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Size Metrics, Longevity, and Growth Rates in Umbellula Encrinus (Cnidaria: Pennatulacea) from the Eastern Canadian Arctic

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Size Metrics, Longevity, and Growth Rates in Umbellula Encrinus (Cnidaria: Pennatulacea) from the Eastern Canadian Arctic Arctic Science Size metrics, longevity, and growth rates in Umbellula encrinus (Cnidaria: Pennatulacea) from the Eastern Canadian Arctic Journal: Arctic Science Manuscript ID AS-2018-0009.R1 Manuscript Type: Article Date Submitted by the Author: 19-Jun-2018 Complete List of Authors: Neves, Bárbara; Memorial University of Newfoundland Department of Biology Edinger, Evan;Draft Memorial University, Geography Wareham Hayes, Vonda; Fisheries and Oceans Canada Newfoundland and Labrador Region Devine, Brynn; Memorial University of Newfoundland Fisheries and Marine Institute Wheeland, Laura; Memorial University of Newfoundland Fisheries and Marine Institute Layne, Graham; Memorial University of Newfoundland, Earth Sciences Deep-water corals, Size distribution, Growth rings, Radiocarbon analysis, Keyword: Trace element analysis Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue?: https://mc06.manuscriptcentral.com/asopen-pubs Page 1 of 63 Arctic Science 1 Size metrics, longevity, and growth rates in Umbellula encrinus (Cnidaria: 2 Pennatulacea) from the Eastern Canadian Arctic. 3 4 Bárbara de Moura Neves1*, Evan Edinger1,2,5, Vonda Wareham Hayes3, Brynn Devine1,4, 5 Laura Wheeland4, Graham Layne5. 6 1Department of Biology, Memorial University of Newfoundland. A1B 3X9, St. John’s, NL 7 Canada. 8 2Department of Geography, Memorial University of Newfoundland. A1B 3X9, St. John’s, 9 NL Canada. 10 3Northwest Atlantic Fisheries Center,Draft Fisheries and Oceans Canada. A1C 5X1, St. John’s, 11 Canada. 12 4Marine Institute, Memorial University of Newfoundland. A1C 5R3, St. John’s, NL 13 Canada. 14 5Department of Earth Sciences, Memorial University of Newfoundland. A1B 3X9, St. 15 John’s, NL Canada. *Corresponding author. E-mail: [email protected]. Present address: Northwest Atlantic Fisheries Center, Fisheries and Oceans Canada. A1C 5X1, St. John’s, NL Canada. 1 https://mc06.manuscriptcentral.com/asopen-pubs Arctic Science Page 2 of 63 16 Abstract 17 Umbellula encrinus Linnaeus, 1758 is a deep-water sea pen commonly found in the Eastern 18 Canadian Arctic. It can reach heights of >2 m, and it has often been caught as fishing 19 bycatch. Here we characterized abundance/density, size metrics, longevity, and growth rates 20 of U. encrinus colonies from Baffin Bay (between Greenland and Canada). No prevalent 21 size classes were identified at most locations, except for Jones Sound and Cape Dyer, 22 where small-size colonies dominated. Average number of growth rings in the internal 23 skeleton (axis) of the examined colonies ranged between 2-68, with a maximum of 75. A 24 bomb-14C analysis yielded 14C curves comparable to those of other deep-water octocorals 25 with annual ring formation. A trace element analysis of Mg/Ca, Sr/Ca, Ba/Ca, and Na/Ca 26 yielded values oscillating along the axisDraft radius, with the number of peaks and growth rings 27 being comparable. Growth rates averaged 0.067 ± 0.015 mm yr-1 (radial extension) and 4.5 28 ± 1.2 cm yr-1 (linear extension), considering rings to be formed annually. Relationships 29 between radial growth rates and depth and surface salinity were weak, but statistically 30 significant. Umbellula encrinus is a long-lived species, vulnerable to various types of 31 fishing gear, with a skeleton that stores biological and environmental information. 32 33 Key words: Deep-water corals, Size distribution, Growth rings, Radiocarbon analysis, 34 Trace element analysis. 2 https://mc06.manuscriptcentral.com/asopen-pubs Page 3 of 63 Arctic Science 35 Introduction 36 Umbellula encrinus Linnaeus, 1758 is a sea pen species of boreal/polar distribution 37 (Broch 1958). Colonies of this sea pen have a proximal peduncle that anchors the colony in 38 soft substrates, followed by an elongated and thin stalk, and a distal crown of polyps 39 (Williams 1995). Colonies can reach heights >2 m (Jørgensen et al. 2015; Neves et al. 40 2018, this study), supported by an internal calcified skeleton (axis) where growth rings can 41 be identified (Ellis 1753; Neves et al. 2018, this study). 42 In the Northwest (NW) Atlantic, U. encrinus is the sea pen species of northernmost 43 distribution, having been recorded as far as northern Baffin Bay (between Greenland and 44 the Canadian Archipelago) (Broch 1958). It has also been recorded in Newfoundland 45 (Wareham and Edinger 2007; WarehamDraft 2009), in the Mid-Atlantic ridge (Molodtsova et al. 46 2008; Mortensen et al. 2008), in Norwegian waters (Buhl-Mortensen et al. 2012; Buhl- 47 Mortensen and Buhl-Mortensen 2014), and in the Barents Sea, where it is considered 48 highly vulnerable to bottom trawling (Jørgensen et al. 2015). 49 In certain areas of Baffin Bay, U. encrinus is believed to be present in high 50 densities. Dense occurrences of Umbellula lindahli Kölliker, 1875 were reported in the 51 Umanak Fjord (West Greenland) as a result of bottom trawls taking place during the Danish 52 Ingolf Expedition (Mortensen 1912). It is commonly caught as fishing bycatch from 53 trawling in both East (Jørgensen et al. 2013) and West Baffin Bay (Wareham 2009; 54 Kenchington et al. 2011). In fact, the first known report of Umbellula sp. dates from 1753, 55 based on samples collected near Greenland (74° N, ~430 m) aboard the Britannia, which 56 was on a whale fishery at that time (Ellis 1753). During recent exploratory fishery surveys 3 https://mc06.manuscriptcentral.com/asopen-pubs Arctic Science Page 4 of 63 57 conducted in Jones Sound and Qikiqtarjuaq (West Baffin Bay) several colonies of U. 58 encrinus were caught as longline, whelk pot and shrimp trap bycatch (this study), indicating 59 that colonies might be abundant in this region. 60 The conspicuous presence of U. encrinus in soft bottom communities in Baffin Bay 61 added to their vulnerability to different types of fishing gear in an increasingly fished area 62 (e.g. Christiansen et al. 2014), raise concerns regarding U. encrinus’ susceptibility to 63 damage/removal. Their exceptional attainable large sizes and the presence of growth rings 64 in their skeleton suggest the long-lived nature of this sea pen (Neves et al. 2018). 65 Furthermore, the possibility of being found in high densities highlights their potential as 66 vulnerable marine ecosystems (VMEs) and habitat for other species (Mortensen 1912; 67 Baillon et al. 2012; De Clippele et al.Draft 2015). 68 In this study our main objectives were to describe abundance/density and size 69 structure patterns, to estimate longevity and growth rates, and to investigate relationships 70 between growth rates and environmental variables for U. encrinus colonies from the 71 Eastern Canadian Arctic. Additionally, for the first time we performed bomb-14C and trace 72 element analyzes in the skeleton of U. encrinus colonies aiming to identify patterns 73 indicative of growth ring formation periodicity. 74 75 Material and methods 76 Sampling 77 The study area comprises eight sites in Baffin Bay: Northeast (NE) Baffin Bay, 78 Jones Sound, Lancaster Sound, Scott Inlet, Home Bay, Qikiqtarjuaq, Cape Dyer, and 4 https://mc06.manuscriptcentral.com/asopen-pubs Page 5 of 63 Arctic Science 79 Cumberland Sound (Fig. 1, Table 1). One sample from Iceland was included in the 14C 80 analysis only. Colonies were obtained using variable gear including bottom trawls, 81 longlines, shrimp traps and whelk pots (Tables 1 and 2). At some sites, more than one gear 82 type were used, and colonies incomplete in size were not included in the size structure 83 portion of the study (Table 1). Samples have been kept frozen at -20 °C since collection. 84 Video data were obtained at Lancaster Sound, Scott Inlet, Home Bay, Qikiqtarjuaq, 85 and Cape Dyer. A Super-Mohawk (SuMo) ROV was deployed in July 2014, in October 86 2015, and in August 2017 aboard the CCGS Amundsen. The SuMo ROV has a high- 87 definition camera (1Cam Alpha, Sub C Imaging, 24.1 megapixels) and it projects a pair of 88 lasers 6 cm apart for size estimation. Colonies were opportunistically video-recorded along 89 transects 0.94-2.7 km long, at depths Draftranging 475-750 m (Table 2). Camera average field of 90 view (width) was 3 m. 91 92 Abundance and size-frequency distribution 93 Relative abundance was estimated by dividing the number of colonies caught or 94 observed by surveyed distance (based on ship speed and survey duration). Density was 95 calculated as the number of colonies m-2 (area determined based on surveyed distance, 96 opening width of trawls, gangion lines, and video field of view). Estimated abundance and 97 density were only determined for samples of known total catch (trawls and longline, Table 1). 98 1). 99 A total of 430 measurements of colony height were included in the size-frequency 100 distribution study (Table 1, Table 4). For this study, colonies were measured in their whole 5 https://mc06.manuscriptcentral.com/asopen-pubs Arctic Science Page 6 of 63 101 length, from the bottom of peduncle to the tip of the polyparium (cluster of polyps) (Fig. 2), 102 except in colonies where the polyparium was missing. Peduncle and rachis (stalk plus 103 polyparium) size were also individually determined. A total of 95 colonies were measured 104 for colony height versus peduncle height relationships, but only 85 were included in the 105 statistical analysis, because ten samples had no information on depth (used as a covariate). 106 For the video analysis, colonies were only measured when the scaling lasers touched the 107 colony or were touching the bottom near the colony. Therefore, some colonies seen in the 108 video could not be measured but were included in our abundance and density estimates. 109 Since the peduncle of sea pens remains buried in the sediment, the peduncle could 110 not be visualized in the videos.
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