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Antarctic ( superba)

Stephen Nicol and So Kawaguchi

General information

Antarctic Krill (Euphausia superba) are one of the larger of krill (Euphausiids – free swimming -like ) reaching a maximum length of 60 mm and a weight of 2 g when mature. They live exclusively in the Southern and have a very wide distribution over a range of habitats. They are a swarming (or schooling species) and much of the krill is to be found in large, dense aggregations that can extend for tens of kilometers. Krill can be found in the surface layer, in mid water and near the ocean floor, and can undertake daily vertical migrations. have a complicated life history, changing size, shape and habitat as they grow (Nicol 2006). They mature at two years old and can live for up to 11 years. Adult krill are capable of living anywhere in the – from the very surface layer to the seafloor, and from inshore areas to the deep open ocean. Larval and juvenile krill are associated with sea ice and feed on that grow on the underside. There is a lack of clarity on whether there are distinct populations of krill around the Antarctic or whether there is regular interchange between centers of distribution. The krill population is large and their production rate is high consequently they are preyed upon by a range of vertebrates including seals, and as well as by invertebrates such as .

Range

The range of krill is estimated to be 19 x 106 km2 (Atkinson et al. 2009). See figure 1.

THE IUCN RED LIST OF THREATENED SPECIES™

Figure 1. Map generated from the KRILLBASE data base* derived from a historical compilation of scientific net hauls from 1926 onwards. Green dots indicate the presence of Antarctic Krill in the catch, red dots indicate absence. The blue area encompasses the range of krill: for a given longitude, the extent is taken as the northernmost latitude of any krill observation in a 30-degree sector centred on that given longitude. The three major frontal systems in the Southern Ocean are indicated. *Krill Base data (version March 2014) has been provided with by the major data contributors to the database (A. Atkinson, S. Hill, V. Siegel, E. Pakhomov, C. Reiss, S. Kawaguchi).

Krill abundance

The overall mean abundance of krill estimated from scientific net surveys conducted between1926-2004 was estimated to be 379 million tons (Atkinson et al. 2009). Atkinson et al. (2009) estimated the krill biomass for January-February 2000 to be 133 million tons. This estimate was based on an acoustic estimate of biomass derived from the CCAMLR2000 survey of the South Atlantic of 37.3 million tons which was estimated to be 28% of the global krill biomass. CCAMLR has subsequently revised this biomass estimate (SC-CCAMLR 2010 para. 3.29) to 60.3 million tons. Applying the process used

in Atkinson et al. (2009) to this new estimate of biomass results in a global krill biomass of 215 million tons in the year 2000.

Generation length

The generation length of Antarctic krill has been calculated to be 5 years using the standard IUCN methodology and the following inputs: • Survival up to 12 month=0.03: Based on average number of juveniles survived from in Nagoya (Hirano et al. 2003). • Survival from 1st to 2nd year = 0.1: Based on Brinton and Townsend (1984). • Survival from 2nd year onwards = 0.4: Based on widely accepted krill post-larval mortality rate (M=0.8) which equals to survival rate of 0.4. • Life span = lives until end of its 6th year (Siegel 1987). • Annual fecundity, 12,000 for age 3+ onwards based on Tarling et al. (2007). Annual fecundity of 6,000 was used for age 2+ krill (spawning during 3rd their summer) due to their smaller body size.

Trends in krill biomass.

Although a decline in krill density in the 1970s and 1980s has been reported from analysis of net survey data in the Southwest Atlantic sector (Atkinson et al. 2004) the magnitude of any such decline is difficult to ascertain. For the purposes of this current assessment the trend in krill biomass over the last 15 years (3generation times) has been used. Two time series of acoustic data (the standard method of biomass assessment used by CCAMLR) exist within the range of Antarctic krill – one for South Georgia (Fielding et al. 2014) and one for the Elephant Island area Cossio et al. 2011). Both of these time series are from the South Atlantic region where 28% of the krill biomass is thought to reside (Atkinson et al. 2009). There is considerable inter-annual (and intra-annual) fluctuation in krill density (either measured acoustically or using nets) at a fixed location; however, in neither of these two acoustic time series is there a significant trend in the data.

Figure 2. (a) Frequency distribution of log10(krill density+1) for each annual mid- season survey. The median (white circles), mean (black crosses), and inter-quartile range (black bar) is shown. (b) The mean WCB krill density from 1997 to 2013 following Jolly and Hampton (1990).

Figure 3. Acoustic estimates based on updated methods from ASAM 2010 of Antarctic Krill biomass (tons) between 1996 and 2011 for the AMLR study area during the January-February (A) and February-March (B) legs. Biomass for each area is calculated if the area was sampled. In several recent years no second leg has been conducted.

The krill :

The has been operating for more than over 40 years. Catches peaked in the early 1980s with Japanese and Soviet vessels catching over half a million tons a year (see below).

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Figure 4. Catches of krill in the Southern Ocean from 1973 to 2014 (data from the CCAMLR Statistical Bulletin).

Today, about 300,000 tons are caught from the South West Atlantic, largely by Norwegian vessels, producing high-end feed and supplements for human consumption (Foster et al., 2011). When the krill fishery was established there was concern that it might cause irreversible damage to the Antarctic , so a unique international treaty was signed to ensure it would be managed using an approach that took into account the needs of the entire ecosystem. This treaty was the Convention on the Conservation of Antarctic Marine Living Resources (CCAMLR). The management of the krill fishery has been guided by its principles since the early 1980s. The fishery is regulated through a series of measures that specify how much can be caught, where it can be caught, acceptable levels of by-catch and other operational restrictions. The amount of krill that can be caught in any one year is set through “Precautionary Catch Limits (PCL)”, which is more conservative than normal fishery quotas because of the sensitivity of the Antarctic krill biomass, the that depend on it as a food source, and the unique environment in which they live (Nicol and Foster 2003). Catch limits are calculated for a particular area by working out how much krill is in that area and by determining the proportion of the krill stock in that area that can be harvested without irreversibly impacting the ecosystem in the long-term. Currently, only 9.3% of the krill

stock is available to the fishery. Precautionary Catch Limits have been set for several large areas of the Southern Ocean, totaling more than 8 million tons per year. In the South Atlantic, where the krill fishery currently operates, there is a Precautionary Catch Limit of 5.6 million tons. As an added element of precaution, CCAMLR has applied a “Trigger Level” of 620,000 tons throughout the main grounds – a level of catch that cannot be exceeded until more advanced management procedures are in place.

Climate change and krill.

The life history and population dynamics of Antarctic krill are likely to be impacted by the due to increasing levels of CO2 in the atmosphere (Flores et al. 2012), and may result in a reduced habitat range (Hill et al. 2013). The reproductive output and recruitment success of krill has been related to the extent, timing and duration of winter sea ice cover. The underside structure of sea ice provides a nursery ground for overwintering krill larvae and a substrate for algae which are their food. Extensive winter sea ice promotes strong spring bloom when retreating in spring which fuels the adults’ reproductive output for their summer spawning season (Quetin and Ross 2001). Krill growth has also been observed to decrease above a temperature optimum of 0.5°C (Atkinson et al. 2006). Their early developmental stage of krill is vulnerable to increased levels of CO2 projected within their habitat range in year 2100 and beyond (Kawaguchi et al. 2013). Overall, the cumulative impact of climate change is most likely to be negative.

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(Euphausia superba) density at South Georgia, Southern Ocean: 1997–2013. ICES Journal of Marine : Journal du Conseil. 71(9): 2578-2588. Flores, H., Atkinson, A., Kawaguchi, S., Krafft, B.A., Milinevsky, G., Nicol, S., Reiss, C., Tarling, G.A., Werner, R., Bravo Rebolledo, E.B., Cirelli, V., Cuzin-Roudy, J., Fielding, S., Groeneveld, J.J., Haraldsson, M., Lombana, A., Marschoff, E., Meyer, B., Pakhomov, E.A., Rombola, E., Schmidt, K., Siegel, V., Teschke, M., Tonkes, H., Toullec, J.Y., Trathan, P.N., Tremblay, N., Van de Putte, A.P., van Franeker, J.A. and Werner, T. (2012) Impact of climate change on Antarctic krill. Marine Ecology Progress Series. 458:1-19. Foster, J., Nicol, S. and Kawaguchi, S. (2011) The use of patent databases to detect trends in the krill fishery. CCAMLR Science. 18: 135-144. Hill, S.L., Phillips, T. and Atkinson, A. (2013) Potential climate change effects on the habitat of Antarctic krill in the Weddell Quadrant of the Southern Ocean. PLoS One. 8(8): e72246 doi:10.1371/journal.pone.0072246 Hirano, Y., Matsuda, T. and Kawaguchi, S. (2003). Breeding Antarctic krill in captivity. Marine and Freshwater Behaviour and Physiology. 36(4): 259-269. Kawaguchi, S., Ishida, A., King, R., Raymond, B., Waller, N., Constable, A., Nicol, S., Wakita, M. and Ishimatsu, A. (2013) Risk maps for Antarctic krill under projected Southern . Nature Climate Change. 3: 843-847. Nicol, S. (2006) Krill, currents and sea ice; the life cycle of Euphausia superba in relation to its changing environment. Bioscience. 56(2): 111-120. Nicol, S. and Foster, J. (2003). Recent trends in the fishery for Antarctic krill. Aquatic Living Resources. 16(1): 42-45. Nicol, S., Foster, J. and Kawaguchi, S. (2011). The fishery for Antarctic krill – recent developments. and doi: 10.1111/j.1467-2979.2011.00406.x Quetin, L.B. and Ross, R.M. (2001) Environmental variability and its impact on the reproductive cycle of Antarctic krill. American Zoologist. 41: 74-2001. Siegel, V. (1987). Age and growth of Antarctic Euphausiacea (Crustacea) under natural conditions. Marine Biology. 96(4): 483-495. Tarling, G.A., Cuzin-Roudy, J., Thorpe, S.E., Shreeve, R.S., Ward, P. and Murphy, E.J. (2007). Recruitment of Antarctic krill Euphausia superba in the South Georgia region: adult fecundity and the fate of larvae. Marine Ecology Progress Series. 331: 161-179.