UHI Research Database pdf download summary
Black guillemot ecology in relation to tidal stream energy generation: An evaluation of current knowledge and information gaps Johnston, Daniel T.; Furness, Robert W.; Robbins, Alexandra M. C.; Tyler, Glen; Taggart, Mark A.; Masden, Elizabeth A. Published in: Marine Environmental Research Publication date: 2018 Publisher rights: Copyright © 2018 Elsevier Ltd. All rights reserved. The re-use license for this item is: CC BY-NC-ND The Document Version you have downloaded here is: Peer reviewed version
The final published version is available direct from the publisher website at: 10.1016/j.marenvres.2018.01.007 Link to author version on UHI Research Database
Citation for published version (APA): Johnston, D. T., Furness, R. W., Robbins, A. M. C., Tyler, G., Taggart, M. A., & Masden, E. A. (2018). Black guillemot ecology in relation to tidal stream energy generation: An evaluation of current knowledge and information gaps. Marine Environmental Research, 134, 121-129. https://doi.org/10.1016/j.marenvres.2018.01.007
General rights Copyright and moral rights for the publications made accessible in the UHI Research Database are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights:
1) Users may download and print one copy of any publication from the UHI Research Database for the purpose of private study or research. 2) You may not further distribute the material or use it for any profit-making activity or commercial gain 3) You may freely distribute the URL identifying the publication in the UHI Research Database
Take down policy If you believe that this document breaches copyright please contact us at [email protected] providing details; we will remove access to the work immediately and investigate your claim.
Download date: 24. Sep. 2021 Accepted Manuscript
Black guillemot ecology in relation to tidal stream energy generation: An evaluation of current knowledge and information gaps
Daniel T. Johnston, Robert W. Furness, Alexandra M.C. Robbins, Glen Tyler, Mark A. Taggart, Elizabeth A. Masden
PII: S0141-1136(17)30385-9 DOI: 10.1016/j.marenvres.2018.01.007 Reference: MERE 4436
To appear in: Marine Environmental Research
Received Date: 21 June 2017 Revised Date: 9 January 2018 Accepted Date: 10 January 2018
Please cite this article as: Johnston, D.T., Furness, R.W., Robbins, A.M.C., Tyler, G., Taggart, M.A., Masden, E.A., Black guillemot ecology in relation to tidal stream energy generation: An evaluation of current knowledge and information gaps, Marine Environmental Research (2018), doi: 10.1016/ j.marenvres.2018.01.007.
This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. 1 ACCEPTED MANUSCRIPT
1 Black guillemot ecology in relation to tidal stream energy generation: An
2 evaluation of current knowledge and information gaps
3 Daniel T. Johnston 1*, Robert W. Furness 2, Alexandra M.C. Robbins 3, Glen Tyler 3, Mark A. Taggart 1, 4 Elizabeth A. Masden 1
5 1 Environmental Research Institute, North Highland College UHI, University of the Highlands and 6 Islands, Thurso, KW14 7EE 2 MacArthur Green Ltd, 95 South Woodside Road, Glasgow, G20 6NT, UK; 3 7 Scottish Natural Heritage, Great Glen House, Leachkin Road, Inverness, IV3 8NW
8 *Correspondence author. email: [email protected]
9 Keywords: renewable energy, conservation, marine spatial planning, environmental impact
10 assessment, seabirds
11 Abstract
12 The black guillemot Cepphus grylle has been identified as a species likely to interact with marine
13 renewable energy devices, specifically tidal turbines, with the potential to experience negative 14 impacts. This likelihood is primarily based on the speciesMANUSCRIPT being a diving seabird, and an inshore, 15 benthic forager often associating with tidal streams. These behavioural properties may bring them
16 into contact with turbine blades, or make them susceptible to alterations to tidal current speed,
17 and/or changes in benthic habitat structure. We examine the knowledge currently available to
18 assess the potential impacts of tidal stream turbines on black guillemot ecology, highlight knowledge
19 gaps and make recommendations for future research. The key ecological aspects investigated
20 include: foraging movements, diving behaviour, seasonal distribution, other sources of disturbance
21 and colony recovery. Relating to foraging behaviour, between studies there is heterogeneity in black
22 guillemot habitatACCEPTED use in relation to season, tide, diurnal cycles, and bathymetry. Currently, there is
23 also little knowledge regarding the benthic habitats associated with foraging. With respect to diving
24 behaviour, there is currently no available research regarding how black guillemots orientate and
25 manoeuvre within the water column. Black guillemots are considered to be a non-migratory species,
26 however little is known about their winter foraging range and habitat. The effect of human 2 ACCEPTED MANUSCRIPT 27 disturbance on breeding habitat and the metapopulation responses to potential mortalities are
28 unknown. It is clear further understanding of black guillemot foraging habitat and behaviour is
29 needed to provide renewable energy developers with the knowledge to sustainably locate tidal
30 turbines and mitigate their impacts.
31 Acknowledgments
32 DTJ is funded through a Marine Alliance for Science and Technology for Scotland (MASTS)
33 studentship supported by Scottish Natural Heritage (SNH). The authors thank Nicola Largey, Nina
34 O’Hanlon, and the anonymous reviewers for their comments and suggested improvements to the
35 manuscript.
36 1. Introduction
37 Seabirds are affected by anthropogenic activities including fishing (Daunt et al., 2008;
38 Furness, 2002), oil spills (Ewins, 1986; Golet et al., 2002), and climate change (Grémillet and 39 Boulinier, 2009; MacDonald et al., 2015). The incre MANUSCRIPTase in anthropogenic activity arising from the 40 growing marine renewable energy industry still has unknown effects on seabirds (Inger et al., 2009).
41 In Scotland, this industry is driven by goals set by the Scottish Government to achieve 100%
42 renewable energy by 2020 (Davies et al., 2014). Tidal stream energy devices, i.e. those based on
43 tidally driven currents channelled through narrow areas between landmasses or around headlands,
44 are currently being tested and installed in advance of potential large-scale implementation. In
45 Scotland, 18 inshore areas have been leased for tidal development (The Crown Estate, 2016). In the 46 Pentland Firth, arraysACCEPTED of up to 398 large tidal stream turbines are planned in areas with potential 47 current speeds of >2 m/s -1 (Rollings, 2011), with 3 trial turbines installed in winter 2016/17.
48 The potential impacts of tidal turbines on diving seabirds are largely unknown since only a
49 few devices have been deployed to date (Savidge et al., 2014; SNH, 2016). In Scotland, the black
50 guillemot Cepphus grylle has been highlighted as the seabird species most at risk from tidal turbines 3 ACCEPTED MANUSCRIPT 51 (Furness et al., 2012; Robbins,
52 2012), in part, due to their
53 association with tidal streams (Nol
54 and Gaskin, 1987; Waggitt et al.,
55 2016b). It has been suggested that
56 the impacts of tidal turbines on
57 diving seabirds may include
58 changes to foraging habitat,
59 foraging efficiency, and diving
60 behaviour (Benjamins et al., 2015;
61 Scott et al., 2014) as well as direct
62 mortality through collision (Furness
63 et al., 2012; Wilson et al., 2006). MANUSCRIPT64 The need for further research 65 regarding impacts to and
Figure 1. Scottish wave and tidal renewable lease areas and black66 responses of seabirds to renewable guillemot Marine Protected Areas 67 energy installations has been
68 highlighted (Inger et al., 2009; Langton et al., 2011; Scott et al., 2014), especially in relation to tidal
69 turbines (Furness et al., 2012; Masden et al., 2013) though it is acknoweldged that this poses
70 challenges (Fox et al., 2018). The sensitivity of diving seabirds to the impacts of tidal turbines is likely
71 to vary in relation to the extent of spatial and temporal overlap (Waggitt and Scott, 2014). In the 72 absence of empiricalACCEPTED data, sensitivity can be expected to relate to foraging locations, diving depth, 73 variation in year-round distribution and behaviour, association with tidal currents, visual nature of
74 foraging, and diurnal rhythms in foraging (Furness et al., 2012; Wilson et al., 2006). Further
75 ecological knowledge of these factors is needed to inform decision makers on the potential impacts
76 related to tidal turbine allocation and operation. 4 ACCEPTED MANUSCRIPT 77 The siting of marine renewable lease sites around Scotland coincides with the designation of
78 six Scottish Marine Protected Areas (MPAs) specifically with black guillemots as a major protected
79 feature (SNH, 2014; Figure 1). A further four MPAs have been established for benthic communities
80 containing kelp which may in turn protect habitat associated with black guillemot prey (blennies and
81 gadoids) (Ewins, 1990; SNH, 2014). This also emphasises the need for more information on black
82 guillemot foraging ecology to ensure MPAs fulfil their desired conservation outcomes by
83 encompassing foraging habitat.
84 The aim of this paper is to provide a synthesis of current knowledge regarding black
85 guillemot ecology and its relevance to potential impacts from tidal stream turbines. Key aspects of
86 black guillemot movement are reviewed, including: (1) foraging movements, (2) diving behaviour,
87 and (3) seasonal distribution. Further factors, which may compound the impact of tidal stream
88 turbines are also considered, including other sources of disturbance to nesting adults and the
89 process of colony recovery. For each topic: (1) the knowledge required to assess the impacts of tidal 90 stream turbines on black guillemots is outlined, (2MANUSCRIPT) current knowledge is reviewed, (3) gaps in 91 current research are highlighted, and recommendations for future research are provided.
92 2. Foraging movements
93 Tidal stream turbines may increase habitat heterogeneity through the addition of previously
94 absent structures, which will potentially form reefs, act as Fish Aggregating Devices (FADs), and may
95 alter seabed sedimentation patterns (Miller et al., 2013; Shields et al., 2009). How susceptible a
96 seabird will be to these changes depends on their ecology (Wilson et al., 2006). Firstly, species which
97 forage within theACCEPTED inshore environment rather than offshore will most likely encounter tidal devices
98 (installed within 5km of shore) (The Crown Estate, 2016). Secondly, temporally varying habitat use
99 (related to season, time of day, or tidal phase) may influence the duration foraging birds experience
100 disturbance from turbine construction, maintenance, or operation (Waggitt and Scott, 2014). Lastly,
101 the dive depths exhibited by a foraging bird may indicate the extent of vertical overlap with turbine 5 ACCEPTED MANUSCRIPT 102 blades (Furness et al., 2012), dependent on the blade depth profile of the class of tidal device
103 encountered, which include floating (3-19m) (Scotrenewables, 2017) and seafloor mounted (8-26m)
104 (Masden et al., 2013; MeyGen, 2012). In terms of black guillemots, a small foraging range, frequent
105 use of inshore areas, and deep/benthic diving behaviour could bring individuals into frequent
106 contact with inshore tidal turbines (Furness et al. 2012). The assessment of the overlap between
107 tidal turbines and foraging black guillemots requires ecological knowledge of both movement and
108 habitat use at sea.
109 2.1 Foraging movements: Current knowledge
110 Black guillemots are inshore, generalist, central place foragers with a small foraging radius.
111 Initial work regarding black guillemot foraging ranges monitored the direction that adults flew from
112 the nest by visually following individual birds (Cairns, 1992; Ewins, 1986; Nol and Gaskin, 1987;
113 Petersen, 1981) (Tab. 1). More recently, electronic tracking studies of adult black guillemots have 114 revealed more accurately the foraging locations and MANUSCRIPTdistances travelled by individuals (Owen, 2015; 115 Sawyer, 1999) (Tab. 1).
Table 1. Summary of breeding season foraging distance studies for black guillemots. Reference Method n Seasons Mean Median (km) Maximum (km) Location sampled (km) Petersen, 1981 Visual observation 65 3 2-4 - 7 Flatey Island, Iceland Ewins, 1986 Visual observation 73 1 - - 2 Shetland, Scotland Sawyer, 1999 Visual observation 623 1 - 5.5 8.4 Holm of Papa, Orkney, (300m offshore) (840m offshore) Scotland Sawyer, 1999 VHF tag 18 3 - 5.5 9 Holm of Papa, Orkney, (275m offshore) (700m offshore) Scotland Shoji et al., 2015 GPS tag 1 1 - - 1.8 Lighthouse Island, Copeland, Northern Ireland
Owen,2015 GPS tagACCEPTED 19 1 - - 7.45 Grass Holm, Shapinsay, Orkney, Scotland 116
117 These relatively small (Thaxter et al., 2012), inshore, foraging ranges are reflected in their
118 diet, and lead to variability in prey species between colonies due to variations in the characteristics 6 ACCEPTED MANUSCRIPT 119 of the local habitat (Barrett and Anker-Nilssen, 1997; Ewins, 1990; Hario, 2001). In Scotland,
120 butterfish Pholis gunnellus tend to dominate diet, followed by sandeels Ammodytes marinus ,
121 gadoids, blenny spp., sculpin spp., and flatfish spp. (Ewins, 1992, 1990; Harris and Riddiford, 1989;
122 Sawyer, 1999; Slater and Slater, 1972).
123 Some seabird species associate with tidal eddies and currents to forage as these may
124 produce upwellings and aggregate prey (Alldredge and Hamner, 1980; Embling et al., 2012; Ladd et
125 al., 2005; St. John et al., 1992). Foraging black guillemots have been shown to associate with
126 moderate current speeds and tidal eddy features of tidal streams (Wade, 2015; Waggitt et al.,
127 2016b). Foraging is most common at speeds of 0.5-1 m/s -1, reducing in frequency at higher current
128 speeds of >2 m/s -1 (Nol and Gaskin, 1987; Wade, 2015; Waggitt et al., 2016a). For black guillemots,
129 the use of tidal streams is a potential trade-off between foraging success and the energy required to
130 move in this energetically demanding environment (Nol and Gaskin, 1987). While swimming costs
131 may be higher in fast flowing water, ‘tidal-conveyor’ foraging behaviour (repeatedly flying upstream 132 and then diving, allowing the current to carry an i ndividualMANUSCRIPT downstream while foraging) may be a 133 strategy which reduces the energy required to search a large transect of the seafloor (Robbins,
134 2017). However, the relationship between black guillemots and tidal streams are not always
135 consistent between studies. For example, when comparing studies of tidal microhabitat use, Waggitt
136 et al. (2017) found heterogeneity between study sites, with the probability of detecting black
137 guillemots in fast habitats greater in 3 of 5 study sites. A number of variables could cause this
138 heterogeneity, including, differing prey communities and the associated foraging behaviour needed
139 to exploit them.
140 Foraging ACCEPTEDin tidal streams can relate to tidal phase (Robbins, 2012) potentially associated to
141 variations in current speeds or depths. Diving depths required to access the seafloor are shallower at
142 low tide, potentially reducing the energetic costs of foraging on the benthos and increasing the
143 accessibility of prey (Vermeer et al., 1993). Shore based observations of foraging black guillemots
144 found distance from shore (Ronconi and Clair, 2002), and density of individuals (Waggitt et al., 7 ACCEPTED MANUSCRIPT 145 2016a) increased at low tide. The influence of low tide on Cepphus spp. colony attendance varies in
146 significance between studies (Ewins, 1985; Petersen, 1981) potentially attributed to local tidal
147 ranges (Vermeer et al., 1993).
148 Temporal cycles may influence black guillemot behaviour. Diel cycles have been shown to
149 affect colony attendance (Hilden, 1994), foraging activity (Robbins, 2012), and foraging dives (Shoji
150 et al., 2015). This is potentially related to light levels and visibility in the water column (Regular et al.,
151 2011) or diel movements of prey (Kamenos et al., 2004). Changes in prey species provisioned to
152 chicks has been correlated with time of day, shifting from a peak sandeels in the morning (0700 hrs)
153 to blenny spp. thereafter (Ewins, 1990). The tidal speeds with which black guillemots associate with
154 while foraging may alter seasonally, however this relationship is variable between studies. Foraging
155 took place in faster flows and a broader range of habitats in the non-breeding season compared to
156 the breeding season in the Fall of Warness, Orkney (Waggitt et al., 2016b), but while the reverse was
157 observed in the Pentland Firth (Wade, 2015). It was speculated that a release from interspecific 158 competition (Waggitt et al., 2016b) or a reduction inMANUSCRIPT prey availability in winter (Ewins, 1990) may 159 cause black guillemots to broaden their foraging niche into high flow areas.
160 Differing habitat use observed among studies may be related to the exploitation of niches
161 which vary locally in substrate type, seaweed species, and tidal dynamics. However, studies
162 highlighting the prevalence of individual behaviour in black guillemots (Owen, 2015; Petersen, 1981;
163 Slater and Slater, 1972), raise the possiblity of inter-individual niches existing within the mean niche
164 (Bolnick et al., 2003). Niches related to tidal stream dynamics could be altered by tidal turbines. The
165 prevelence of inter-individual niches suggests that individuals within a colony could associate with
166 tidal streams to ACCEPTED different extents, and so are affected by habitat change related to tidal turbines
167 differently. Generalist and specialist foraging behaviours prevelent in black guillemots (Cairns 1987,
168 Ewins 1990, Barrett and Anker-Nilssen 1997) and pigeon guillemots (Litzow et al., 2004, 2000, 1998),
169 may dictate those individuals best suited to adapt to habitat change. 8 ACCEPTED MANUSCRIPT 170 2.2 Foraging movements: Knowledge gaps
171 In assessing the extent to which black guillemots use tidal currents, previous studies
172 highlight that while tidal streams are a common foraging habitat, a large degree of heterogeneity
173 exists in their use (Nol and Gaskin, 1987; Wade, 2015; Waggitt et al., 2017, 2016b). The reasons
174 behind these differing tidal associations between study sites has not been investigated in-depth.
175 Previous studies have considered differences to be potentially related to the pressures of
176 interspecific competition or differing benthic communities (Wade, 2015; Waggitt et al., 2017,
177 2016b). Few studies have investigated the relationship between tidal currents, bathymetry and
178 benthos. There is currently a knowledge gap in terms of the interactions between these three
179 properties, and further investigation could highlight a common driver (or drivers) behind black
180 guillemot foraging distributions within tidal streams. By identifying these habitat associations,
181 predictions can also be made as to how they may be altered by tidal turbines. To identify the
182 sensitivity and responses of birds to tidal stream turbines, future research should focus on the 183 ecological relationships that intertwine with tidal streams,MANUSCRIPT explicitly the benthos and associated 184 prey, and how certain tidal dynamics may be advantageous to foraging. GPS data, such as those
185 presented in Owen (2015) identify horizontal movement patterns and potential foraging locations,
186 but, the environmental characteristics of these foraging locations have not yet been detailed. It is
187 possible that tidal stream turbines may significantly alter the conditions that foraging birds respond
188 to, but until the drivers behind foraging are established, it is not possible to fully consider the risks
189 and potential impacts.
190 What areACCEPTED the characteristics of black guillemot foraging habitats, what changes will tidal 191 turbines exert on these habitats, and how will black guillemots use these habitats, are all primary
192 questions developers need to understand when siting tidal turbines. Potential outcomes of tidal
193 turbine installation include shifts in current flows previously associated with foraging, or blocked
194 access to benthic habitat (Savidge et al., 2014). Tidal devices may increase turbulence within the 9 ACCEPTED MANUSCRIPT 195 current flow (Churchfield et al., 2013; Edmunds et al., 2014), having ecological impacts downstream
196 (Shields et al., 2011). Alternatively, tidal turbines may act as FADs (Scott et al., 2014; Wilhelmsson et
197 al., 2006), potentially attracting more foraging birds (Savidge et al., 2014). The formation of reefs
198 may create new habitat features producing heterogeneous biodiversity in comparison to the
199 adjacent seabed (Broadhurst and Orme, 2014; Shields et al., 2009; Wilhelmsson et al., 2006). Though
200 FADs may increase fish abundance, fish may aggregate around turbines to avoid predation,
201 potentially reducing prey availability to seabirds and/or encouraging them to forage in areas of
202 higher collision risk (Langton et al., 2011). Where high current flows create physical stress on benthic
203 communities, a reduction in kinetic energy through tidal turbine structures may precipitate an
204 increase in biodiversity and species abundance (Moore and Roberts, 2011).
205 The advantages of individual-specific variation in foraging location (Owen, 2015), or diet
206 (Slater and Slater, 1972) are not well understood. Knowledge of the extent of individual variation in
207 foraging behaviour driven by individual preference, life-stage or sex, is required in order to define a 208 sample size of foraging site or diet observations that MANUSCRIPT is representative of the colony (Scott et al., 209 2014; Soanes et al., 2013). Individually varying threats to habitat use or collision risk posed by tidal
210 turbines can indicate the proportion of a colony at risk.
211 With the potential for habitat use and diet to vary diurnally (Ewins, 1990), tidal turbine
212 operation, construction, and maintenance may obstruct foraging birds at key periods, i.e., when
213 foraging is most efficient and high quality prey items are normally present in the diet. Distinct
214 colonies have differing behavioural responses to tidal phase and diurnal cycles. While there have 215 been some suggestionsACCEPTED for the reasons for these differences, including tidal height and diet, no 216 research has positively identified the underlying drivers (Ewins, 1990; Petersen, 1981; Vermeer et
217 al., 1993). Further study of the ecological interactions driving temporal variation in movement may
218 help inform and mitigate against the periods during which disturbance from construction and
219 maintenance of tidal turbines is most deleterious to foraging birds. 10 ACCEPTED MANUSCRIPT 220 Modern tagging techniques, such as GPS (Owen, 2015), and time-depth recorders (TDRs)
221 (Masden et al., 2013; Shoji et al., 2015) are powerful methods of identifying the horizontal and
222 vertical foraging movements of individual movement. Associating foraging location and behaviour to
223 habitat or prey items would provide useful information on the environmental characteristics related
224 to foraging strategies (Elliott et al., 2008; Woo et al., 2008). While several studies have suggested
225 that foraging in black guillemots is associated with kelp, few have made a definitive link (Ewins,
226 1992). Combining tracking techniques, prey observations, and environmental data could identify
227 these links. Previously, studies have related seabird diet to movement using data loggers in relation
228 to variables of depth (Barrett and Furness, 1990; Woo et al., 2008), foraging trip duration
229 (Weimerskirch, 1998), and foraging location (Caron-Beaudoin et al., 2013; Linnebjerg et al., 2013;
230 Votier et al., 2010). Understanding black guillemot horizontal foraging location and associated prey
231 would provide developers and licensing authorities with invaluable knowledge regarding habitat use
232 and ecology within tidal lease areas and MPAs.
233 3. Diving behaviour MANUSCRIPT
234 Turbine arrays have been suggested to act as barriers to diving seabirds (Savidge et al., 2014)
235 and have been identified as particularly hazardous to black guillemots due to their benthic foraging
236 behaviour (Furness et al., 2012, Masden et al., 2013; Robbins, 2012; Wade, 2015). The behavioural
237 responses of diving birds to tidal turbine blades have yet to be fully understood, including the
238 potential risks of collision or affects to foraging efficiency (Wilson et al., 2007). It is currently
239 unknown how black guillemots use tidal currents when diving, currents which tidal turbines may 240 alter by obstructingACCEPTED water flow (Shields et al. 2009). In foraging areas associated with tidal stream 241 turbines, knowledge of black guillemot diving behaviour in relation to direction, profile, and use of
242 currents could indicate the potential for underwater interactions with turbine blades.
243 3.1 Diving behaviour: Current knowledge 11 ACCEPTED MANUSCRIPT 244 A wide range of diving depths have been reported for black guillemots, although they have
245 not been recorded to exceed 50m (Ewins, 1986; Masden et al., 2013; Nol and Gaskin, 1987; Piatt and
246 Nettleship, 1985; Shoji et al., 2015) (Tab.2).
Table 2. Summary of breeding season dive depth studies for black guillemots. Reference Method n Seasons Mean SD Median Maximum Location sampled (m) (m) (m) Ewins, 1986 Visual observation 73 1 30-50 - - 50 Shetland, Scotland Piatt and Nettleship, 1985 Fisheries by-catch 36 2 13.6 - - 50 Newfoundland, Canada Nol and Gaskin, 1987 Visual observation 799 2 26.4 14.9 - 50 Bay of Fundy, Canada Cairns, 1992 Visual observation 130 1 21.1 10.5 22.5 48 Northeast Hudson Bay, Canada Masden et al., 2013 TDR tag 2 1 - - 32 43 Stroma, Caithness, Scotland Shoji et al., 2015 TDR tag 4 1 9.3 2.8 - 18.7 Bangor Harbour and Lighthouse Island, Copeland, Northern Ireland 247
248 When foraging in the water column black guillemots have been shown to undertake W-
249 shaped, V-shaped and U-shaped dives (Shoji et al., 2015). V-shaped and W-shaped dives exploit the
250 midwater, while U-shaped dives are indicative of foraging lower in the water column, suggesting 251 that black guillemot diving behaviour is often related MANUSCRIPTto the seafloor (Shoji et al., 2015). 252 Tidal-conveyor foraging entails an individual drifting downstream with the force of a tidal
253 current before flying upstream, landing and then diving (Robbins, 2017). Fraenkel (2006) has
254 suggested that in relation to tidal turbines, diving birds will likely be swept between blades within
255 the current flow (through ‘entrainment’), or, possess the manoeuvrability to avoid contact with
256 blades. Langton et al. (2011) countered this assumption, indicating that for diving birds to
257 manoeuvre around turbines they must first be able to see, recognise and predict blade movements.
258 Wade (2015) suggests that seabirds diving within tidal streams actively face into the direction of the
259 current; this was ACCEPTEDconcluded from surface observations of locations where individuals submerged and
260 resurfaced, and assumptions that fusiform body shapes are more suited to swimming against the
261 current, minimizing drag and energy expenditure (Lovvorn et al., 2001). The fact that diving birds
262 may actively swim into the direction of the current may increase the likelihood of collision, as
263 turbines approached from upstream may be outside their field of vision (Wade, 2015). 12 ACCEPTED MANUSCRIPT 264 3.2 Diving behaviour: Knowledge gaps
265 Current literature describes black guillemots diving to depths that overlap with turbine
266 blades (Masden et al., 2013). However, whether these dives will overlap with the horizontal
267 locations of turbines in sites with high tidal flow is as yet unknown. Furthermore, there is currently
268 no available research to indicate how black guillemot’s orientate and manoeuvre within the water
269 column and may behave when encountering a tidal turbine.
270 To address this, knowledge of black guillemots diving behaviour in relation to current flow
271 speeds, and manoeuvrability in relation to obstacles or arrays of turbines is a priority. To prevent
272 collisions, blade avoidance would need to occur while birds are swimming through strong tidal
273 currents whilst also ascending or descending through the water column actively seeking prey.
274 Sediment disturbed by tidal turbines may increase turbidity and affect visibility within the water
275 column (Shields et al. 2009). Decreased visibility may significantly affect the foraging ability of diving 276 birds, whilst also reducing their ability to avoid collisionMANUSCRIPT with devices (Grecian et al., 2010; Wilson et 277 al., 2006). Black guillemots, as with other auks, a re visual foragers (Martin and Wanless, 2015),
278 though little is known of the extent to which turbidity will effect foraging behaviour. Tactile cues
279 may be employed by diving birds for navigation in low light situations, but these have yet to be
280 identified (Regular et al., 2011), making vision the primary known sense to be potentially affected.
281 Turbidity has been shown to reduce visual resolution in great cormorants Phalacrocorax carbo (Strod
282 et al., 2004) . Similar work is needed on the extent to which visibility relates to black guillemot
283 foraging efficiency, and, how their optical orientation (Martin and Wanless, 2015) and depth of 284 vision (Katzir andACCEPTED Howland, 2003) relates to their foraging movements and their potential to react to 285 structures in their underwater environment (Martin, 2011).
286 Further research combining diving depth data with horizontal location information may
287 reveal how black guillemots use the water column in three-dimensions and potentially quantify the
288 extent of overlap with tidal turbine blades. One major information gap to be addressed is the 13 ACCEPTED MANUSCRIPT 289 relationship of dive depth relationship with conditions such as bathymetry and the benthos.
290 Differences in dive depths between Masden et al. (2013) and Shoji et al. (2015) were primarily
291 attributed to variation in bathymetry between the differing study sites. Approaches which may help
292 improve our knowledge of underwater seabird behaviour might include combined deployments of
293 TDRs, animal-borne cameras, accelerometers and GPS devices (Berlincourt and Arnould, 2014; Sato
294 et al., 2008). By combining accelerometers with TDRs, foraging events at certain depths can be
295 inferred, and have been used to identify foraging habits in diving seabirds (Laich et al., 2010;
296 Watanuki et al., 2006). Animal-borne cameras can record the field of vision of the animal, identifying
297 foraging events and behaviour within track data (i.e., prey captures and species). In other studies
298 (which could be mimicked for black guillemots), the relationship between dive depth and sediment
299 types were investigated in European shags Phalacrocorax aristotelis through the use of combined
300 camera and depth sensor deployments (Watanuki et al., 2008). However, the technology is not yet
301 small enough to deploy on smaller diving birds, including black guillemots (Chimienti et al., 2016).
302 4. Seasonal distribution MANUSCRIPT
303 Assessing seasonal variations in the overlap of black guillemot foraging distributions with
304 tidal stream turbines arrays is the first step required in quantifying periods of risk. Seabird
305 distributions may shift due to seasonal migrations, however, as a non-migratory seabird, black
306 guillemots remain within the inshore year round (Ewins and Tasker, 1985). Despite this, black
307 guillemot habitat use or distributions can vary in response to climatic conditions, prey behaviour, or
308 inter-annual variations in relation to climate change.
309 4.1 Seasonal distribution:ACCEPTED Current knowledge
310 As year-round inshore residents (Ewins and Kirk, 1988) black guillemots must adapt to
311 changing inshore conditions. For example, in Shetland, black guillemots are known to move to less
312 exposed areas of coastline in the winter (Ewins and Kirk, 1988). The winter range of black guillemots 14 ACCEPTED MANUSCRIPT 313 has also shifted over decadal scales (Potvin et al., 2016; Veit and Manne, 2015) attributed to climate
314 change.
315 In the non-breeding season, birds are no longer central-place foragers as they do not need
316 to return to a single nest location with prey, which will potentially alter their foraging locations
317 (Mehlum and Gabrielsen, 1993). As well as modulating their foraging priorities, foraging habitat may
318 broaden with the opening up of prey resources previously unavailable during the breeding season
319 due to intra-specific competition with species that have migrated (Chase, 2011; Waggitt et al.,
320 2016b).
321 Foraging locations may change to correspond with changes in prey behaviour (Ewins and
322 Kirk, 1988). Some benthic prey species migrate to deeper water in winter, such as butterfish (Shorty
323 and Gannon, 2013), whilst others (i.e., sandeels) remain buried for most of the winter except to
324 spawn. Ewins (1990) explored the differences in breeding (May-August) and non-breeding 325 (September-April) diet in adult birds. Taking into accountMANUSCRIPT different sample sizes between seasons, 326 the variety of prey items observed in non-breeding diet was larger than in the breeding season.
327 4.2 Seasonal distribution: Knowledge gaps
328 Seasonal differences in movement may dictate the periods which black guillemots are most
329 likely to overlap with tidal arrays. To investigate these movements, counts through visual
330 observations (Ewins and Kirk, 1988) and aerial surveys (Gaston and Mclaren, 1990; Heinänen et al.,
331 2017; Prach and Smith, 1992) may be carried out year-round, and are useful methods to investigate 332 population scale ACCEPTEDmovements. Observational survey methods can highlight fine scale foraging habitat 333 in both breeding and non-breeding seasons (Waggitt and Scott, 2014), however, individual
334 movements studied through the use of tracking deployments are difficult to study outside of the
335 breeding season. During the non-breeding season, birds do not display site fidelity to a nest, making
336 their capture difficult. The use of geolocators (GLS) has made tracking over long periods feasible, 15 ACCEPTED MANUSCRIPT 337 allowing capture during the breeding season and data collection carrying on into the winter, this has
338 provided data on non-breeding movements of some alcids (Harris et al., 2015, 2013; Jessopp et al.,
339 2013; McFarlane Tranquilla et al., 2014). The use of GLS technology has contributed significantly to
340 improving understanding of the non-breeding movements of arctic black guillemots (Divoky et al.,
341 2016). GLS technology assesses individual movement within an error of several hundred kilometres,
342 making it a viable method to study movements relating to migration (Jessopp et al., 2013), and large
343 scale sea-ice boundary movement (Divoky et al., 2016).
344 Diet could be used as a tool to investigate black guillemot foraging ecology, which may be a
345 useful indicator of potential seasonal shifts in foraging habitat related to tidal turbines. Current
346 literature provides limited knowledge of the inshore behaviour and diet of black guillemots during
347 the non-breeding season (Ewins and Kirk, 1988; Gaston and Mclaren, 1990; Wade, 2015; Waggitt et
348 al., 2016b). Studies of black guillemot diet have primarily been conducted during the breeding
349 season and therefore largely reflect chick diet (Barrett and Furness, 1990; Cairns, 1987a; Ewins, 350 1990; Furness and Barrett, 1985; Harris and Riddiford, MANUSCRIPT 1989; Petersen, 1981; Weslawski et al., 1994). 351 These studies are also limited to a relatively short period of the year and to restrictive foraging
352 ranges (Brown and Ewins, 1996). This makes adult diet outside the breeding season a more robust,
353 but difficult to ascertain, measure of black guillemot foraging ecology in relation to habitats
354 associated with tidal turbines (Barrett et al., 2016).
355 5. Sources of disturbance and colony recovery
356 Tidal turbine arrays may affect the nesting habitat of black guillemots through the
357 construction of infrastructureACCEPTED onshore to manage the electricity generated. Human disturbance
358 through ongoing turbine maintenance and operation also has the potential to displace black
359 guillemots, affect breeding success, and disturb nesting habitat (Cairns, 1980). When assessing the
360 extent of the impact created by tidal turbines on breeding black guillemots, it is imperative that 16 ACCEPTED MANUSCRIPT 361 existing sources of background disturbance and mortality at nest sites are taken into account, such
362 as that caused by invasive predators (Craik, 1995) and adverse weather (Hario, 2001).
363 Should colonies be disturbed by human activities related to tidal turbines (or other factors),
364 mechanisms of recovery need to be better understood. Dispersal patterns may indicate the potential
365 for metapopulation rescue of colonies that have experienced significant mortalities due to tidal
366 turbines. These dispersal patterns may relate to the amount of philopatry and emigration occurring
367 within colonies, which vary depending on several factors including competition, disturbance and
368 availability of suitable breeding habitat (Harris et al., 1996; Lavers et al., 2007; Moum and Árnason,
369 2001) . Understanding the extent of emigration and philopatry, including breeding and natal
370 dispersal distances within a metapopulation, may indicate the distance at which source-sink
371 relationships between colonies may develop between those potentially affected by tidal turbine
372 related disturbance or mortalities.
373 5.1 Sources of disturbance and colony recovery: Current knowledge MANUSCRIPT 374 Direct human disturbance at black guillemot nest sites has been suggested to cause
375 individuals to reduce incubation times or abandon breeding attempts (Cairns, 1980). However,
376 colonies have been shown to exist where human activity is high and predictable, such as harbours
377 and on ferry terminals (Greenwood, 2002). Black guillemots are regarded as crevice nesters,
378 primarily choosing gaps within boulder beaches to nest (Greenwood, 2002). They are also known to
379 nest under, or within, a range of natural and manmade objects (Greenwood, 2002). Regarded as less
380 colonial than other alcids (Harris and Birkhead 1985), black guillemot colonies are small and
381 dispersed (Cairns,ACCEPTED 1980).
382 Nesting behaviour in black guillemots is vulnerable to certain mammalian invasive species
383 (Ewins, 1990; Magnusdottir et al., 2014; Nordström, 2002). There are now numerous accounts of
384 island seabird colonies, including black guillemots, being impacted by American mink Neovison vison 17 ACCEPTED MANUSCRIPT 385 predation (Barrett and Anker-Nilssen, 1997; Clode and Macdonald, 2002; Craik, 1997, 1995;
386 Magnusdottir et al., 2014). The primary impact of mink presence is mortality, leading to a loss of
387 breeding habitat, possibly due to an effort by black guillemots to avoid predation, this is a major
388 confining aspect of black guillemot abundance, and may lead to emigration (Ewins and Tasker, 1985;
389 Mitchell et al., 2004; Nordström and Korpimäki, 2004; Petersen, 1981). Adverse weather through
390 intense precipitation or storm swells may wash out coastal nests; and the frequency of these
391 incidents may increase with climate change (Lowe et al., 2001). Nest failure caused by heavy rain
392 and high waves has previously been seen for black guillemots in Finland (Hario, 2001).
393 The ability of birds to disperse is important in colony/population recovery and persistence,
394 and the transfer of individuals between colonies may reduce the potential for colony extinctions
395 (Frederiksen and Petersen 2000). Recolonization by immature black guillemots may (for example)
396 have driven local population recovery in Sullom Voe following the Esso Bernicia oil spill in 1979
397 (Ewins, 1986; Heubeck, 1995). Dispersal distances derived from UK ring recoveries vary in relation to 398 the geographic characteristics of natal sites; with median MANUSCRIPT dispersal distances for the isolated islands 399 of Fair Isle and Foula of 56km (n = 36) while at the island groups of Orkney and Shetland distances
400 were only 6km (n = 35) (Ewins, 1988). Rates of philopatry in alcids is generally regarded as high
401 (Coulson, 2016). On Flatey Island (Iceland), 34% (n = 322) of marked black guillemots were
402 philopatric (Frederiksen and Petersen, 1999), with higher site fidelity exhibited by breeding birds,
403 and dispersal behaviour being more prevalent in natal birds (Frederiksen and Petersen, 2000).
404 However, dispersal distances did not differ between adult and immature birds in Shetland (Ewins,
405 1988). ACCEPTED 406 5.2 Sources of disturbance and colony recovery: Knowledge gaps
407 Understanding how human disturbance affects black guillemot nesting habitat could help
408 developers identify impacts associated with the construction, operation, and maintenance of tidal
409 turbines. To delineate human caused mortality and disturbance from that occurring naturally, 18 ACCEPTED MANUSCRIPT 410 background sources need to be identified. Should mortality or disturbance occur, an understanding
411 of the mechanisms of colony recovery could help assess the long-term consequences of these
412 impacts. Current literature highlights the potential pressures facing nesting black guillemots
413 (primarily terrestrial predators), however, little is known about the effects of human disturbance
414 (Ronconi and Clair, 2002). This lack of research makes identifying the response of black guillemots to
415 anthropogenic activity difficult. When assessing disturbance generated by tidal turbines, observing
416 local population changes in breeding success could also indicate the potential effects of tidal
417 turbines. At each colony, threats from terrestrial predators, human disturbance, and climate change
418 (EcoWatt 2050, 2017), may all produce larger impacts than those from tidal turbines.
419 Little is currently known about the transfer of individuals between geographically disparate
420 black guillemot colonies. How the metapoplation interacts and will respond to colony extinction is
421 relatively unknown. Current literature suggests some metapopulation connectivity with individuals
422 transferring between island groups (Ewins, 1988; Frederiksen and Petersen, 2000) and dispersal 423 from isolated islands in the absence of suitable winter MANUSCRIPT shelter (Ewins, 1988). Methods such as mark- 424 recapture of individuals and genetic analysis may be used to address knowledge gaps regarding
425 dispersal patterns within metapopulations (Barlow et al., 2013; Greenwood, 1980). While mark-
426 recapture and colour ringing are useful, they may be limited by the cryptic nature of crevice nesting
427 black guillemots, making it more difficult to study life-history aspects such as dispersal and
428 philopatry (Coulson, 2016). Limitations to mark-recapture methods could be alleviated by DNA
429 marker analysis, a method that has been suggested to be underutilized in the assessment of
430 philopatry (Coulson, 2016). DNA markers can also provide the potential for historic samples to be
431 used to study metapopulationACCEPTED composition (Wink, 2006).
432 6. Summary
433 Black guillemot foraging behaviour and year round presence within the inshore marine
434 environment have highlighted the species to be particularly vulnerable to interacting with tidal 19 ACCEPTED MANUSCRIPT 435 stream turbines (Furness et al., 2012). To assess the potential impacts of tidal stream turbines on
436 black guillemots, several gaps in our current ecological knowledge require addressing. Required
437 information includes: 1) the characteristics of the foraging habitat and diet preferences; 2)
438 underwater foraging behaviour, including dive profile and body orientation; 3) temporal shifts in
439 behaviour and foraging location related to seasonal, diel, and tidal cycles; 4) sources of additional
440 pressure on black guillemots and the potential of colony recovery from deleterious effects.
441 While some knowledge exists about the amount of time black guillemots dive at the depth
442 of turbine blades (Masden et al., 2013), little is known about how dives horizontally and vertically
443 overlap with turbines in space. Fine-scale knowledge of behaviour at specific foraging locations is
444 essential to investigate how black guillemots may interact with turbines (Waggitt et al., 2016b).
445 Methods to investigate these behaviours include visual observation (shore based or aerial), tracking,
446 and diet studies (Grémillet and Boulinier, 2009; Lascelles et al., 2016; Ronconi et al., 2012; Soanes et
447 al., 2013). Once foraging locations and behaviours are identified, collating knowledge of prey 448 distribution, benthic flora compositions, bathymetry, MANUSCRIPTtidal dynamics, and substrate, will provide vital 449 information to identify the characteristics typical of black guillemot foraging habitat.
450 Current knowledge shows that black guillemot foraging habitat, including use of tidal
451 streams, are heterogeneous between study sites (Waggitt et al., 2017), seasons (Wade, 2015;
452 Waggitt et al., 2016b) and individuals (Owen, 2015). Drivers of this heterogeneity have not been
453 robustly identified, though have so far been attributed to abiotic and biotic spatial and temporal
454 covariates (Waggitt et al., 2017). Identifying predictable or consistent drivers of habitat use across 455 studies may provideACCEPTED information to developers on which environmental characteristics will have the 456 largest impact on foraging.
457 The response of breeding birds to terrestrial predators (Craik, 1997) or humans (Ronconi and
458 Clair, 2002) could include disturbance, desertion, or nest failure. These disturbances could persist
459 leading to local population declines, or alternatively, once such pressures are removed, 20 ACCEPTED MANUSCRIPT 460 recolonization from the metapopulation could occur (Frederiksen and Petersen, 2000). Therefore,
461 further understanding of philopatry and dispersal distances by immatures from nearby colonies may
462 inform the potential for recolonization, or creation of a source-sink scenario, should any deleterious
463 impacts from tidal turbines occur.
464 Progress toward large scale tidal stream turbine installation has highlighted multiple gaps in
465 our understanding of black guillemot and (more broadly) inshore diving seabird ecology. Researching
466 these gaps will shed new light on this understudied species, which could potentially help developers
467 mitigate against deleterious effects of tidal turbine construction and operation in the future. A
468 better understanding of black guillemot ecology will put MPAs in a better position to serve their
469 purpose and, the knowledge gained will have relevance to a wider set of biota that may be impacted
470 by marine renewable developments in the future.
471 Compliance with Ethical Standards 472 The correspondence author is funded through a stude MANUSCRIPTntship awarded by Marine Alliance for Science 473 and Technology for Scotland (MASTS) supported by Scottish Natural Heritage (SNH). The authors
474 declare that they have no conflict of interest with these funding bodies. For this study, a literature
475 review, formal consent for animal research was not required, and this review does not contain any
476 previously unpublished studies with animals performed by any of the authors.
477 References
478 Alldredge, A.L., Hamner, W.M., 1980. Recurring aggregation of zooplankton by a tidal current. 479 Estuar. Coast.ACCEPTED Mar. Sci. 10, 31–37. doi:10.1016/S0302-3524(80)80047-8
480 Barlow, E.J., Daunt, F., Wanless, S., Reid, J.M., 2013. Estimating dispersal distributions at multiple
481 scales: Within-colony and among-colony dispersal rates, distances and directions in European
482 Shags Phalacrocorax aristotelis. Ibis (Lond. 1859). 155, 762–778. doi:10.1111/ibi.12060 21 ACCEPTED MANUSCRIPT 483 Barrett, R.T., Anker-Nilssen, T., 1997. Egg-laying, chick growth and food of Black Guillemots Cepphus
484 grylle in North Norway. Fauna norv. Ser. C., Cinclus 20, 69–79.
485 Barrett, R.T., Christensen-Dalsgaard, S., Anker-Nilssen, T., Langset, M., Fangel, K., 2016. Diet of adult
486 and immature North Norwegian Black Guillemots Cepphus grylle. Seabird 29, 1–14.
487 Barrett, R.T., Furness, R.W., 1990. The prey and diving depth of seabirds on Hornoy, North Norway
488 after a decrease in the Barents Sea capelin stocks. Ornis Scand. 21, 179–186.
489 doi:10.2307/3676777
490 Benjamins, S., Dale, A., Hastie, G., Waggitt, J.J., Lea, M.-A., Scott, B.E., Wilson, B., 2015. Confusion
491 reigns? A review of marine megafauna interactions with tidal-stream environments. Oceanogr.
492 Mar. Biol. An Annu. Rev. 53, 1–54. doi:10.1201/b18733-2
493 Berlincourt, M., Arnould, J.P.Y., 2014. At-sea associations in foraging little penguins. PLoS One 9,
494 e105065. doi:10.1371/journal.pone.0105065 MANUSCRIPT 495 Bolnick, D.I., Svanbäck, R., Fordyce, J. a, Yang, L.H., Davis, J.M., Hulsey, C.D., Forister, M.L., 2003. The
496 ecology of individuals: incidence and implications of individual specialization. Am. Nat. 161, 1–
497 28. doi:10.1086/343878
498 Broadhurst, M., Orme, C.D.L., 2014. Spatial and temporal benthic species assemblage responses with
499 a deployed marine tidal energy device: A small scaled study. Mar. Environ. Res. 99, 76–84.
500 doi:10.1016/j.marenvres.2014.03.012 501 Brown, K.M., Ewins,ACCEPTED P.J., 1996. Technique-dependent biases in determination of diet composition: 502 An example with Ring-Billed Gulls. Condor 98, 34–41. doi:10.2307/1369505
503 Cairns, D.K., 1992. Diving behavior of Black Guillemots in Northeastern Hudson-Bay. Colon.
504 Waterbirds 15, 245–248. doi:10.2307/1521461
505 Cairns, D.K., 1987. The ecology and energetics of chick provisioning by Black Guillemots. Condor 89, 22 ACCEPTED MANUSCRIPT 506 627–635. doi:10.2307/1368652
507 Cairns, D.K., 1980. Nesting density , habitat structure and human disturbance as factors in Black
508 Guillemot reproduction. Wilson Bull. 92, 352–361.
509 Caron-Beaudoin, É., Gentes, M.L., Patenaude-Monette, M., Hélie, J.F., Giroux, J.F., Verreault, J.,
510 2013. Combined usage of stable isotopes and GPS-based telemetry to understand the feeding
511 ecology of an omnivorous bird, the Ring-billed Gull (Larus delawarensis). Can. J. Zool. 91, 689–
512 697. doi:10.1675/1524-4695
513 Chase, J.M., 2011. Ecological niche theory, in: The Theory of Ecology. pp. 93–108.
514 Chimienti, M., Cornulier, T., Owen, E., Bolton, M., Davies, I.M., Travis, J.M.J., Scott, B.E., 2016. The
515 use of an unsupervised learning approach for characterizing latent behaviors in accelerometer
516 data. Ecol. Evol. 6, 727–741. doi:10.1002/ece3.1914 517 Churchfield, M.J., Li, Y., Moriarty, P.J., 2013. A large-eddyMANUSCRIPT simulation study of wake propagation and 518 power production in an array of tidal-current turbi nes. Philos. Trans. R. Soc. A Math. Phys. Eng.
519 Sci. 371, 20120421–20120421. doi:10.1098/rsta.2012.0421
520 Clode, D., Macdonald, D.W., 2002. Invasive predators and the conservation of island birds: The case
521 of American Mink Mustela vison and terns Sterna spp. in the Western Isles, Scotland. Bird
522 Study 49, 118–123. doi:10.1080/00063650209461255
523 Coulson, J.C., 2016. A review of philopatry in seabirds and comparisons with other waterbird species. 524 Waterbirds ACCEPTED39, 229–240. doi:10.1675/063.039.0302 525 Craik, C., 1997. Long-term effects of North American Mink Mustela vison on seabirds in western
526 Scotland. Bird Study 44, 303–309. doi:10.1080/00063659709461065
527 Craik, C., 1995. Effects of North American mink on the breeding success of terns and smaller gulls in
528 west Scotland. Seabird 17, 3–11. 23 ACCEPTED MANUSCRIPT 529 Daunt, F., Wanless, S., Greenstreet, S.P.R., Jensen, H., Hamer, K.C., Harris, M.P., 2008. The impact of
530 the sandeel fishery closure in the northwestern North Sea on seabird food consumption,
531 distribution and productivity. Can. J. Fish. Aquat. Sci. 65, 362–381. doi:10.1139/f07-164
532 Davies, I.M., Watret, R., Gubbins, M., 2014. Spatial planning for sustainable marine renewable
533 energy developments in Scotland. Ocean Coast. Manag. 99, 72–81.
534 doi:10.1016/j.ocecoaman.2014.05.013
535 Divoky, G.J., Douglas, D.C., Stenhouse, I.J., 2016. Arctic sea ice a major determinant in Mandt ’ s
536 Black Guillemot movement and distribution during non-breeding season. Biol. Lett. 12,
537 20160275. doi:http://dx.doi.org/10.1098/rsbl.2016.0275
538 EcoWatt 2050, 2017. EcoWatt 2050 Project Summary 2017.
539 Edmunds, M., Malki, R., Williams, A.J., Masters, I., Croft, T.N., 2014. Aspects of tidal stream turbine
540 modelling in the natural environment using a coupled BEM-CFD model. Int. J. Mar. Energy 7, 541 20–42. doi:10.1016/j.ijome.2014.07.001 MANUSCRIPT
542 Elliott, K.H., Woo, K., Gaston, A.J., Benvenuti, S., Dall’Antonia, L., Davoren, G.K., 2008. Seabird
543 foraging behaviour indicates prey type. Mar. Ecol. Prog. Ser. 354, 289–303.
544 doi:10.3354/meps07221
545 Embling, C.B., Illian, J., Armstrong, E., van der Kooij, J., Sharples, J., Camphuysen, K.C.J., Scott, B.E.,
546 2012. Investigating fine-scale spatio-temporal predator-prey patterns in dynamic marine
547 ecosystems: A functional data analysis approach. J. Appl. Ecol. 49, 481–492.
548 doi:10.1111/j.1365-2664.2012.02114.xACCEPTED
549 Ewins, P.J., 1992. Growth of Black Guillemot Cepphus grylle chicks in Shetland in 1983-84. Seabird
550 14, 3–14.
551 Ewins, P.J., 1990. The diet of Black Guillemots Cepphus grylle in Shetland. Holarct. Ecol. 13, 90–97. 24 ACCEPTED MANUSCRIPT 552 doi:10.1111/j.1600-0587.1990.tb00593.x
553 Ewins, P.J., 1988. An analysis of ringing recoveries of Black Guillemots Cepphus grylle in Britain and
554 Ireland. Ringing Migr. 9, 95–102. doi:10.1080/03078698.1988.9673932
555 Ewins, P.J., 1986. The ecology of Black Guillemots Cepphus grylle in Shetland. PhD Thesis. University
556 of Oxford.
557 Ewins, P.J., 1985. Colony attendance and censusing of Black Guillemots. Bird Study 32, 176–185.
558 doi:10.1080/00063658509476876
559 Ewins, P.J., Kirk, D.A., 1988. The distribution of Shetland Black Guillemots Cepphus grylle outside the
560 breeding season. Seabird 11, 50–61.
561 Ewins, P.J., Tasker, M.L., 1985. The breeding distribution of Black Guillemots Cepphus grylle in
562 Orkney and Shetland, 1982-84. Bird Study 32, 37–41. doi:10.1080/00063658509476877 563 Fox, C.J., Benjamins, S., Masden, E.A., Miller, R., 2018. MANUSCRIPT Challenges and opportunities in monitoring 564 the impacts of tidal-stream energy devices on marine vertebrates. Renew. Sustain. Energy Rev.
565 doi:10.1016/j.rser.2017.06.004
566 Fraenkel, P.L., 2006. Tidal current energy technologies. Ibis (Lond. 1859). 148, 145–151.
567 doi:10.1111/j.1474-919X.2006.00518.x
568 Frederiksen, M., Petersen, A., 2000. The importance of natal dispersal in a colonial seabird, the Black
569 Guillemot Cepphus grylle. Ibis (Lond. 1859). 142, 48–57. doi:10.1111/j.1474- 570 919X.2000.tb07683.xACCEPTED
571 Frederiksen, M., Petersen, A., 1999. Philopatry and dispersal within a black guillemot colony.
572 Waterbirds 22, 274–281. doi:10.2307/1522215
573 Furness, R.W., 2002. Management implications of interactions between fisheries and sandeel- 25 ACCEPTED MANUSCRIPT 574 dependent seabirds and seals in the North Sea. ICES J. Mar. Sci. 59, 261–269.
575 doi:10.1006/jmsc.2001.1155
576 Furness, R.W., Barrett, R.T., 1985. The food-requirements and ecological relationships of a seabird
577 community in North Norway. Ornis Scand. 16, 305–313. doi:10.2307/3676695
578 Furness, R.W., Wade, H.M., Robbins, A.M.C., Masden, E.A., 2012. Assessing the sensitivity of seabird
579 populations to adverse effects from tidal stream turbines and wave energy devices. ICES J. Mar.
580 Sci. 69, 1466–1479. doi:10.1093/icesjms/fss131
581 Gaston, A.J., Mclaren, P.L., 1990. Winter observations of Black Guillemots. Stud. Avian Biol 67–70.
582 Golet, G., Seiser, P., McGuire, A., Roby, D., Fischer, J., Kuletz, K., Irons, D., Dean, T., Jewett, S.,
583 Newman, S., 2002. Long-term direct and indirect effects of the “Exxon Valdez” oil spill on
584 Pigeon Guillemots in Prince William Sound, Alaska. Mar. Ecol. Prog. Ser. 241, 287–304.
585 doi:10.3354/meps241287 MANUSCRIPT 586 Grecian, W.J., Inger, R., Attrill, M.J., Bearhop, S., Godley, B.J., Witt, M.J., Votier, S.C., 2010. Potential
587 impacts of wave-powered marine renewable energy installations on marine birds. Ibis (Lond.
588 1859). 152, 683–697. doi:10.1111/j.1474-919X.2010.01048.x
589 Greenwood, J.G., 2002. Nest cavity choice by Black Guillemots Cepphus grylle. Atl. Seabirds 4, 119–
590 122.
591 Greenwood, P.J., 1980. Mating systems, philopatry and dispersal in birds and mammals. Anim. 592 Behav. 28, 1140–1162.ACCEPTED doi:10.1016/S0003-3472(80)80103-5 593 Grémillet, D., Boulinier, T., 2009. Spatial ecology and conservation of seabirds facing global climate
594 change: A review. Mar. Ecol. Prog. Ser. 391, 121–137. doi:10.3354/meps08212
595 Hario, M., 2001. Chick growth and nest departure in Baltic Black Guillemots Cepphus grylle. Ornis
596 Fenn. 78, 97–108. 26 ACCEPTED MANUSCRIPT 597 Harris, M.P., Daunt, F., Bogdanova, M.I., Lahoz-Monfort, J.J., Newell, M.A., Phillips, R.A., Wanless, S.,
598 2013. Inter-year differences in survival of Atlantic puffins Fratercula arctica are not associated
599 with winter distribution. Mar. Biol. 160, 2877–2889. doi:10.1007/s00227-013-2278-5
600 Harris, M.P., Halley, D.J., Wanless, S., 1996. Philopatry in the Common Guillemot Uria aalge. Bird
601 Study 43, 134–137. doi:10.1080/00063659609461005
602 Harris, M.P., Riddiford, N.J., 1989. The food of some young seabirds on Fair Isle in 1986-88. Scottish
603 Birds 15, 119–125.
604 Harris, M.P., Wanless, S., Ballesteros, M., Moe, B., Daunt, F., Erikstad, K.E., 2015. Geolocators reveal
605 an unsuspected moulting area for Isle of May Common Guillemots Uria aalge. Bird Study 62,
606 267–270. doi:10.1080/00063657.2015.1006164
607 Heinänen, S., Žydelis, R., Dorsch, M., Nehls, G., Skov, H., 2017. High-resolution sea duck distribution
608 modeling: Relating aerial and ship survey data to food resources, anthropogenic pressures, and 609 topographic variables. Condor 119, 175–190. doi:10. MANUSCRIPT1650/CONDOR-16-57.1
610 Heubeck, M., 1995. Shetland beached bird surveys: national and European context. Proc. R. Soc.
611 Edinburgh 103, 165–179.
612 Hilden, O., 1994. Diurnal rhythm of colony attendance and optimal census time for the black
613 guillemot Cepphus grylle in the Baltic Sea. Ornis Fenn. 71, 61–67.
614 Inger, R., Attrill, M., Bearhop, S., Broderick, A., James Grecian, W., Hodgson, D., Mills, C., Sheehan, E., 615 Votier, S., Witt,ACCEPTED M., Godley, B., 2009. Marine renewable energy: potential benefits to 616 biodiversity? An urgent call for research. J. Appl. Ecol.
617 Jessopp, M.J., Cronin, M., Doyle, T.K., Wilson, M., McQuatters-Gollop, A., Newton, S., Phillips, R.A.,
618 2013. Transatlantic migration by post-breeding puffins: A strategy to exploit a temporarily
619 abundant food resource? Mar. Biol. 160, 2755–2762. doi:10.1007/s00227-013-2268-7 27 ACCEPTED MANUSCRIPT 620 Kamenos, N.A., Moore, P.G., Hall-Spencer, J.M., 2004. Small-scale distribution of juvenile gadoids in
621 shallow inshore waters; what role does maerl play? ICES J. Mar. Sci. 61, 422–429.
622 doi:10.1016/j.icesjms.2004.02.004
623 Katzir, G., Howland, H.C., 2003. Corneal power and underwater accommodation in great cormorants
624 (Phalacrocorax carbo sinensis). J. Exp. Biol. 206, 833–841. doi:10.1242/jeb.00142
625 Ladd, C., Jahncke, J., Hunt, G.L., Coyle, K.O., Stabeno, P.J., 2005. Hydrographic features and seabird
626 foraging in Aleutian Passes. Fish. Oceanogr. 14, 178–195. doi:10.1111/j.1365-
627 2419.2005.00374.x
628 Laich, A.G., Wilson, R.P., Quintana, F., Shepard, E.L.C., 2010. Identification of imperial cormorant
629 Phalacrocorax atriceps behaviour using accelerometers. Endanger. Species Res. 10, 29–37.
630 doi:10.3354/esr00091
631 Langton, R., Davies, I.M., Scott, B.E., 2011. Seabird conservation and tidal stream and wave power 632 generation: Information needs for predicting and MANUSCRIPT managing potential impacts. Mar. Policy 35, 633 623–630. doi:10.1016/j.marpol.2011.02.002
634 Lascelles, B.G., Taylor, P.R., Miller, M.G.R., Dias, M.P., Oppel, S., Torres, L., Hedd, A., Le Corre, M.,
635 Phillips, R.A., Shaffer, S.A., Weimerskirch, H., Small, C., 2016. Applying global criteria to tracking
636 data to define important areas for marine conservation. Divers. Distrib. 22, 422–431.
637 doi:10.1111/ddi.12411
638 Lavers, J.L., Jones, I.L., Diamond, A.W., 2007. Natal and breeding dispersal of razorbills (Alca torda) in
639 Eastern NorthACCEPTED America. Waterbirds 30, 588–594. doi:10.1675/1524-
640 4695(2007)030[0588:NABDOR]2.0.CO;2
641 Linnebjerg, J.F., Fort, J., Guilford, T., Reuleaux, A., Mosbech, A., Frederiksen, M., 2013. Sympatric
642 breeding auks shift between dietary and spatial resource rartitioning across the annual cycle. 28 ACCEPTED MANUSCRIPT 643 PLoS One 8, e72987. doi:10.1371/journal.pone.0072987
644 Litzow, M.A., Piatt, J.F., Abookire, A.A., Speckman, S.G., Arimitsu, M.L., Figurski, J.D., 2004.
645 Spatiotemporal predictability of schooling and nonschooling prey of Pigeon Guillemots. Condor
646 106, 410–415.
647 Litzow, M.A., Piatt, J.F., Abookire, A., Prichard, A., Robards, M., 2000. Monitoring temporal and
648 spatial variability in sandeel (Ammodytes hexapterus) abundance with pigeon guillemot
649 (Cepphus columba) diets. ICES J. Mar. Sci. 57, 976–986. doi:10.1006/jmsc.2000.0583
650 Litzow, M.A., Piatt, J.F., Figurski, J.D., 1998. Hermit crabs in the diet of Pigeon Guillemots at
651 Kachemak Bay, Alaska. Colon. Waterbirds 21, 242–244. doi:10.2307/1521913
652 Lovvorn, J., Liggins, G. a, Borstad, M.H., Calisal, S.M., Mikkelsen, J., 2001. Hydrodynamic drag of
653 diving birds: effects of body size, body shape and feathers at steady speeds. J. Exp. Biol. 204,
654 1547–1557. MANUSCRIPT 655 Lowe, J.A., Gregory, J.M., Flather, R.A., 2001. Changes in the occurrence of storm surges around the
656 United Kingdom under a future climate scenario using a dynamic storm surge model driven by
657 Hadley Centre climate models. Clim. Dyn. 18, 179–188. doi:10.1007/s003820100163
658 MacDonald, A., Heath, M., Edwards, M., Furness, R., Pinnegar, J.K., Wanless, S., Speirs, D.,
659 Greenstreet, S., 2015. Climate driven trophic cascades affecting seabirds around the British
660 Isles. Oceanogr. Mar. Biol. - An Annu. Rev. 53, 55–79. doi:10.1201/b18733-3 661 Magnusdottir, R.,ACCEPTED von Schmalensee, M., Stefansson, R.A., Macdonald, D.W., Hersteinsson, P., 2014. 662 A foe in woe: American mink (Neovison vison) diet changes during a population decrease.
663 Mamm. Biol. 79, 58–63. doi:10.1016/j.mambio.2013.08.002
664 Martin, G.R., 2011. Understanding bird collisions with man-made objects: A sensory ecology
665 approach. Ibis (Lond. 1859). doi:10.1111/j.1474-919X.2011.01117.x 29 ACCEPTED MANUSCRIPT 666 Martin, G.R., Wanless, S., 2015. The visual fields of Common Guillemots Uria aalge and Atlantic
667 Puffins Fratercula arctica: Foraging, vigilance and collision vulnerability. Ibis (Lond. 1859). 157,
668 798–807. doi:10.1111/ibi.12297
669 Masden, E.A., Foster, S., Jackson, A.C., 2013. Diving behaviour of Black Guillemots Cepphus grylle in
670 the Pentland Firth, UK: potential for interactions with tidal stream energy developments. Bird
671 Study 60, 547–549. doi:10.1080/00063657.2013.842538
672 McFarlane Tranquilla, L.A., Montevecchi, W.A., Fifield, D.A., Hedd, A., Gaston, A.J., Robertson, G.J.,
673 Phillips, R.A., 2014. Individual winter movement strategies in two species of murre (Uria spp.)
674 in the Northwest Atlantic. PLoS One 9, e90583. doi:10.1371/journal.pone.0090583
675 Mehlum, F., Gabrielsen, G.W., 1993. Energy expenditure by black guillemots (Cepphus grylle) during
676 chick-rearing. Colon. Waterbirds 16, 45–52. doi:10.2307/1521555
677 MeyGen, 2012. MeyGen Tidal Energy Project Phase 1 Environmental Statement. MANUSCRIPT 678 Miller, R.G., Hutchison, Z.L., Macleod, A.K., Burrows, M.T., Cook, E.J., Last, K.S., Wilson, B., 2013.
679 Marine renewable energy development: Assessing the Benthic Footprint at multiple scales.
680 Front. Ecol. Environ. 11, 433–440. doi:10.1890/120089
681 Mitchell, P.I., Newton, S.F., Ratcliffe, N., Dunn, T.E., 2004. Seabird Populations of Britain and Ireland:
682 results of the Seabird 2000 census.
683 Moore, C.G., Roberts, J.M., 2011. An assessment of the conservation importance of species and 684 habitats identifiedACCEPTED during a series of recent research cruises around Scotland. SNH 685 Commissioned Report No. 446.
686 Moum, T., Árnason, E., 2001. Genetic diversity and population history of two related seabird species
687 based on mitochondrial DNA control region sequences. Mol. Ecol. 10, 2463–2478.
688 doi:10.1046/j.0962-1083.2001.01375.x 30 ACCEPTED MANUSCRIPT 689 Nol, E., Gaskin, D.E., 1987. Distribution and movement of Black Guillemots (Cepphus grylle) in coastal
690 waters of the southwestern Bay of Fundy, Canada. Can. J. Zool. 65, 2682–2689.
691 doi:10.1139/z87-407
692 Nordström, M., 2002. Effects of feral mink removal on seabirds , waders and passerines on small
693 islands in the Baltic Sea. Biological 109, 3336550.
694 Nordström, M., Korpimäki, E., 2004. Effects of island isolation and feral mink removal on bird
695 communities on small islands in the Baltic Sea. J. Anim. Ecol. 73, 424–433. doi:10.1111/j.0021-
696 8790.2004.00816.x
697 Owen, E., 2015. Black guillemot (Cepphus grylle) tracking in Orkney, 2013 and 2014. Scottish Natural
698 Heritage Commissioned Report No. 903.
699 Petersen, Æ., 1981. Breeding biology and feeding ecology of Black guillemots. University of Oxford. 700 Piatt, J.F., Nettleship, D.N., 1985. Diving depths of fourMANUSCRIPT alcids. Am. Ornithol. Union 102, 293–297. 701 doi:10.2307/4086771
702 Potvin, D.A., Välimäki, K., Lehikoinen, A., 2016. Differences in shifts of wintering and breeding ranges
703 lead to changing migration distances in European birds. J. Avian Biol. 47, 619–628.
704 doi:10.1111/jav.00941
705 Prach, R.W., Smith, A.R., 1992. Breeding distribution and numbers of Black Guillemots in Jones
706 Sound, NWT. Arctic 45, 111–114.
707 Regular, P.M., Hedd,ACCEPTED A., Montevecchi, W.A., 2011. Fishing in the dark: A pursuit-diving seabird 708 modifies foraging behaviour in response to nocturnal light levels. PLoS One 6, e26763.
709 doi:10.1371/journal.pone.0026763
710 Robbins, A.M.C., 2017. Seabird ecology in high-energy environments : approaches to assessing
711 impacts of marine renewables. PhD Thesis. University of Glasgow. 31 ACCEPTED MANUSCRIPT 712 Robbins, A.M.C., 2012. Analysis of bird and marine mammal data for the Fall of Warness tidal test
713 site, Orkney. Scottish Natural Heritage Commissioned Report No. 614.
714 Rollings, E., 2011. MeyGen phase 1 EIA scoping document.
715 Ronconi, R.A., Clair, C.C.S., 2002. Management options to reduce boat disturbance on foraging black
716 guillemots (Cepphus grylle) in the Bay of Fundy. Biol. Conserv. 108, 265–271.
717 doi:10.1016/S0006-3207(02)00126-X
718 Ronconi, R.A., Lascelles, B.G., Langham, G.M., Reid, J.B., Oro, D., 2012. The role of seabirds in Marine
719 Protected Area identification, delineation, and monitoring: Introduction and synthesis. Biol.
720 Conserv. doi:10.1016/j.biocon.2012.02.016
721 Sato, K., Daunt, F., Watanuki, Y., Takahashi, A., Wanless, S., 2008. A new method to quantify prey
722 acquisition in diving seabirds using wing stroke frequency. J. Exp. Biol. 211, 58–65.
723 doi:10.1242/jeb.009811 MANUSCRIPT 724 Savidge, G., Ainsworth, D., Bearhop, S., Christen, N., Elsaesser, B., Fortune, F., Inger, R., Kennedy, R.,
725 McRobert, A., Plummer, K.E., Pritchard, D.W., Sparling, C.E., Whittaker, T.J.T., 2014. Strangford
726 Lough and the SeaGen tidal turbine. Mar. Renew. Energy Technol. Environ. Interact. Humanit.
727 Sea 153–172. doi:10.1007/978-94-017-8002-5
728 Sawyer, T.R., 1999. Habitat use and breeding performance in an inshore foraging seabird, the Black
729 Guillemot Cepphus grylle. University of Glasgow. 730 Scotrenewables, ACCEPTED2017. SR2000 [WWW Document]. URL 731 http://www.scotrenewables.com/technology-development/sr2000 (accessed 10.28.17).
732 Scott, B.E., Langton, R., Philpott, E., Waggitt, J.J., 2014. Seabirds and Marine Renewables: Are we
733 asking the right questions?, in: Marine Renewable Energy Technology and Environmental
734 Interactions. pp. 81–92. doi:10.1007/978-94-017-8002-5 32 ACCEPTED MANUSCRIPT 735 Shields, A.M.A., Woolf, D.K., Grist, E.P.M.M., Kerr, S.A., C, A., Harris, R.E., Bell, M.C., Beharie, R.,
736 Want, A., Gibb, S.W., Side, J., Shields, M.A., Woolf, D.K., Grist, E.P.M.M., Kerr, S.A., Jackson,
737 A.C., Harris, R.E., Bell, M.C., Beharie, R., Want, A., Osalusi, E., Gibb, S.W., Side, J., 2011. Marine
738 renewable energy: The ecological implications of altering the hydrodynamics of the marine
739 environment. Ocean Coast. Manag. 54, 2–9. doi:10.1016/j.ocecoaman.2010.10.036
740 Shields, M.A., Dillon, L.J., Woolf, D.K., Ford, A.T., 2009. Strategic priorities for assessing ecological
741 impacts of marine renewable energy devices in the Pentland Firth (Scotland, UK). Mar. Policy
742 33, 635–642. doi:10.1016/j.marpol.2008.12.013
743 Shoji, A., Elliott, K.H., Greenwood, J.G., McClean, L., Leonard, K., Perrins, C.M., Fayet, A., Guilford, T.,
744 2015. Diving behaviour of benthic feeding Black Guillemots. Bird Study 3657, 1–6.
745 doi:10.1080/00063657.2015.1017800
746 Shorty, J.T., Gannon, D.P., 2013. Habitat selection by the Rock Gunnel, Pholis gunnellus L. (Pholidae). 747 Northeast. Nat. 20, 155–170. doi:10.1656/045.020.01 MANUSCRIPT13 748 Slater, P.J.B., Slater, E.P., 1972. Behaviour of the Tystie during feeding of the young. Bird Study 19,
749 105–113. doi:10.1080/00063657209476332
750 SNH, 2016. Assessing collision risk between underwater turbines and marine wildlife. SNH guidance
751 note.
752 SNH, 2014. SNH’s advice on selected responses to the 2013 Marine Scotland consultation on Nature
753 Conservation Marine Protected Areas (MPAs)., Scottish Natural Heritage Commissioned Report
754 No. 747. ACCEPTED
755 Soanes, L.M., Arnould, J.P.Y., Dodd, S.G., Sumner, M.D., Green, J.A., 2013. How many seabirds do we
756 need to track to define home-range area? J. Appl. Ecol. 50, 671–679. doi:10.1111/1365-
757 2664.12069 33 ACCEPTED MANUSCRIPT 758 St. John, M.A., Harrison, P.J., Parsons, T.R., 1992. Tidal wake-mixing: localized effects on primary
759 production and zooplankton distributions in the Strait of Georgia, British Columbia. J. Exp. Mar.
760 Bio. Ecol. 164, 261–274. doi:10.1016/0022-0981(92)90179-E
761 Strod, T., Arad, Z., Izhaki, I., Katzir, G., 2004. Cormorants keep their power: Visual resolution in a
762 pursuit-diving bird under amphibious and turbid conditions. Curr. Biol.
763 doi:10.1016/j.cub.2004.05.009
764 Thaxter, C.B., Lascelles, B., Sugar, K., Cook, A., Roos, S., Bolton, M., Langston, R.H.W., Burton, N.H.K.,
765 2012. Seabird foraging ranges as a tool for identifying Marine Protected Areas. Biol. Conserv.
766 156, 53–61.
767 The Crown Estate, 2016. Crown Estate tidal and wave lease areas | Marine Scotland Information
768 [WWW Document]. URL http://marine.gov.scot/information/crown-estate-tidal-and-wave-
769 lease-areas (accessed 4.21.17).
770 Veit, R.R., Manne, L.L., 2015. Climate and changing winter MANUSCRIPT distribution of alcids in the Northwest 771 Atlantic. Front. Ecol. Evol. 3, 1–9. doi:10.3389/fevo.2015.00038
772 Vermeer, K., Morgan, K.H., Smith, G.E.J., 1993. Attendance of Pigeon Guillemots as related to tide
773 height and time of day. Colon. Waterbirds 16, 1–8. doi:10.2307/1521550
774 Votier, S.C., Bearhop, S., Witt, M.J., Inger, R., Thompson, D., Newton, J., 2010. Individual responses
775 of seabirds to commercial fisheries revealed using GPS tracking, stable isotopes and vessel
776 monitoring systems. J. Appl. Ecol. 47, 487–497. doi:10.1111/j.1365-2664.2010.01790.x ACCEPTED 777 Wade, H.M., 2015. Investigating the potential effects of marine renewable energy developments on
778 seabirds. PhD Thesis. University of Aberdeen.
779 Waggitt, J.J., Cazenave, P.W., Torres, R., Williamson, B.J., Scott, B.E., 2016a. Predictable
780 hydrodynamic conditions explain temporal variations in the density of benthic foraging 34 ACCEPTED MANUSCRIPT 781 seabirds in a tidal stream environment. ICES J. Mar. Sci. 73, 2677–2686.
782 doi:10.1093/icesjms/fsw100
783 Waggitt, J.J., Cazenave, P.W., Torres, R., Williamson, B.J., Scott, B.E., Siriwardena, G., Torres, P.,
784 Williamson, B.J., Scott, B.E., 2016b. Quantifying pursuit-diving seabirds’ associations with fine-
785 scale physical features in tidal stream environments. J. Appl. Ecol. 53, 1653–1666.
786 doi:10.1111/1365-2664.12646
787 Waggitt, J.J., Robbins, A.M.C., Wade, H.M., Masden, E.A., Furness, R.W., Jackson, A.C., Scott, B.E.,
788 2017. Comparative studies reveal variability in the use of tidal stream environments by
789 seabirds. Mar. Policy 81, 143–152. doi:10.1016/j.marpol.2017.03.023
790 Waggitt, J.J., Scott, B.E., 2014. Using a spatial overlap approach to estimate the risk of collisions
791 between deep diving seabirds and tidal stream turbines: A review of potential methods and
792 approaches. Mar. Policy 44, 90–97. doi:10.1016/j.marpol.2013.07.007
793 Watanuki, Y., Daunt, F., Takahashi, A., Newell, M., Wanless, MANUSCRIPT S., Sato, K., Miyazaki, N., 2008. 794 Microhabitat use and prey capture of a bottom-feeding top predator, the European shag,
795 shown by camera loggers. Mar. Ecol. Prog. Ser. 356, 283–293. doi:10.3354/meps07266
796 Watanuki, Y., Wanless, S., Harris, M., Lovvorn, J.R., Miyazaki, M., Tanaka, H., Sato, K., 2006. Swim
797 speeds and stroke patterns in wing-propelled divers: a comparison among alcids and a penguin.
798 J. Exp. Biol. 209, 1217–1230. doi:10.1242/jeb.02128
799 Weimerskirch, H., 1998. How can a pelagic seabird provision its chick when relying on a distant food
800 resource? CyclicACCEPTED attendance at the colony, foraging decision and body condition in sooty
801 shearwaters. J. Anim. Ecol. 67, 99–109. doi:10.1046/j.1365-2656.1998.00180.x
802 Weslawski, J.M., Stempniewicz, L., Galaktionov, K., 1994. Summer diet of seabirds from the Franz-
803 Josef-Land Archipelago, Russian Arctic. Polar Res. 13, 173–181. doi:10.1111/j.1751- 35 ACCEPTED MANUSCRIPT 804 8369.1994.tb00447.x
805 Wilhelmsson, D., Malm, T., Öhman, M.C., 2006. The influence of offshore windpower on demersal
806 fish. ICES J. Mar. Sci. 63, 775–784. doi:10.1016/j.icesjms.2006.02.001
807 Wilson, B., Batty, R., Daunt, F., Carter, C., 2006. Collision risks between marine renewable energy
808 devices and mammals, fish and diving birds 1–105.
809 Wink, M., 2006. Use of DNA markers to study bird migration. J. Ornithol. 147, 234–244.
810 doi:10.1007/s10336-006-0065-5
811 Woo, K.J., Elliott, K.H., Davidson, M., Gaston, A.J., Davoren, G.K., 2008. Individual specialization in
812 diet by a generalist marine predator reflects specialization in foraging behaviour. J. Anim. Ecol.
813 77, 1082–1091. doi:10.1111/j.1365-2656.2008.01429.x
814 815 MANUSCRIPT 816
ACCEPTED ACCEPTED MANUSCRIPT • Black guillemots have been highlighted as at risk to tidal stream turbines (TSTs). • Further knowledge of foraging is required to assess the risk of overlap with TSTs. • We synthesise the knowledge of black guillemot ecology and its relevance to TSTs. • Aspects of black guillemot foraging, diving, and seasonal movements are reviewed. • Disturbance to nesting adults and processes of colony recovery are considered.
MANUSCRIPT
ACCEPTED