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Larval and Adult Fish Assemblages Along the Northwest Passage: the Shallow Kitikmeot and the Ice-Covered Parry Channel As Potential Barriers to Dispersal

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Larval and Adult Fish Assemblages Along the Northwest Passage: the Shallow Kitikmeot and the Ice-Covered Parry Channel As Potential Barriers to Dispersal Arctic Science Larval and adult fish assemblages along the Northwest Passage: the shallow Kitikmeot and the ice-covered Parry Channel as potential barriers to dispersal Journal: Arctic Science Manuscript ID AS-2018-0003.R1 Manuscript Type: Article Date Submitted by the 15-Jun-2018 Author: Complete List of Authors: Bouchard, Caroline; Université Laval, Québec-Océan; Gronlands Naturinstitut, Greenland Climate Research Centre Geoffroy, Maxime; Memorial University of Newfoundland Fisheries and Marine Institute LeBlanc, Mathieu; Université Laval, Québec-Océan Fortier, Louis;Draft Université Laval, Québec-Océan Arctic cod, polar cod, fish larvae, Canadian Arctic Archipelago, spatial Keyword: connectivity Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue?: https://mc06.manuscriptcentral.com/asopen-pubs Page 1 of 22 Arctic Science 1 Larval and adult fish assemblages along the Northwest Passage: the shallow Kitikmeot 2 and the ice-covered Parry Channel as potential barriers to dispersal 3 4 Caroline Bouchard, Maxime Geoffroy, Mathieu LeBlanc, and Louis Fortier 5 6 C. Bouchard*, M. LeBlanc, and L. Fortier. Québec-Océan, Département de Biologie, 7 Université Laval, Québec, QC G1V 0A6, Canada. ([email protected]; 8 [email protected]; [email protected]) 9 M. Geoffroy: Centre for Fisheries Ecosytems Research, Marine Institute of Memorial 10 University of Newfoundland, St. John’s, NL A1C 5R3, Canada. 11 ([email protected]) 12 Corresponding author: C. BouchardDraft (email: [email protected]). *Current address: Greenland 13 Institute of Natural Resources, Greenland Climate Research 14 Centre, 3900 Nuuk, Greenland. https://mc06.manuscriptcentral.com/asopen-pubs Arctic Science Page 2 of 22 15 Abstract 16 Climate warming and sea ice decline are expected to increase fish population movements in 17 the circumpolar Arctic, including across the Canadian Arctic Archipelago (CAA). Wanting 18 information on present distribution, habitat uses, barriers to dispersal, and population 19 connectivity along the Northwest Passage (NWP) hinders the management of fish in the 20 North American Arctic. Pelagic trawl, bottom trawl and ichthyoplankton net collections from 21 the US Beaufort Sea to Baffin Bay between 2005 and 2017 are used to map fish distribution 22 along the NWP and identify potential zoogeographic barriers. In the Kitikmeot (southern 23 CAA), the combination of shallow depths, sub-zero temperatures and slow water circulation 24 may represent a physical barrier reducing the dispersal of marine fish between western and 25 eastern regions. In contrast, the Parry Channel (northern CAA) may exemplify a disappearing 26 sea ice barrier as climate warming unfolds and allow new genetic exchanges. 27 28 29 Keywords: Arctic cod, polar cod, fishDraft larvae, Canadian Arctic Archipelago, spatial 30 connectivity 31 https://mc06.manuscriptcentral.com/asopen-pubs Page 3 of 22 Arctic Science 32 Introduction 33 In the ocean, identifying barriers to dispersal and patterns of population connectivity is often 34 challenging. Yet, understanding how populations are connected is crucial to manage fisheries 35 resources and to predict the response of marine ecosystems to climate change and other 36 anthropogenic disruptions (e.g. Gerber et al., 2014, Carvalho and Hauser, 1994). Habitat 37 fragmentation is one of the main types of physical barriers, and can create population 38 discontinuities even on short geographical distances (e.g. Blanco Gonzalez et al., 2016). In 39 Arctic seas, sea ice acts as a barrier to the dispersal of many marine mammal species, and a 40 diminishing ice cover results in increased hybridization between subspecies and closely 41 related species (Kelly et al., 2010). 42 43 Warmer temperatures and reduced ice cover, by favoring the northward range expansion of 44 boreal fish species, are expected to increase the interchange of fish populations between the 45 Atlantic and the Pacific via the Northwest and Northeast Passages (Wisz et al., 2015). The 46 Canadian Arctic Archipelago (CAA) Draftconnects the Pacific-influenced waters of the Beaufort 47 Sea with the Atlantic-influenced waters of Baffin Bay (Fig. 1a) and will be at the forefront of 48 this interchange. However, our knowledge of fish population structure, habitat use and larval 49 ecology for the region is limited (Fortier et al., 2015). For instance, despite its dominance of 50 the pelagic fish assemblage (Benoit et al., 2008) and its pivotal role in arctic marine 51 ecosystems (Welch et al., 1992), the different habitat uses of Arctic cod (Boreogadus saida) 52 and their relative importance remain debated. Immature Arctic cod colonize anfractuosities in 53 the ice pack (e.g. David et al., 2015) and sometimes form dense surface schools in shallow 54 embayments (e.g. Welch et al., 1992, Matley et al., 2013, Drost et al., 2014). In the North 55 American Arctic, mature individuals concentrate over the continental slope in the relatively 56 warm Atlantic waters at depth > 200 m (e.g. Benoit et al., 2008, Parker-Stetter et al., 2011, 57 Geoffroy et al., 2011). 58 59 Shallow regions of the Kitikmeot (e.g. Queen Maud Gulf, max. depth ca. 100 m, Fig. 1a), 60 may represent a suboptimal habitat for mature Arctic cod and some barrier to dispersal and 61 gene fluxes across the CAA. Dispersal could be possible further north through the deeper 62 western Parry Channel (Fig. 1a). Yet, extreme environmental conditions in the central and 63 northern Archipelago, including a frequent ice cover, low SSTs and absence of freshwater 64 input may shorten the hatching season and limit the early survival of Arctic cod (Bouchard https://mc06.manuscriptcentral.com/asopen-pubs Arctic Science Page 4 of 22 65 and Fortier, 2008, Bouchard and Fortier, 2011, Bouchard et al., 2017). The genetic population 66 structure of Arctic cod, showing a phylogenetical break between the Beaufort Sea and Baffin 67 Bay (R.J. Nelson, Fisheries and Oceans Canada, unpublished data), also suggest the presence 68 of barriers to dispersal in the CAA. Several benthic arctic fish of the families Stichaeidae, 69 Cottidae, Liparidae, Agonidae and Ammoditidae, typical of shallow waters (Fortier et al., 70 2015), are less mobile as adults than the pelagic Arctic cod and depend on their planktonic 71 larvae for dispersal. The harsh environment of the central and northern CAA may also reduce 72 connectivity among populations of these species. 73 74 In the present study, we document larval and adult fish assemblages along the Northwest 75 Passage and explore possible barriers to dispersal of fish populations between western and 76 eastern regions. In particular, we test the hypothesis that the Kitikmeot represents a 77 zoogeographical barrier for Arctic cod, a key species in arctic marine ecosystems. 78 79 Materials and methods Draft 80 Study areas 81 The Northwest Passage links the Beaufort Sea to Baffin Bay and is composed of two main 82 waterways: (1) the Parry Channel which runs from M’Clure Strait to Lancaster Sound in a 83 relatively straight line, and (2) the Kitikmeot formed of complex coastlines and 84 heterogeneous bathymetry from the Amundsen Gulf to Lancaster Sound, via M’Clintock 85 Channel or Peel Sound (Fig. 1a). Both waterways are relatively contrasted in terms of fish 86 habitats. The Parry Channel is deep (max. of ~700 m), generally ice covered and receive very 87 little coastal freshwater. The Kitikmeot is shallower (< 100 m in the southern waterways of 88 Dease Strait and Queen Maud Gulf and < 350 m in the northern waterways of Larsen Sound, 89 M’Clintock Channel and Peel Sound), has longer ice-free seasons and receives freshwater 90 from numerous rivers. For the analyses, the general study area was divided into four areas: 91 the Western Arctic, the Parry Channel, the Kitikmeot and the Eastern Arctic, each of them 92 composed of several geographical regions (Fig. 1a). 93 94 Oceanographic data 95 Data collected with a SBE-911 plus® conductivity, temperature and depth probe (CTD) 96 deployed from the research icebreaker CCGS Amundsen at 11 stations along the Northwest 97 Passage from the Amundsen Gulf to Baffin Bay in July-August 2015 were used to illustrate https://mc06.manuscriptcentral.com/asopen-pubs Page 5 of 22 Arctic Science 98 the water masses in the sudy area. Atlantic Water masses were distinguished with the 0°C 99 isotherm at depths > 200 m (Carmack and Macdonald 2002). 100 101 Ichthyoplankton sampling 102 Ichthyoplankton samples were collected between 2005 and 2017 from the Amundsen at 103 different stations over the study area (Fig. 1b). A Double Square Net (DSN) carrying two 1- 104 m2 aperture nets (mesh size 500 µm and 750 µm) was deployed obliquely at maximum 105 sampling depths of ca. 90 m (or 10 m above the bottom at shallower stations) and at a ship 106 speed of 2 knots. The duration of the DSN tow averaged 14.6 ± 4.6 minutes (mean ± standard 107 deviation) and varied between 4.3 to 36.7 minutes. Larval and juvenile fish were sorted out 108 and individually preserved in 95% ethanol. Back in the laboratory, all individuals were 109 identified to the family. Catch per unit effort (CPUE in fish min-1) were calculated by 110 dividing the number of larval and juvenile fish from each family by the DSN deployment 111 duration. To account for the intense mortality occurring in larval fish populations during their 112 first few months in the plankton, ichthyoplanktonDraft abundances were estimated separately for 113 collection dates in July-August and September-October. The CPUE values were then 114 averaged for each region. 115 116 Adult fish sampling 117 Pelagic fish were sampled with an Isaacs-Kidd Midwater Trawl (IKMT, rectangular 9-m2 118 mouth aperture net with mesh size of 11 mm in the first section and 5 mm in the last section) 119 deployed between June and October from 2014 to 2017 (Fig 1c).
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