Recovery of Holothuroidea Population Density, Community Composition, and Respiration Activity After a Deep-Sea Disturbance Experiment
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Heriot-Watt University Research Gateway Recovery of Holothuroidea population density, community composition, and respiration activity after a deep-sea disturbance experiment Citation for published version: Stratmann, T, Voorsmit, I, Gebruk, A, Brown, A, Purser, A, Marcon, Y, Sweetman, AK, Jones, DOB & van Oevelen, D 2018, 'Recovery of Holothuroidea population density, community composition, and respiration activity after a deep-sea disturbance experiment', Limnology and Oceanography, vol. 63, no. 5, pp. 2140- 2153. https://doi.org/10.1002/lno.10929 Digital Object Identifier (DOI): 10.1002/lno.10929 Link: Link to publication record in Heriot-Watt Research Portal Document Version: Peer reviewed version Published In: Limnology and Oceanography Publisher Rights Statement: Copyright © and Moral Rights for this work are retained by the author and/or other copyright owners. A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. 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Oct. 2021 1 Title: Recovery of Holothuroidea population density, community composition and 2 respiration activity after a deep-sea disturbance experiment 3 Authors and affiliations: Tanja Stratmann1*, Ilja Voorsmit2, Andrey Gebruk3, Alastair Brown4, 4 Autun Purser5, Yann Marcon5,6, Andrew K. Sweetman7, Daniel O. B. Jones8, Dick van Oevelen1 5 1NIOZ Royal Netherlands Institute for Sea Research, Department of Estuarine and Delta 6 Systems, and Utrecht University, P.O. Box 140, 4400 AC Yerseke, The Netherlands. 7 2Faculty of Mathematics and Natural Sciences, University of Groningen, P.O. Box 11103, 9700 8 CC Groningen, The Netherlands. 9 3P.P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Nakhimovsky Pr., 36, 10 Moscow 117997, Russia. 11 4Ocean and Earth Science, University of Southampton, National Oceanography Centre 12 Southampton, European Way, Southampton, SO14 3ZH, United Kingdom. 13 5Deep Sea Ecology and Technology, Alfred Wegener Institute, Am Handelshafen 12, 27570 14 Bremerhaven, Germany. 15 6MARUM, Center for Marine Environmental Sciences, Leobener Str., 28359 Bremen, 16 Germany. 17 7The Marine Benthic Ecology, Biogeochemistry and In-situ Technology Research Group, The 18 Lyell Centre for Earth and Marine Science and Technology, Heriot-Watt University, Edinburgh 19 EH14 4AS, United Kingdom. 20 8Ocean Biogeochemistry and Ecosystems, National Oceanography Centre Southampton, 21 European Way, Southampton SO14 3ZH, United Kingdom. 22 *Corresponding author 1 23 e-mail addresses: T Stratmann: [email protected], Ilja Voorsmit: 24 [email protected], Andrey Gebruk: [email protected], Alastair Brown: 25 [email protected], Autun Purser: [email protected], Yann Marcon: 26 ymar•[email protected], Andrew K. Sweetman: [email protected], Daniel O. B. Jones: 27 [email protected], Dick van Oevelen: [email protected] 28 29 Running head: Holothuroidea recovery after disturbance 2 30 1. Abstract 31 Mining polymetallic nodules on abyssal plains will have adverse impacts on deep-sea 32 ecosystems, but it is largely unknown whether the impacted ecosystem will recover, and if so at 33 what rate. In 1989 the ‘DISturbance and reCOLonization’ (DISCOL) experiment was conducted 34 in the Peru Basin where the seafloor was disturbed with a plough harrow construction to explore 35 the effect of small-scale sediment disturbance from deep-sea mining. Densities of Holothuroidea 36 in the region were last investigated seven years post-disturbance, before nineteen years later, the 37 DISCOL site was re-visited in 2015. An ‘ocean floor observatory system’ was used to 38 photograph the seabed across ploughed and unploughed seafloor and at reference sites. The 39 images were analyzed to determine the Holothuroidea population density and community 40 composition, which were combined with in situ respiration measurements of individual 41 Holothuroidea to generate a respiration budget of the study area. For the first time since the 42 experimental disturbance, similar Holothuroidea densities were observed at the DISCOL site and 43 at reference sites. The Holothuroidea assemblage was dominated by Amperima sp., Mesothuria 44 sp. and Benthodytes typica, together contributing 46% to the Holothuroidea population density. 45 Biomass and respiration were similar among sites, with a Holothuroidea community respiration 46 of 5.84×10-4±8.74×10-5 mmol C m-2 d-1 at reference sites. Although these results indicate 47 recovery of Holothuroidea, extrapolations regarding recovery from deep-sea mining activities 48 must be made with caution: results presented here are based on a relatively small-scale 49 disturbance experiment as compared to industrial-scale nodule mining, and also only represent 50 one taxonomic class of the megafauna. 51 52 2. Key words 3 53 Anthropogenic disturbance, manganese nodule, echinoderm, holothurian, OFOS, south-east 54 Pacific, community composition, DISCOL 55 56 3. Introduction 57 Interest in mining polymetallic nodules from abyssal plains has increased substantially since the 58 1960s (Glasby 2000; Jones et al. 2017). Polymetallic nodules contain valuable metals such as 59 copper, cobalt, nickel and rare earth elements (Wang and Müller 2009) and are therefore 60 considered economically interesting resources. However, deep-sea mining activities will have 61 negative impacts on these vulnerable deep-sea ecosystems through the removal of hard 62 substratum (i.e. polymetallic nodules) essential for sessile (e.g. Amon et al. 2016; Vanreusel et 63 al. 2016) and mobile (Leitner et al. 2016; Purser et al. 2016) fauna. Further negative impacts may 64 result from the disturbance and/or removal of the upper sediment layer (Thiel and Tiefsee- 65 Umweltschutz 2001), i.e. the habitat that contains the majority of organisms and their food 66 sources (Danovaro et al. 1993; Danovaro et al. 1995; Haeckel et al. 2001). Sessile fauna and 67 infauna will be removed in the mining tracks (Bluhm 2001; Borowski and Thiel 1998; Borowski 68 2001), and fauna in and on adjacent sediments may be blanketed by mechanically displaced 69 sediment (Thiel and Tiefsee-Umweltschutz 2001). Re-suspended sediment and sediment plumes 70 may smother both sessile and mobile fauna over extended areas, or potentially adversely impact 71 filter feeding organisms (Brooke et al. 2009; Jankowski and Zielke 2001; Thiel and Tiefsee- 72 Umweltschutz 2001). 73 74 Ecological concerns about deep-sea mining have resulted in several deep-sea experiments 75 studying ecosystem recovery following disturbance (Jones et al. 2017). To investigate the 4 76 recovery from small scale disturbance experiments employing isolated individual disturbance 77 tracks, Vanreusel et al. (2016) used ROV video surveys to compare sessile and mobile metazoan 78 epifauna densities at disturbed and undisturbed sites with high and low “polymetallic nodule” 79 coverage in the Clarion-Clipperton Zone (CCZ, central Pacific). The mobile metazoan epifauna 80 in a 20-year-old experimental mining track at a nodule-free site comprised mainly Holothuroidea 81 and Ophiuroidea (1,500 ind. ha-1), whereas the density at the reference site was substantially 82 higher (3,000 ind. ha-1). In contrast, the same study showed that mobile metazoan epifauna at a 83 nodule-rich site in another, 37-year-old mining track in the CCZ consisted exclusively of 84 Echinoidea (1,000 ind. ha-1) and had a 70% lower mobile epifauna density than the reference 85 site, where Holothuroidea and Ophiuroidea were also present. 86 87 The most extensively studied and largest-scale disturbance experiment conducted to date is the 88 ‘DISturbance and reCOLonization experiment’ (DISCOL) in the Peru Basin. This experiment 89 was conducted in a 10.8 km2 circular area, DISCOL Disturbance Area or ‘DEA’ in short, which 90 was ploughed 78 times on diametric courses in 1989 to mimic a small-scale deep-sea mining 91 disturbance event (Bluhm 2001; Foell et al. 1990; Foell et al. 1992). The DEA can be considered 92 ‘small-scale’ in comparison to commercial mining as the 10.8 km2 represent between 1.4 and 93 3.6% of the area projected to be disturbed by one individual mining operation per year (Smith et 94 al. 2008). The imposed disturbance occurred on two levels: 1) direct impacts inside ‘plough 95 tracks‘ where polymetallic nodules were ploughed into the sediment (~22% of DEA) and 2) 96 indirect impacts from plume exposure associated with the ploughing outside plough tracks (~70- 97 75% of the DEA; Bluhm and Thiel 1996; Thiel and Schriever 1989). Four sites, each 5 98 approximately 4 km away from the DEA served as reference areas