; −0.35)
Walking, horse-riding, cycling (G3; −0.42)
Motorized vehicles (G4; −0.29)
Off-road driving (G5; −0.32)
Other outdoor sports and leisure (G6; −0.30)
PC4
PC5
PC6
PC7
Mammalian and avian predators were among the
factors strongly associated with Hen Harrier territories.
O’Donoghue (2010) attributed 55% of all nest failures
in south and west Ireland in 2007 and 2008 to
predation events and Red Foxes (Vulpes vulpes) have
been observed depredating Hen Harrier chicks via
remote-sensing camera traps (Irwin et al. 2012,
Fernández-Bellon et al. 2018a). Other potential
predators of Hen Harrier nests in Ireland include Pine
Marten Martes martes, American Mink Neovison vison,
Stoat Mustela erminea, Raven Corvus corax, and
Hooded Crow Corvus cornix (Picozzi 1984, FernándezBellon et al. 2018a). These predators can have
substantial negative impacts on ground-nesting birds
(Paton 1994) as eggs and young chicks are particularly
vulnerable to predation when parents are absent.
Populations of generalist predators may be bolstered by
changes in land-use and management, including
afforestation and other forms of habitat fragmentation
(Prestt 1965, Hayden & Harrington 2000, Chalfoun
et al. 2002, Twining et al. 2019). However, data on the
abundance and activity of upland predators in Ireland
are scarce; efforts to investigate such may be of
considerable benefit to the conservation of Hen Harriers.
It is notable that wind energy production was
recorded more frequently within SPA boundaries than
outside but was rarely recorded in survey squares that
contained breeding Hen Harriers (3.3%) in this study,
perhaps indicating avoidance of wind farms for
± SE
t
0.025
0.007
3.52
−0.046
0.009
−5.05
∗∗∗
0.044
0.011
−3.91
∗∗
0.010
−0.004
−0.005
0.006
0.013
0.014
0.014
0.015
0.78
−0.27
−0.38
0.41
∗∗
breeding purposes (see also Wilson et al. 2017).
Indeed, windfarm construction activity has been
implicated in the desertion of traditional breeding sites
in Ireland (O’Donoghue et al. 2011). The construction
and operation of wind turbines can have both lethal
and sub-lethal impacts on birds (Drewitt & Langston
2006, Marques et al. 2014, Balotari-Chiebao et al. 2016,
Smith & Dwyer 2016, Fernández-Bellon et al. 2018b,
Thaker et al. 2018). The Republic of Ireland is
committed to European Union targets on renewable
energy including a national target of 40% electricity
from renewables by 2020, which is likely to involve the
construction of additional wind farms (DCCAE 2010).
Wind energy developments tend to be upland-focussed
and future, large-scale expansion may pose a threat to
breeding Hen Harriers. A bird sensitivity mapping tool
has been developed to guide the siting of future wind
energy developments in Ireland in relation to the
distribution of species of conservation concern,
including Hen Harrier (McGuinness et al. 2015).
However, there is as yet no mandatory obligation on
developers to use this tool.
The timing of disturbance events may be a key
consideration and many sources of disturbance may
already be present at the onset of breeding, when pairs
are establishing territories, potentially resulting in nest
abandonment. Other pressures such as peat extraction
or illegal burning may not occur until after laying and,
hence, can impact on parental care and, ultimately,
BIRD STUDY
breeding success. Such activities also effectively sterilise
breeding habitat in the longer-term. Current mitigation
measures for Hen Harriers in Ireland adopt a reactive
approach where circular ‘High Likelihood Nesting
Areas’ (HNLA, formerly Red Areas) of high sensitivity
to Hen Harriers, that contain nesting pairs and with a
radius of 1.2 km, are added to the HNLA network
when new breeding pairs are identified (NPWS 2015).
Some forestry operations that may cause disturbance
are regulated within these HNLAs during the breeding
season (Forest Service 2012). However, the protection
afforded by HNLAs only applies to known pairs within
the SPA network. Therefore, all other pairs that are
outside of the SPA network (>50% of the breeding
population; Ruddock et al. 2016) remain vulnerable to
direct disturbance from forest management activities
during the nesting season. Moreover, breeding Hen
Harriers have been recorded travelling as far as 11 km
from active nests (Irwin et al. 2012, Arroyo et al. 2014)
and human activities and impacts in the wider
landscape can have impacts on the physiology (Abbasi
et al. 2017) and mortality (Ferrer & Hiraldo 1993) of
birds. Human activities within the foraging range of
breeding Hen Harriers could result in patch avoidance
and/or stress-related responses in foraging birds,
potentially keeping them away from the nest for longer
periods of time and subsequently increasing chick
vulnerability. It is possible that Hen Harriers in Ireland
have the capacity to develop a tolerance for human
activities during the breeding season, though this may
be highly dependent on temporal and spatial variation
of disturbance sources (Ruddock & Whitfield 2007,
Whitfield et al. 2008). However, given their small
population size and conservation status, the
precautionary principle suggests that human activities
should be strictly regulated in areas of suitable Hen
Harrier breeding and foraging habitat, particularly
during key breeding months. Furthermore, pressures
and their potential impacts on breeding Hen Harriers
must be placed in a broader context that includes the
timing of pressure occurrence, the composition of the
wider landscape, and the conservation of suitable habitat.
The pressures described herein represent potential
disturbances to Hen Harriers throughout their breeding
cycle and therefore may have important consequences
for long-term population persistence or recovery. Recent
research suggests that the same pressures impact
another upland bird of prey, the Short-eared Owl Asio
flammeus, across their European range (FernándezBellon, unpubl. data). Thus, we recommend the
following actions to enhance conservation benefits for
Hen Harriers and other sensitive upland species and
habitats: (i) restrict forestry activities within the known
467
Hen Harrier range during the Hen Harrier breeding
season (March–August) by using targeted surveys to
detect Hen Harrier presence, thereby ensuring that
forest management activities can be undertaken in areas
that do not hold Hen Harriers during the summer
months; (ii) review and identify risks during the
wintering season for Hen Harriers, particularly as many
of the upland breeding sites can be used by roosting
birds during winter; (iii) quantify the abundance and
activity of upland predators and explore options for
predator control, where appropriate; (iv) avoid
recreation and non-licensed forestry-related activities in
areas known to hold Hen Harriers, throughout the
breeding season, supported by a programme of
community engagement, awareness-raising and upland
signage; and (v) improve lines of communication
between stakeholders so that potentially disturbing or
damaging activities can be identified at the earliest stages.
Failure to mitigate anthropogenic disturbances in
upland areas of potentially suitable Hen Harrier
breeding habitat, whether inside or outside of SPAs,
could have negative consequences for this already
vulnerable population. To date, none of the SPAs in the
Hen Harrier Natura 2000 network possess management
plans, one of the key requirements of such sites, over a
decade on from designation in 2007, and a Hen Harrier
Threat Response Plan, initiated by the National Parks &
Wildlife Service in the Republic of Ireland in 2016 with
wide stakeholder consultation, has yet to be published.
Furthermore, connecting multiple pressures is a key
issue for conservation management, and Hen Harrier
conservation policies must comprehensively account for
cumulative anthropogenic impacts at regional level.
Successful mitigation and management would represent
a significant step towards the conservation of Hen
Harriers in Ireland and serve as an example for upland
conservation initiatives in Europe.
Acknowledgments
The authors thank the many people who collected data for the
2015 National Survey of Breeding Hen Harrier in Ireland
which was undertaken by the Golden Eagle Trust, Irish
Raptor Study Group and Birdwatch Ireland with National
Parks & Wildlife Service (NPWS). Authors also extend
thanks to members of the stakeholder and scientific steering
groups for the Supporting Hen Harriers in Novel
Environments (SHINE) research project at UCC. We also
thank the Editor and two anonymous reviewers whose
feedback helped improve this manuscript.
Funding
The SHINE research project was funded by the Department of
Agriculture, Food and the Marine.
468
A. CARAVAGGI ET AL.
ORCID
Anthony Caravaggi http://orcid.org/0000-0002-1763-8970
John O’Halloran http://orcid.org/0000-0002-8150-7510
References
Abbasi, N.A., Arukwe, A., Veerle, L.B.J., Eulaers, I.,
Mennilo, E., Ibor, O.R., Frantz, A., Covaci, A. & Malik,
R.N. 2017. Oxidative stress responses in relationship to
persistent organic pollutant levels in feathers and blood of
two predatory bird species from Pakistan. Sci. Total.
Environ. 580: 26–33.
Amar, A., Arroyo, B., Meek, E., Redpath, S. & Riley, H. 2008.
Influence of habitat on breeding performance of Hen
Harriers Circus cyaneus in Orkney. Ibis 150: 400–404.
Arroyo, B., Leckie, F., Amar, A., McCluskie, A. & Redpath,
S. 2014. Ranging behaviour of Hen Harriers breeding in
special protection areas in Scotland. Bird Study 61: 48–55.
Balotari-Chiebao, F., Brommer, J.E., Niinimäki, T. &
Laaksonen, T. 2016. Proximity to wind-power plants
reduces the breeding success of the white-tailed eagle.
Anim. Conserv. 19: 265–272.
Barton, C., Pollock, C., Norriss, D.W., Nagle, T., Oliver,
G.A. & Newston, S. 2006. The second national survey of
breeding Hen Harriers Circus cyaneus in Ireland. Irish
Birds 8: 1–20.
Bötsch, Y., Tablado, Z. & Jenni, L. 2017. Experimental
evidence of human recreational disturbance effects on
bird-territory establishment. Philos. Trans. R. Soc.
Lond. B. Biol. Sci. 284: 20170846.
Burnham, K. & Anderson, D. 2002. Model Selection and
Multi-Model Inference: a practical information-theoretic
approach. Springer, New York.
Carroll, C., Rohlf, D.J., Li, Y.-W., Hartl, B., Phillips, M.K. &
Noss, R.F. 2015. Connectivity conservation and endangered
species recovery: a study in the challenges of defining
conservation-reliant species. Conserv. Lett. 8: 132–138.
Chalfoun, A.D., Thompson, F.R. & Ratnaswamy, M.J. 2002.
Nest predators and fragmentation: a review and metaanalysis. Conserv. Biol 16: 306–318.
Ciuti, S., Northrup, J.M., Muhly, T.B., Simi, S., Musiani, M.,
Pitt, J.A., Boyce, M.S. & Moreira, N. 2012. Effects of
humans on behaviour of wildlife exceed those of natural
predators in a landscape of fear. PLOS ONE 7: e50611.
Coetzee, B.W.T. & Chown, S.L. 2016. A meta-analysis of
human disturbance impacts on Antarctic wildlife. Biol.
Rev. 91: 578–596.
Colhoun, K. & Cummins, S. 2013. Birds of conservation
concern in Ireland. Irish Birds 9: 523–544.
Department of Communications, Climate Action &
Environment [DCCAE]. 2010. The National Renewable
Energy Action Plan: submitted under Article 4 of Directive
2009/28/EC. Department of Communications, Climate
Action & Environment, Dublin.
Di Minin, E., Slotow, R., Hunter, L.T.B., Montesino
Pouzols, F., Toivonen, T., Verburg, P.H., LeaderWilliams, N., Petracca, L. & Moilanen, A. 2016. Global
priorities for national carnivore conservation under land
use change. Sci. Rep. 6: 23814.
Drewitt, A.L. & Langston, R.H.W. 2006. Assessing the
impacts of wind farms on birds. Ibis 148: 29–42.
Everitt, B., Landau, S. & Leese, M. 2009. Cluster Analysis. 4th
ed. Arnold , Oxford University Press, London.
Fernández-Bellon, D., Irwin, S., Wilson, M. & O’Halloran, J.
2015. Reproductive output of Hen Harriers Circus cyaneus
in relation to wind turbine proximity. Irish Birds 10: 143–
150.
Fernández-Bellon, D., Wilson, M., Irwin, S., Kelly, T.C.,
O’Mahony, B. & O’Halloran, J. 2018a. Video evidence of
siblicide and cannibalism, movement of nestlings by
adults, and interactions with predators in nesting hen
harriers. J. Raptor Res. 52: 393–399.
Fernández-Bellon, D., Wilson, M., Irwin, S. & O’Halloran, J.
2018b. Effects of development of wind energy and
associated changes in land use on bird densities in upland
areas. Conserv. Biol. 33: 413–422.
Ferrer, M. & Hiraldo, F. 1993. Evaluation of management
techniques for the Spanish Imperial Eagle. Biol. Conserv.
63: 436–442.
Forest Service. 2012. Forest Service Appropriate Assessment
Procedure. Appendix C: Appropriate Assessment Procedure
(AAP) Requirements Regarding Hen Harrier SPAs, Felling
and Other Disturbance Operations. Forest Service, Dublin.
Galili, T. 2015. Dendextend: an R package for visualizing,
adjusting, and comparing trees of hierarchical clustering.
Bioinformatics. 31: 3718–3720.
Hayden, T. & Harrington, R. 2000. Exploring Irish Mammals.
Town House and Country House Ltd, Dublin.
Hynes, S. & Buckley, C. 2007. Recreational pursuits on
marginal farm land: a discrete-choice model of Irish farm
commonage recreation. Econ. Soc. Rev. 38: 63–84.
Irwin, S., Wilson, M., O’Donoghue, B., O’Mahony, B., Kelly,
T. & O’Halloran, J. 2012. Optimum Scenarios for Hen
Harrier Conservation in Ireland. Department of
Agriculture, Food and the Marine by the School of
Biological, Earth and Environmental Sciences, University
College Cork, Cork.
Johnson, C.J., Boyce, M.S., Case, R.L., Cluff, H.D., Gau, R.J.,
Gunn, A. & Mulders, R. 2005. Cumulative effects of human
developments on Arctic wildlife. Wildlife Monogr. 160:
1–36.
Jolliffe, I.T. & Cadima, J. 2016. Principal component analysis:
a review and recent developments. Philos. Tran. R. S. A. 374:
20150202.
Kareiva, P. & Marvier, M. 2012. What is conservation
science? BioScience 62: 962–969.
Kaufman, L. & Rousseeuw, P.J. 1990. Finding Groups in Data.
John Wiley & Sons, Inc., Hoboken, NJ.
Kennedy, C.M., Miteva, D.A., Baumgarten, L., Hawthorne,
P.L., Sochi, K., Polasky, K., Oakleaf, J.R., Uhlhorn, E.M.
& Kiesecker, J. 2016. Bigger is better: improved nature
conservation and economic returns from landscape-level
mitigation. Sci. Adv. 2: e1501021–e1501021.
Kuhn, M. 2017. caret: Classification and regression training.
Available at: https://CRAN.R-project.org/package=caret.
Larson, C.L., Reed, S.E., Merenlander, A.M., Crooks, K.R. &
Doi, H. 2016. Effects of recreation on animals revealed as
widespread through a global systematic review. PLOS
ONE 11 (12): e0167259.
Maechler, M., Rousseuw, P., Struyf, A., Hubert, M. &
Hornik, K. 2018. cluster: Cluster ANALYSIS BASICS AND
EXTENsions. Available at: https://CRAN.R-project.org/
package=rgdal.
BIRD STUDY
Marques, A.T., Batalha, H., Rodrigues, S., Costa, H., Pereira,
M.J.R., Fonseca, C., Mascarenhas, M. & Bernardino, J.
2014. Understanding bird collisions at wind farms: An
updated review on the causes and possible mitigation
strategies. Biol. Conserv. 179: 40–52.
McGuinness, S., Muldoon, C., Tierney, N., Cummins, S.,
Murray, A., Egan, S. & Crowe, O. 2015. Bird Sensitivity
Mapping for Wind Energy Developments and Associated
Infrastructure in the Republic of Ireland. BirdWatch
Ireland, Kilcoole.
Mead, C.J. 1973. Movements of British raptors. Bird Study 20:
259–286.
Murgatroyd, M., Redpath, S.M., Murphy, S.G., Douglas, D.J.,
Saunders, R. & Amar, A. 2019. Patterns of satellite tagged hen
harrier disappearances suggest widespread illegal killing on
British grouse moors. Nat. Commun. 10: 1094.
National Parks & Wildlife Service [NPWS]. 2015. Hen
Harrier Conservation and the Forestry Sector in Ireland.
National Parks & Wildlife Service, Dublin.
National Red Grouse Steering Commitee. 2013. Red Grouse
Species Action Plan 2013. University College Dublin,
Dublin, Ireland.
O’Donoghue, B. 2010. The Ecology and conservation of Hen
Harriers (Circus cyaneus) in Ireland. PhD thesis, University
College Cork.
O’Donoghue, B., O’Donoghue, T.A. & King, F. 2011. The Hen
Harrier in Ireland: conservation issues for the 21st century.
Biol. Env. Proc. R. Irish Acad. 111B: 83–93.
O’Riordan, M., Mahon, M. & McDonagh, J. 2015. Power,
discourse and participation in nature conflicts: the case of
turf cutters in the governance of Ireland’s raised bog
designations. J. Env. Policy Plann. 17: 127–145.
O’Rourke, E. & Kramm, N. 2009. Changes in the
management of the Irish uplands: a case-study from the
Iveragh Peninsula. Eur. Countryside 1: 53–66.
Paton, P.W.C. 1994. The effect of edge on avian nest success:
how strong is the evidence? Conserv. Biol. 8: 17–26.
Philip Whitfield, D., Ruddock, M. & Bullman, R. 2008. Expert
opinion as a tool for quantifying bird tolerance to human
disturbance. Biol. Conserv. 141: 2708–2717.
Picozzi, N. 1984. Breeding biology of polygynous Hen
Harriers Circus c. cyaneus in Orkney. Ornis Scand. 15: 1–10.
Pinheiro, J., Bates, D., DebRoy, S., Sarkar, D. & R Core
Team 2017. nlme: Linear and nonlinear mixed effects
models. Available at: https://CRAN.R-project.org/package=
rgdal.
Prestt, I. 1965. An enquiry into the recent breeding status of
some of the smaller birds of prey and crows in Britain.
Bird Study 12: 196–221.
R Core Team. 2017. R: a language and environment for statistical
computing. R Foundation for Statistical Computing, Vienna.
Available at: https://www.r-project.org/.
Redpath, S.M., Madders, M., Donnelly, E., Anderson, B.,
Thirgood, S., Martin, A. & McLeod, D. 1998. Nest
site selection by Hen Harriers in Scotland. Bird Study 45:
51–61.
Redpath, S.M., Amar, A., Smith, A., Thompson, D.B. &
Thirgood, S. 2010. People and nature in conflict: can we
reconcile hen harrier conservation and game management.
In J. Baxter & C. A. Galbraith. (ed) Species Management:
challenges and solutions for the 21st century, 335–350.
Stationery Office Books, Edinburgh.
469
Renou-Wilson, F., Bolger, T., Bullock, C., Convery, F.,
Curry, J., Ward, S., Wilson, D. & Müller, C. 2011.
BOGLAND: sustainable management of Peatlands in
Ireland – final report. Johnstown Castle, Environmental
Protection Agency, Co Wexford.
Ruddock, M. & Whitfield, D.P. 2007. A Review of Disturbance
Distances in Selected Bird Species. Report from Natural
Research (Projects) Ltd to Scottish Natural Heritage.
Natural Research, Banchory, UK.
Ruddock, M., Dunlop, B., O’Toole, L., Mee, A. & Nagle, T.
2012. Republic of Ireland National Hen Harrier Survey 2010.
National Parks and Wildlife Service, Department of Arts,
Heritage and the Gaeltacht, Ireland.
Ruddock, M., Mee, A., Lusby, J., Nagle, T., O’Neill, S. &
O’Toole, L. 2016. The 2015 National Survey of
Breeding Hen Harrier in Ireland. Irish Wildlife Manuals
No. 93. National Parks and Wildlife Service,
Department of the Arts, Heritage and the Gaeltacht,
Dublin.
Ruhlen, T.D., Abbott, S., Stenzel, L.E. & Page, G.W. 2003.
Evidence that human disturbance reduces Snowy Plover
chick survival. J. Field Ornithol. 74: 300–304.
Smart, J., Amar, A., Sim, I.M., Etheridge, B., Cameron, D.,
Christie, G. & Wilson, J.D. 2010. Illegal killing slows
population recovery of a re-introduced raptor of high
conservation concern – the red kite Milvus milvus. Biol.
Conserv. 143: 1278–1286.
Smith, J.A. & Dwyer, J.F. 2016. Avian interactions with
renewable energy infrastructure: an update. Condor 118:
411–423.
Soule, M.E. 1985. What is conservation biology? BioScience
35: 727–734.
Steven, R., Pickering, C. & Castley, J.G. 2011. A review of the
impacts of nature based recreation on birds. J. Env. Manage.
92: 2287–2294.
Suzuki, R. & Shimodaira, H. 2006. Pvclust: an R package for
assessing the uncertainty in hierarchical clustering.
Bioinformatics. 22: 1540–1542.
Suzuki, R. & Shimodaira, H. 2015. pvclust: hierarchical
clustering with P-values via multiscale bootstrap
resampling. Available at: https://CRAN.R-project.org/
package=pdist.
Taylor, A.R. & Knight, R.L. 2003. Wildlife responses to
recreation and associated visitor perceptions. Ecol. Appl.
13: 951–963.
Thaker, M., Zambre, A. & Bhosale, H. 2018. Wind farms
have cascading impacts on ecosystems across trophic
levels. Nat. Ecol. Evol. 2: 1854–1858.
Twining, J.P., Montgomery, I., Fitzpatrick, V., Marks, N.,
Scantlebury, D.M. & Tosh, D.G. 2019. Seasonal,
geographical, and habitat effects on the diet of a
recovering predator population: the European pine
marten (Martes martes) in Ireland. Euro. J. Wildlife Res.
65: 51.
Vangansbeke, P., Blondeel, H., Landuyt, D., De Frenne, P.,
Gorissen, L. & Verheyen, K. 2017. Spatially
combining wood production and recreation with
biodiversity conservation. Biodivers. Conserv. 26: 3213–3239.
Ward, J.H. 1963. Hierarchical grouping to optimize an
objective function. J. Am. Stat. Assoc. 58: 236–244.
Watson, D. 2017. The Hen Harrier. Bloomsbury Natural
History, London.
470
A. CARAVAGGI ET AL.
Webber, A.F., Heath, J.A. & Fischer, R.A. 2013. Human
disturbance and stage-specific habitat requirements
influence snowy plover site occupancy during the
breeding season. Ecol. Evol. 3: 853–863.
White, C.M. & Thurow, T.L. 1985. Reproduction of
Ferruginous Hawks exposed to controlled disturbance.
Condor 87: 14–22.
Whitfield, D.P., Ruddock, M. & Bullman, R. 2008. Expert
opinion as a tool for quantifying bird tolerance to human
disturbance. Biol. Conserv. 141: 2708–2717.
Wilcove, D.S., Rothstein, D., Dubow, J., Phillips, A. &
Losos, E. 1998. Quantifying threats to imperiled species in
the United States. BioScience 48: 607–615.
Wilson, M.W., Irwin, S., Norriss, D.W., Newton, S.F., Collins,
K., Kelly, T.C. & O’Halloran, J. 2009. The importance of prethicket conifer plantations for nesting Hen Harriers Circus
cyaneus in Ireland. Ibis 151: 332–343.
Wilson, M.W., O’Donoghue, B., O’Mahony, B., Cullen, C.,
O’Donoghue, T., Oliver, G., Ryan, B., Troake, P., Irwin,
S., Kelly, T.C., Rotella, J.J. & O’Halloran, J. 2012.
Mismatches between breeding success and habitat
preferences in Hen Harriers Circus cyaneus breeding in
forested landscapes. Ibis 154: 578–589.
Wilson, M.W., Fernández-Bellon, D., Irwin, S. &
O’Halloran, J. 2017. Hen Harrier Circus cyaneus
population trends in relation to wind farms. Bird Study
64: 20–29.
Young, J., Watt, A., Nowicki, P., Alard, D., Clitherow, J.,
Henle, K., Johnson, R., Laczko, E., McCracken, D.,
Matouch, S., Niemela, J. & Richards, C. 2005.
Towards sustainable land use: identifying and
managing the conflicts between human activities and
biodiversity conservation in Europe. Biodivers. Conserv.
14: 1641–1661.