Arctic Science

Spring distribution of ringed seals (Pusa hispida) in Eclipse Sound and , : implications for potential ice-breaking activities

Journal: Arctic Science

Manuscript ID AS-2018-0020.R2

Manuscript Type: Note

Date Submitted by the 27-Sep-2018 Author:

Complete List of Authors: Yurkowski, David; University of Manitoba, Department of Biological Sciences; Fisheries and Oceans Canada Central and Arctic Region Young, Brent;Draft Fisheries and Oceans Canada Central and Arctic Region Dunn, Blair; Fisheries and Oceans Canada Central and Arctic Region Ferguson, Steven; Fisheries and Oceans Canada Central and Arctic Region

Aerial survey, Anthropogenic stressor, Conservation, Hotspot, Infrared Keyword: imagery

Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue?:

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1 Spring distribution of ringed seals (Pusa hispida) in Eclipse Sound and Milne 2 Inlet, Nunavut: implications for potential ice-breaking activities 3 4 David J. Yurkowski*1,2, Brent G. Young2, J. Blair Dunn2, and Steven H. Ferguson1,2 5 6 1 Department of Biological Sciences, University of Manitoba, Winnipeg, Manitoba, R3T 7 2N2, Canada (DJY: [email protected]) 8 2 Fisheries and Oceans Canada, Winnipeg, Manitoba, R3T 2N6, Canada (BG: 9 [email protected]; BD: [email protected]; SHF: 10 [email protected]) 11 12 Corresponding author*: David Yurkowski – Email: [email protected] 13

14 Type of article: Research Note

15 Number of tables: 1

16 Number of figures: 2

17 Number of references: 31 Draft

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18 Abstract

19 Resource development in Arctic waters is proceeding rapidly leading to increased

20 interactions with Arctic wildlife. As sea ice extent decreases, the demand for shipping

21 and ice-breaking operations will expand into winter and spring with greater impact on

22 ice-dependent pinnipeds. However, knowledge of the distribution of these species, such

23 as ringed seals (Pusa hispida), during spring within areas of resource development is

24 lacking. Baffinland’s Mary River iron ore port in southern Milne Inlet, Nunavut opened in

25 2015 with proposed ice-breaking activities in spring – an important period in ringed seal

26 seasonal life-history. We conducted infrared and photographic aerial surveys in June

27 2016 and 2017 to overlay the proposed ice-breaking route with ringed seal hotspots (i.e.

28 areas of higher density). We identifiedDraft four areas of overlap where proposed ice-

29 breaking would traverse through ringed seal hotspots: eastern and western Eclipse

30 Sound (a ringed seal pupping ground identified by local knowledge), middle of Milne

31 Inlet, and southern Milne Inlet. We identified potential negative implications of spring

32 ice-breaking operations on ringed seals such as displacement, separation of mothers

33 and pups, destruction of resting and birth lairs, and vessel-seal collisions. Results are

34 relevant to policy decision-makers who can develop mitigation strategies in the rapidly

35 thawing and developing Arctic.

36

37 Key words: Aerial survey, Anthropogenic stressor, Conservation, Hotspot, Infrared

38 imagery

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39 Introduction

40 The distribution of animals is affected by a broad suite of abiotic, biotic and

41 anthropogenic factors that influence habitat use at numerous spatiotemporal scales

42 (Levin 1992). Therefore, quantifying animal space-use and density patterns is critical to

43 conservation and management, particularly in environments undergoing rapid change

44 such as the Arctic marine ecosystem (IPCC 2013). Climate change is most pronounced

45 in the Arctic and receding sea ice allows a longer season of anthropogenic

46 disturbances, such as shipping and resource development, placing new stresses on

47 Arctic wildlife (Post et al. 2013; Pizzolato et al. 2014). For example, shipping in Arctic

48 waters is developing rapidly with increases in polar tourism, and natural resource

49 extraction leading to increased interactionsDraft with Arctic marine mammals. Possible

50 interactions include the potential for collisions, higher noise levels in the Arctic

51 soundscape, and destruction of sea ice habitat through spring ice-breaking activities

52 (Reeves et al. 2014; Laidre et al. 2015; Halliday et al. 2017; Hauser et al. 2018).

53 Pagophilic (i.e. ice-dependent) pinnipeds can be highly sensitive to ice-breaking

54 activities, which for example in Caspian seals (Pusa capsica), has resulted in increased

55 mortality rates (Harkonen et al. 2008), as well as increased the potential for non-

56 consumptive (i.e. sub-lethal) effects (e.g. higher incidences of mother-pup separation;

57 Wilson et al. 2017). Ringed seals, who are endemic to the circumpolar Arctic and are a

58 pillar to Inuit culture, typically rely on shore-fast ice for building subnivean resting and

59 birthing lairs in winter and spring, and as a basking platform during their annual spring

60 molt (McLaren 1958; Smith 1987; Kenny and Chan 2017). However, ringed seal

61 distribution and hotspots (i.e. areas of significantly higher density) during spring within

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62 areas of rapid resource development and potential ice-breaking activities is poorly

63 documented.

64 Baffinland’s Mary River iron ore mine and port in southern Milne Inlet, Nunavut

65 (see Fig. 1) opened in 2015 with high shipping activity occurring during the open water

66 season and proposed ice-breaking activities in spring – an important period in ringed

67 seals’ seasonal life-history due to their establishment of territories and subsequent

68 parturition, nursing, breeding and basking activities (McLaren 1958; Smith 1987; See

69 Table 1). Therefore, ice-breaking activities in the Eclipse Sound area, which includes

70 , Tremblay Sound, Milne Inlet, Koluktoo Bay and Eclipse Sound could

71 have negative implications on ringed seal productivity and the population.

72 Here, we assessed the springDraft distribution of ringed seals in the Eclipse Sound

73 area by performing infrared and photographic aerial surveys in June 2016 and 2017.

74 We quantified inter-annual variability of ringed seal hotspots between 2016 and 2017,

75 and compared the proposed ice-breaking shipping route with ringed seal surface

76 density. Our hotspot methods can be used in future research to inform decisions related

77 to potential ice-breaking activities in Arctic regions undergoing increased shipping and

78 for the development of mitigation strategies.

79

80 Materials and methods

81 Survey design

82 Aerial surveys were conducted in four strata: Eclipse Sound, Milne Inlet,

83 Tremblay Sound, and Navy Board Inlet, Nunavut. Surveys were flown in June of 2016

84 (June 17 to 22) and 2017 (June 6 to 8) using a DeHavilland Twin Otter (DH-6) equipped

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85 with bubble windows and a camera port at the rear underbelly of the plane. In 2016,

86 most snow cover had melted by the time each survey was conducted while in 2017,

87 there was still significant snow cover at the time of the surveys. However, both surveys

88 were flown after the ringed seal subnivean period. In addition, the distribution of ringed

89 seals and their hotspots during the basking period in June are also likely similar to that

90 in March, as ringed seals exhibit minimal movement and small home ranges throughout

91 the ice-covered period (Luque et al. 2014; Yurkowski et al. 2016). A Global Positioning

92 System (GPS) was used to log the position, altitude, speed and heading of the aircraft

93 every second. Surveys were flown at a target ground speed of 204 km/h (110 knots)

94 and a target altitude of 305 m (1000 ft). In 2016, all four survey strata were completed

95 and repeated (see Fig. 1). In 2017, oneDraft survey of Eclipse Sound and Navy Board Inlet

96 were completed while Milne Inlet and Tremblay Sound could not be surveyed due to

97 adverse weather conditions.

98 Infrared technology is a reliable, accurate method to survey ice-associated

99 pinnipeds (Udevitz et al. 2008; Conn et al. 2014; Young et al. In Revision). Therefore,

100 we used a forward-looking infrared camera (FLIR T1030sc) with a 45° lens and a Nikon

101 D810 digital single-lens reflex (DSLR) camera with a 35 mm lens. Thermal infrared

102 imagery (video files) and DSLR photographs were obtained from a strip directly below

103 the aircraft. Photographs were taken at an interval of two seconds, providing overlap

104 between consecutive photos. At the target altitude of 305 m, the strip width covered by

105 both the DSLR camera (resolution of 7360 x 4912 pixels) and the infrared camera

106 (resolution of 1024 x 768 pixels) was 250 m. Observations of ringed seals were

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107 analysed using strip transect methods and ringed seal density was estimated using the

108 standard ratio estimate (Buckland et al. 2001).

109 Infrared video files were analysed by a single observer using FLIR ResearchIR

110 Max software version 4.30.1.70 (FLIR Systems, Inc., Wilsonville, OR, USA) to detect

111 heat signatures indicative of potential seals. All potential seals observed in the infrared

112 imagery were verified by checking the corresponding images obtained from the Nikon

113 DSLR camera. In this verification process, only the DSLR photographs that

114 corresponded to potential seals in the infrared imagery were analysed. We analyzed a

115 random set of 802 photographs to assess the rate of successful detection of the infrared

116 camera where 33 out of 34 (97%) of the observed seals were also detected in the

117 infrared imagery, offering high confidenceDraft in the infrared imagery method (see Young et

118 al. In Revision for more details). GPS coordinates of the observed seals were extracted

119 from the photograph metadata to perform analysis of seal distribution and space use.

120 Data analysis

121 We constructed a spatial distribution map of ringed seals in the study area by summing

122 the number of individuals within 5 km x 5 km grid cells using ArcGIS 10.5 (ESRI Inc.,

123 USA). In 2016, when multiple surveys of each stratum were conducted, we only used

124 observations of ringed seals from the survey that had the highest density of individuals

125 and densities were calculated using the standard ratio estimate (see Young et al. In

126 Revision). Briefly, density estimates were highest in Milne Inlet (1.40 individuals/km2;

127 flown on June 22, 2016) followed by Eclipse Sound (0.98 individuals/km2; flown on June

128 19 and 22, 2016) and Navy Board Inlet (0.74 individuals/km2; flown on June 22, 2016).

129 All data were projected to a Lambert Azimuthal Equal Area projection before analysis.

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130 We then performed a spatial hotspot analysis in the form of Getis-Ord Gi* statistic (Getis

131 and Ord 1992) in ArcGIS to quantify specific areas of high spatial clustering and

132 significance. The Getis-Ord Gi* statistic tests the spatial clustering of grid cell values

133 that are higher (hotspot) or lower (coldspot) than is expected by a random distribution.

134 Significance tests using z-scores were performed between nearby grid cells in a

135 surrounding neighbourhood area (Getis and Ord 1992). A z-score above 1.96 (red) or

136 below -1.96 (blue) is significant at the α = 0.05 level and indicates a hotspot and

137 coldspot for ringed seals, respectively. To conceptualize the spatial relationship, we

138 used the more conservative polygon contiguity conceptualization (i.e. contiguity edges

139 corners in ArcGIS) – a common approach when equal area grid units are used within

140 areal datasets (Harvey et al. 2017). DraftWe calculated the spatial and percentage overlap

141 (km2 and % area, respectively) of ringed seal hotspots between 2016 and 2017 using

142 the Union geoprocessing tool in ArcGIS. The proposed ice-breaking route was obtained

143 from Baffinland and the Nunavut Planning Commission, and georeferenced in ArcGIS

144 from CBC 2017).

145

146 Results

147 The number of ringed seals observed per 5 km x 5 km grid cell ranged from 0 to 14

148 individuals. Observations of ringed seals were more common in Eclipse Sound and

149 Milne Inlet than in Navy Board Inlet (Fig. 2a, c). In 2016, ringed seal hotspots were

150 identified in several areas: 1) eastern Eclipse Sound near the entrance to Baffin Bay, 2)

151 in western Eclipse Sound near the southern point of , 3) northern section of

152 Tremblay Sound, 4) in the middle of Milne Inlet, and 5) in southern Milne Inlet northward

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153 of Koluktoo Bay (Fig. 2b). The proposed ice-breaking route overlapped ringed seal

154 hotspots in four different areas (eastern and western Eclipse Sound, midway in Milne

155 Inlet and in southern Milne Inlet; Fig. 2b). In 2017, ringed seal hotspots occurred within

156 the proposed ice-breaking route in eastern, western, and southwestern Eclipse Sound

157 (Fig. 2c, d). Note that Milne Inlet could not be surveyed in 2017 due to inclement

158 weather. The amount of spatial overlap of ringed seal hotspots in Eclipse Sound and

159 Navy Board Inlet between 2016 and 2017 was low (75 km2 and 12%) due to a smaller

160 hotspot size in 2016 than in 2017 in western Eclipse Sound and due to a shift of 10 km

161 to the east of the eastern Eclipse Sound hotspot in 2017 (Fig 2). However, general

162 locations of ringed seals hotspots in western and eastern Eclipse Sound were

163 cartographically similar inter-annually.Draft

164

165 Discussion and conclusion

166 We identified several ringed seal hotspots in Eclipse Sound and Milne Inlet which

167 proposed ice-breaking would traverse through in spring. An area of particular

168 importance is the ringed seal hotspot in western Eclipse Sound extending southward

169 from Bylot Island, which has been documented as a ringed seal pupping ground through

170 traditional ecological knowledge (Baffinland 2012). Furthermore, hotspots from the 2017

171 aerial survey also encompassed part of this large ringed seal pupping ground further

172 highlighting its importance. Southern Milne Inlet is also a ringed seal hotspot and had

173 relatively high densities of ringed seals in a 2008 aerial survey (2.2 individuals/km2 in

174 Koluktoo Bay and 1.3 individuals/km2 in Milne Inlet; Baffinland 2012). Eastern Eclipse

175 Sound and Tremblay Sound were also ringed seal hotspots which are areas of popular

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176 tourist floe edge camps in spring (i.e. Arctic Adventures and Black Feather) and attract

177 high concentrations of narwhal in summer, respectively (Doniol-Valcroze et al. 2015).

178 Future research to quantify ringed seal hotspots relative to dynamic variables such as

179 sea ice topography and thickness, bathymetry, prey availability and polar bear predation

180 pressure are needed to determine important factors that support preferred ringed seal

181 winter habitat and its inter-annual persistence in particular areas. Identifying areas

182 where ringed seal densities are highest and understanding why they might shift over

183 time would provide important information for management to evaluate anthropogenic

184 stressors and to develop mitigation strategies of potential spring ice-breaking activities

185 on Arctic wildlife in the area.

186 Anthropogenic disturbances causeDraft animal behaviour and physiology responses

187 (i.e. stress and energetic burdens) at the individual level that can propagate up to

188 demographic consequences at the population level (McHuron et al. 2017). For example,

189 high levels of vessel noise can disrupt foraging behaviour and may be associated with

190 chronic stress in cetaceans (Rolland et al. 2012; Wisniewska et al. 2018). Ice-breaking

191 can significantly displace and disturb Caspian seal mother-pup pairs from their resting

192 position within 200m of the vessel (Wilson et al. 2017). A distance of at least 250 m and

193 speeds ≤ 2.2 knots were found to minimize disturbance to mothers and pups in the

194 Caspian Sea (Wilson et al. 2017). However, most Arctic ice-breakers are larger, more

195 powerful and can navigate through thicker ice than Caspian Sea ice-breaking vessels

196 which range from 66-96 m in length (Wilson et al. 2017). Therefore, Arctic ice-breakers

197 would likely create more noise and disturbance, and equate to a much larger ‘safe’

198 distance than 250 m. Separation of ringed seal mother and pups during the critical three

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199 month neonatal and nursing period (March to May) would have detrimental

200 consequences to pup survival by ending the lactation period prematurely. However, a

201 disturbance diameter of 500m along the vessel path would make it difficult to navigate

202 from Eclipse Sound entrance to the port site at the base of Milne Inlet while avoiding

203 ringed seal hotspots, especially if subsequent trips use different paths through the ice to

204 avoid refrozen rubble. For example, in southern Milne Inlet northward of Koluktoo Bay,

205 distances between shorelines range from 7 to 14 km, therefore shipping disturbance on

206 ringed seals is likely unavoidable in this narrow area.

207 In 2017, Baffinland proposed a 189 m long icebreaking cargo vessel, the MV

208 Nunavik, to traverse through Eclipse Sound and Milne Inlet to its port site during late

209 March – a critical time period in ringedDraft seal seasonal life-history (see CBC 2017). As

210 such, this ice-breaking vessel would likely destroy critical ringed seal subnivean birth

211 and nursing habitat, breathing holes, resting lairs and haulout platforms in the shore-fast

212 ice. Subnivean birth lairs maintained by adult females provides pups protection against

213 inclement weather and predation from polar bears (Ursus maritimus) and Arctic foxes

214 (Vulpes vulpes; Smith and Stirling 1975). As observed in Caspian seals, there will also

215 be an incidence of vessel-ringed seal collisions resulting in mortality, to which pups are

216 most susceptible (Wilson et al. 2017). Other pinnipeds, such as Atlantic walrus

217 (Odobenus rosmarus rosmarus) are also susceptible to collisions if ice-breakers

218 traverse through preferred walrus habitat (Stewart et al. 2014). Wilson et al. (2017) also

219 documented that collisions were most frequent at speeds ≥ 4 knots and at night possibly

220 due to individuals being dazzled by ship lights.

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221 To minimize negative impacts of ice-breaking traffic on ringed seals in the

222 Eclipse Sound area, mitigation strategies need to be developed and integrated prior to

223 these activities occurring. As such, having no ice-breaking activities during the ringed

224 seal parturition, nursing and breeding period would greatly reduce negative impacts on

225 the ringed seal population. Such mitigation would require supply logistics to already be

226 in place prior to this critical time period. In addition, the ice-breaking route should avoid

227 ringed seal hotspots and pupping grounds in western eclipse Sound. For example, ice-

228 breaking vessels could be routed through areas with lower densities of ringed seals

229 such as Navy Board Inlet, though ice-breaking would be unavoidable in ringed seal

230 hotspots in southern Milne Inlet. To lower the probability of collisions, restricting speeds

231 could likely allow successful manoeuvrabilityDraft of ships around hauled out seals while

232 also maintaining a ‘safe distance’ of at least 250m. However, more precaution in speed

233 and a farther ‘safe distance’ would likely be needed for larger Arctic ice-breaking

234 vessels. Given rapid resource and industrial development across the Arctic, more

235 studies like this are required to inform management and policy decision-makers who

236 can then develop effective mitigation strategies and cumulative effects assessments in

237 a rapidly developing world.

238

239 Acknowledgements

240 Funding for this research was provided by Polar Continental Shelf Project (PCSP),

241 Environment and Climate Change Canada (ECCC), World Wildlife Fund Canada

242 (WWF), Fisheries and Oceans Canada (DFO) and ArcticNet. We thank our Twin Otter

243 captains, first officers, and engineers from Kenn Borek Air Ltd. We also thank

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244 Mittimatalik Hunters and Trappers Organization of , NU for supporting this

245 research. We thank Jeff Higdon for input and Jarrett Friesen for georeferencing the

246 proposed ice-breaking route.

247

248 Conflict of Interest

249 The authors have conflicts of interest to report.

250

251 References

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323 324 McLaren, I.A. 1958. The biology of the ringed seal (Phoca hispida Schreber) in the 325 eastern Canadian Arctic. Bull. Fish. Res. B. Can. 118: 1-97. 326 327 Pizzolato, L., Howell, S.E.L., Derksen, C., Dawson, J. and Copland, L. 2014. Changing 328 sea ice conditions and marine transportation activity in Canadian Arctic waters 329 between 1990 and 2012. Clim. Change 123: 161-173. Doi.org/10.1007/s10584- 330 013-1038-3 331 332 Post, E., Bhatt, U.S., Bitz, C.M., Brodie, J.F., Fulton, T.L., Hebblewhite, M., Kerby, J., 333 Kutz, S.J., Stirling, I. and Walker, D.A. 2013. Ecological consequences of sea-ice 334 decline. Science 341: 519-524. Dor.org/ 10.1126/science.1235225 335 336 Reeves, R.R., Ewins, P.J., Agbayani, S., Heide-Jørgensen M.P., Kovacs, K.M., 337 Lydersen, C., Suydam, R., Elliott, W., Polet, G., van Dijk, Y. and Blijleven R. 338 2014. Distribution of endemic cetaceans in relation to hydrocarbon development 339 and commercial shipping in a warming Arctic. Mar. Pol. 44: 375-389. 340 doi.org/10.1016/j.marpol.2013.10.005 341 342 Rolland, R.M., Parks, S.E., Hunt, K.E., Castellote, M., Corkeron, P.J., Nowacek, D.P., 343 Wasser, S.K. and Kraus, S.D.Draft 2012. Evidence that ship noise increases stress in 344 right whales. Proc. R. Soc. B 279: 2363-2368. doi.org/10.1098/rspb.2011.2429 345 346 Smith, T.G. 1987. The ringed seal, Phoca hispida, of the Canadian western Arctic. Can. 347 Bull. Fish. Aquat. Sci. 216: 1-81. 348 349 Smith, T.G. and Hammill, M.O. 1981. Ecology of the ringed seal, Phoca hispida, in its 350 fast ice breeding habitat. Can. J. Zool. 59: 966-981. doi.org/10.1139/z81-135 351 352 Smith, T.G. and Stirling, I. 1975. The breeding habitat of the ringed seal (Phoca 353 hispida). The birth lair and associated structures. Can. J. Zool. 53: 1297-1305. 354 doi.org/10.1139/z75-155 355 356 Stewart, D.B., Higdon, J.W. and Stewart, R.E.A. 2014. Development threats and effects 357 pathways of shipping related to non-renewable resource developments on 358 Atlantic walruses (Odobenus rosmarus rosmarus) in Hudson Strait and Foxe 359 Basin. Prepared by Arctic Biological Consultants and Higdon Wildlife Consulting, 360 Winnipeg for Fisheries and Oceans Canada. viii + 57 pp. 361 362 Udevitz, M.S., Burn, D.M. and Webber, M.A. 2008. Estimation of walrus populations on 363 sea ice with infrared imagery and aerial photography. Mar. Mamm. Sci. 24: 57- 364 70. doi.org/10.1111/j.1748-7692.2007.00169.x 365 366 Wilson, S.C., Trukhanova, I., Dmitrieva, L., Dolgova, E., Crawford, I., Baimukanov, M., 367 Baimukanov, T., Ismagambetov, B., Pazylbekov, M., Jüssi, M. and Goodman, 368 S.J. 2017. Assessment of impacts and potential mitigation for icebreaking

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369 vessels transiting pupping areas of an ice-breeding seal. Biol. Cons. 214: 213- 370 222. doi.org/10.1016/j.biocon.2017.05.028 371 372 Wisniewska, D.M., Johnson, M., Teilmann, J., Siebert, U., Galatius, A., Dietz, R. and 373 Madsen, P.T. 2018. High rates of vessel noise disrupt foraging in wild harbour 374 porpoises (Phoecena phoecena). Proc. R. Soc. B 285: 20172314. 375 doi.org/10.1098/rspb.2017.2314 376 377 Young, B.G. Yurkowski, D.J., Dunn, B. and Ferguson, S.H. 2018. Comparing the use of 378 infrared imagery with traditional aerial survey methods to estimate ringed seal 379 density on ice. Wild. Soc. Bull. In Revision. 380 381 Yurkowski, D.J., Semeniuk, C.A.D., Harwood, L.A., Rosing-Asvid, A., Dietz, R., Brown, 382 T.M., Clackett, S., Grgicak-Mannion, A., Fisk, A.T and Ferguson, S.H. 2016. 383 Influence of sea ice phenology on the movement of ringed seals across their 384 latitudinal range. Mar. Ecol. Prog. Ser. 562: 237-250. 385 doi.org/10.3354/meps11950 Draft

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386 Table 1. Key ringed seal behavioural and life-history parameters by month during the 387 ice-covered winter and spring (January to July).

388 Territories Parturition Nursing Mating Basking January x February x March x x x April x x x x May x x x x June x July x 389 References: McLaren (1958); Smith and Stirling (1975); Smith and Hammill (1981); 390 Smith (1987); Hammill et al. (1991); Chambellant et al. (2012)

Draft

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Figure captions

Figure 1. Map of Eclipse Sound, NU highlighting the community of Pond Inlet, the shipping port and proposed shipping/ice-breaking route (red; A). Survey strata and transects flown in the Eclipse Sound area during aerial surveys of ringed seals in June of 2016 (black; B). Map was generated using ArcGIS 10.5 (ESRI Inc., USA). NBI: Navy Board Inlet, MI: Milne Inlet, KB: Koluktoo Bay, TS: Tremblay Sound

Figure 2. The 2016 (A) and 2017 (C) spatial distribution of ringed seals per 5 km x 5 km grid cell in Eclipse Sound, Milne Inlet, Tremblay Sound and Navy Board Inlet. Associated 2016 (B) and 2017 (D) ringed seal hotspots are provided per 5 km x 5 km grid cell across the study area (B). Yellow and red signify an insignificant and significant grid cell, respectively. Black line represents the proposed shipping/ice-breaking route in (B) and (D). Milne Inlet transect was not surveyed in 2017 due to inclement weather (see Materials and methods). Map was generated using ArcGIS 10.5 (ESRI Inc., USA).

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