Biomass of Antarctic Krill in the Scotia Sea in January/February 2000 and Its Use in Revising an Estimate of Precautionary Yield
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ARTICLE IN PRESS Deep-Sea Research II 51 (2004) 1215–1236 www.elsevier.com/locate/dsr2 Biomass of Antarctic krill in the Scotia Sea in January/February 2000 and its use in revising an estimate of precautionary yield Roger P. Hewitta,Ã, Jon Watkinsb, Mikio Naganobuc, Viacheslav Sushind, Andrew S. Brierleyb,1, David Demera, Svetlana Kasatkinad, Yoshimi Takaoe, Cathy Gossb, Alexander Malyshkod, Mark Brandonf, So Kawaguchic, Volker Siegelg, Philip Trathanb, Jennifer Emerya, Inigo Eversonb, Denzil Millerh aSouthwest Fisheries Science Center, 8604 La Jolla Shores Drive, La Jolla, CA 92037, USA bBritish Antarctic Survey, NERC, High Cross, Madingley Road, Cambridge CB3 0ET, UK cNational Research Institute of Far Seas Fisheries, Orido 5-7-1, Shimizu, Shizuoka 424, Japan dAtlantNIRO, 5 Dimitry Donskoy Street, Kaliningrad 236000, Russia eNational Research Institute of Fisheries Engineering, Ebidai Hasaki, Kashima-gun, Ibaraki 314-0421, Japan fEarth Sciences, The Open University, Milton Keynes MK7-6AA, UK gInstitut fu¨r Seefischerei, Palmaille 9, D-22767 Hamburg, Germany hMarine and Coastal Management, Private Bag X2, Roggebaai 8012, South Africa Accepted 18 June 2004 Available online 23 September 2004 Abstract In January and February 2000, a collaborative survey designed to assess the biomass of Antarctic krill across the Scotia Sea was conducted aboard research vessels from Japan, Russia, the UK and the USA using active acoustic and net sampling. Survey design, sampling protocols, and data analysis procedures are described. Mean krill density across the survey area was estimated to be 21.4 g mÀ2, and total biomass was estimated to be 44.3 million tonnes (CV 11.4%). This biomass estimate leads to a revised estimate of precautionary yield for krill in the Scotia Sea of 4 million tonnes. However, before the fishery can be permitted to expand to this level, it will be necessary to establish mechanisms to avoid concentration of fishing effort, particularly near colonies of land-breeding krill predators, and to consider the effects of krill immigrating into the region from multiple sources. Published by Elsevier Ltd. ÃCorresponding author. E-mail address: [email protected] (R.P. Hewitt). 1Present address: Getty Marine Laboratory, University of St. Andrews, St. Andrews, Fife, Scotland. 0967-0645/$ - see front matter Published by Elsevier Ltd. doi:10.1016/j.dsr2.2004.06.011 ARTICLE IN PRESS 1216 R.P. Hewitt et al. / Deep-Sea Research II 51 (2004) 1215–1236 1. Introduction one year should be less than 10%. The second criterion is to protect the viability of krill predator The highest concentrations of Antarctic krill populations; this is defined such that the median (Euphausia superba), krill predators, and krill- population level should be at least 75% of the fishing effort in the Southern Ocean are located in unexploited median population level. The third the Scotia Sea (Agnew and Nicol, 1996; Laws, criterion is to evaluate these risks using population 1985; Marr, 1962). The internationalfishery is trajectories that extend at least 20 years. The value regulated in accordance with the Convention for of the harvest rate, expressed as a proportion of the Conservation of Antarctic Marine Living the unexploited biomass (g), that meets these Resources, which is part of the Antarctic Treaty criteria is accepted as the most precautionary. system. In principle, the Commission for the This value, together with an estimate of B0; is used Conservation of Antarctic Marine Living Re- to set the precautionary yield for krill. sources (CCAMLR) has adopted a feedback Two important parameters in this analysis are approach to management of the krill fishery, by B0 and its associated variance. These values are which management measures are adjusted in used to generate a distribution of population response to ecosystem monitoring (Constable et biomasses from which an initialbiomass is drawn al., 2000; Hewitt and Linen Low, 2000). However, for each population trajectory. Initially, an esti- such a scheme remains to be fully developed. In mate of krill biomass was generated from acoustic the interim, a complementary approach, which data collected during the first international BIO- defines and implements provisions of Article II2 of MASS experiment (FIBEX)3 in 1981 (Trathan et the Convention in reference to the Scotia Sea krill al., 1992), the only large-scale acoustic survey in stock, was adopted in order to set a precautionary this region prior to 2000. Because exploitation had catch limit (Butterworth et al., 1991, 1994). historically been low relative to the size of the The approach is to set the proportion (g)of fished resource an estimate of the standing stock unexploited population biomass (B0) that can be was assumed to approximate B0: Recent reports of harvested under defined management criteria. The the Scientific Committee of CCAMLR have allowable catch, referred to as the precautionary questioned the current relevance of this estimate yield (Y), is thus defined as (e.g. CCAMLR, 1995, Annex 4, para 4.61) and have recommended a new survey. Y ¼ gB : (1) 0 The reasons for conducting a new survey were The risks of exceeding the criteria are evaluated recognition that: (1) severaltechnicalimprove- by comparing statisticaldistributions of popula- ments had been made in the assessment of krill tion biomasses generated from simulated popula- biomass using active acoustic methods since the tion trajectories, with and without fishing FIBEX survey (Everson et al., 1990; Greene et al., mortality. Uncertainty is accommodated by using 1991; Hewitt and Demer, 1991); (2) the FIBEX values of abundance, recruitment, growth, and survey area was substantially less than the known mortality drawn from appropriate statistical dis- habitat of krill in the Scotia Sea (CCAMLR, tributions. The first criterion is to protect the 1995); and (3) the krill population in the Scotia Sea viability of the harvested population and for may not be stable. Recently, published evidence Antarctic krill is defined such that the probability suggests that krill reproductive success may be that the spawning biomass declines to less than dependent on multi-year changes in the physical 20% of its unexploited median level during any environment (Brierley et al., 1999; Loeb et al., 2Article II of the Convention mandates that fisheries be 3In the early 1980s, the Scientific Committee for Antarctic managed such that: (a) the size of harvested populations is Research (SCAR) established the BIOMASS Program (Biolo- sufficient to ensure stable recruitment; (b) ecological relation- gicalInvestigations of Antarctic Systems and Stocks). FIBEX ships between harvested and dependent populations are was a multi-national, multi-ship effort to conduct large-scale maintained; and (c) changes to the marine system that cannot acoustic surveys over large areas of the Southern Ocean. See be reversed over two to three decades are prevented. also El-Sayed (1994). ARTICLE IN PRESS R.P. Hewitt et al. / Deep-Sea Research II 51 (2004) 1215–1236 1217 1997; Naganobu et al., 1999; Nicolet al.,2000 ; CCAMLR subsequently adopted a revised pre- White and Peterson, 1996). During periods of cautionary catch limit for krill (CCAMLR, 2000b; northward excursions of the Southern Boundary Hewitt et al., 2002). Much of the information of the Antarctic Circumpolar Current (SBACC), presented here is drawn from these reports. the development of winter sea ice is more extensive, populations of Salpa thompsoni (a pelagic tunicate postulated to be a competitor with krill for access to the spring phytoplankton 2. Survey design and data collection protocols bloom) are displaced offshore, and both krill reproductive output and survivalof their larvae The defining physicalfeature of the Scotia Sea is are enhanced. During periods of southward its southern boundary along the Scotia Ridge, excursion of the SBACC, the development of extending from the South Shetland Islands east winter sea ice is less extensive, salps are more and north through the South Orkney Islands, the abundant closer to shore, and krill reproductive South Sandwich Islands, and South Georgia (Fig. success is depressed. These interactions may be 1). This ridge influences the direction and intensity confounded by a warming trend observed in the of the ACC. Antarctic krill appear to move region of the Antarctic Peninsula over the last 50 eastward through the Scotia Sea via the ACC, years (Vaughan and Doake, 1996). The intention although the relative importance of passive trans- was to anchor the estimate of precautionary yield port versus active migration is uncertain. Likely with the most recent and accurate assessment of sources of immigrants to the Scotia Sea are the Antarctic krill in the Scotia Sea possible. Because Bellingshausen Sea to the west and the Weddell harvest rates continue to be low relative to the size Sea to the south. Differences in mitochondrial of the fished resource, it was again assumed that DNA sequences suggest that krill from these an estimate of the current standing stock was regions may be genetically distinct (Zane et al., 1998). Within the Scotia Sea, zones of water equivalent to B0: Plans for the survey developed over a period of convergence, eddies, and gyres are loci for krill 5 years through a series of working papers, concentrations (Makarov et al., 1988; Witek et al., discussions at the meetings of the Scientific 1988). Krill spawn in the vicinity of the South Committee of CCAMLR and its working groups, Shetland and South Orkney Islands. Although and more formalworkshops ( CCAMLR, 1995, they are abundant further to the north and east Annex 4, paras 4.62–4.67; CCAMLR, 1996, near South Georgia, they do not spawn there in Annex 4, paras 3.72–3.75; CCAMLR, 1997, great numbers and few larvae are found (Fraser, Annex 4, paras 8.121–8.129; CCAMLR, 1998, 1936; Siegel, 2000). Consumption of krill through- Annex 4, paras 9.49–9.90; CCAMLR, 1999, out the Scotia Sea by baleen whales, crabeater and Annex 4, paras 8.1–8.74 and Appendix D).