Cod Behaves in Mysterious Ways:Shifting Distribution in the North Sea During the Last Century. ICES CM 2011/D:06
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International Council for the Exploration of the Sea ICES CM 2011/D:06 Theme Session D Linking the History to the Present: Understanding the History of Fish, Fisheries and Management Cod behaves in mysterious ways: shifting distribution in the North Sea during the last century Georg H. Engelhard*, David A. Righton, Tina K. Kerby, and John K. Pinnegar Centre for Environment, Fisheries and Aquaculture Science, Pakefield Road, Lowestoft NR33 0HT, UK * Author for correspondence ([email protected]) Summary The distribution of cod within the North Sea has shown major shifts over the course of the last century. This has become evident from an analysis of almost one-hundred years of British commercial fisheries data, digitised from Cefas archives. Combined with contemporary fisheries data, these span the period 1913–2010 (excepting both World Wars), at the spatially detailed level of ICES rectangle (0.5° Latitude, 1° Longitude). New analysis of old data reveals that during most of the Twentieth Century, North Sea cod distribution was very different from that in the most recent period (2000– 2010). Whereas historically, densities of cod catches were especially concentrated in the central and north-western North Sea, they are presently concentrated in the northern- and north-easternmost part of the North Sea. We attempt to reveal the extent to which climate, fishing, or both have contributed to the recent shift in distribution. Keywords: cod, distribution shift, climate, North Sea 1 INTRODUCTION Historically, cod Gadus morhua has arguably been the most important of commercial fish species, and its fishery has played a prominent role in the economical and political development of Europe as well as North America over the past centuries (e.g. Kurlansky 1997). More recently, disputes over cod fishing grounds have led to such well-known conflicts as the British-Icelandic ‘Cod Wars’ of 1950s– 1970s (Gilchrist 1978; Jónsson 1982), which ultimately contributed to the international establishment of 200-miles Exclusive Economic Zones. Currently, cod remains one of the world’s most important fish stocks, and in 2004 ranked 12th among all fish species in terms of global landings (and even higher in terms of value of landings; FAO 2007). This is in spite of significant declines in landings of most cod stocks during recent decades (review: Engelhard et al. 2010a) and a collapse of populations off eastern Canada and the U.S. (Kurlansky 1997). The North Sea cod stock has shown a particularly marked decline since the 1980s, even more noteworthy since it followed a period of very high productivity known as the ‘gadoid outburst’ (late 1960s to mid-1980s: Daan 1978; Hislop 1996). During the gadoid outburst, not only cod but also haddock Melanogrammus aeglefinus, whiting Merlangius merlangus, saithe Pollachius virens and Norway pout Trisopterus esmarki produced some of the largest year-classes on record (Holden 1978; Jones & Hislop 1978). Whilst landings of cod had remained fairly stable throughout the period 1900– 1950, they rose enormously during the 1960s and remained very high until the mid-1980s, when the recent marked decline began. Survey indices and stock assessment have indicated that the high landings were not only due to substantially increased fishing effort but also to a considerable extent, to very high recruitment and biomass levels (Pope & Macer 1996). Unfortunately, the recruitment of North Sea cod has remained low since the late 1990s, and the spawning stock biomass has been at historically low levels for almost two decades now (ICES 2010). The past three decades have also seen a shift in the mean distribution of cod within the North Sea, to more northerly and on average deeper waters; this was reported by studies based on research survey data (International Bottom Trawl Surveys, IBTS; see Hedger et al. 2004; Perry et al. 2005; Rindorf & Lewy 2006; Dulvy et al. 2008). As this shift coincided with the population decline it is important to understand its causes, not only because of predicted links between distribution and abundance (e.g., Blanchard et al. 2005) but also because changes in fish distributions may have knock-on effects upon fisheries (Pinnegar et al. 2010). Two main hypotheses have been put forward (Engelhard et al. 2010b). Firstly, climate change is expected to result in contractions, expansions, or shifts in fish distribution (review: Rijnsdorp et al. 2009). In the North Sea, a warming trend has happened over the past 30 years; this has coincided with a northward shift in many, but not all, North Sea fish species (Beare et al. 2004; Perry et al. 2005), and an average, deepening shift of around –3.6 m per decade since the 1980s (Dulvy et al. 2008). Secondly, fishing pressure has over the same period been consistently higher in the southern compared to northern North Sea (Jennings et al. 1999); with a greater rate of fishery-induced depletion in the south, a northward shift in mean population distribution is to be expected. However, the rather short time-span of fishery-independent survey data (past three decades) makes it very hard to disentangle these two hypotheses (which are not mutually exclusive). This has motivated the current study based on a unique dataset of British commercial catch per unit effort 2 (cpue) records spanning a far longer timespan—approaching one century—covering both warming and cooling periods, and including periods of contrasting levels of fishing effort. The aims of the present paper are to: 1. Describe long-term distribution shifts of cod within the North Sea over the period 1913–2010; 2. Compare the current (2000–2010) distribution with that throughout the Twentieth Century; 3. Examine to what extent distribution shifts can be explained by variables related to climate change, fishing pressure, and/or stock abundance. METHODS Data and modelling of cod distributions In broad lines the methodology follows that of our companion paper describing the long-term distribution shifts of sole Solea solea and plaice Pleuronectes platessa in the North Sea (Engelhard et al. 2011). For the period 1913–1980, cod data were obtained from historical fisheries ‘statistical charts’ (catalogued in Engelhard 2005) that were produced by the UK Ministry of Agriculture, Fisheries and Food (MAFF; now the UK Department for Environment, Food and Rural Affairs [Defra]). These show fishing effort (hours fished) and fish landings by British otter trawlers (either steam- or motor-driven) for each ICES rectangle (0.5° Latitude, by 1° Longitude) in the North Sea. These data record all fish that were landed by the otter trawl fleet into England and Wales (1913, 1968–1980) or into England, Scotland and Wales (1920–1967). For the period 1968–2010, data on otter trawler landings into Scotland were obtained from the Fisheries Management Database (FMD) of Marine Scotland (cf. Greenstreet et al. 1999). For 1982–2010, data on otter trawl landings into England and Wales were obtained from the Fisheries Activity Database (FAD) of Defra/Cefas. Over the time-span examined, important improvements have occurred in the cod fishing power, or technical efficiency, of otter trawlers (Robinson 2000; Engelhard 2008). Our aim is not to analyse temporal changes in absolute cpue values as proxies for the abundance of cod, but rather to look at trends in spatial distribution of catches. We therefore normalised the cpue values in any given year (divided by the annual mean cpue for the entire area), to overcome the confusing effect of an increase in fishing power of trawlers. We are assuming that relative cpue by the commercial fleet gives an appropriate indication of the spatial distribution of the species. We acknowledge that potential bias might arise from uneven spatial distribution of effort by more or less powerful vessels within the North Sea. By decade, we calculated cpue values by rectangle for a large area encompassing the majority of the North Sea (shaded in Figure 2). As an approach to quantify shifts in population distribution, we calculated the ‘centres of gravity’ of the latitudinal, longitudinal, and depth distributions of North Sea cod, using methods akin to those developed and applied by Heino et al. (2008). This analysis was based on a slightly less extensive area within the North Sea that included only those rectangles with cpue data for the majority of years in our time-series (see polygon line in Figure 2). Within this polygon, the latitudinal (or longitudinal) centre of gravity of distribution in a given year was calculated as the average of the 3 latitudes (or longitudes) of all rectangle centres, weighted by the cpue value in each rectangle. Weighted standard deviations and standard errors of the weighted mean longitudes were calculated (Bevington 1969). The centres of gravity of depth distributions were calculated analogously and, given that cod generally live close to the seafloor, based on the mean sea depth in any given rectangle. Modelling distribution shifts in relation to climate, abundance and fishing pressure We examined cod distribution in relation to: (1) climatic variables and (2) cod fishing pressure and abundance. As a broad-scale climate indicator, the North Atlantic Oscillation (NAO) winter index (December of the previous year to March of the focal year) for 1913–2007 was taken from Jones et al. (1997), with updated values provided online by the Climatic Research Unit, University of East Anglia, Norwich, UK (www.cru.uea.ac.uk/~timo/projpages/nao_update.htm; see Figure 1a). The NAO is associated with speed and direction of westerly winds across the North Atlantic, and is particularly important in winter when it exerts a strong influence on European weather patterns, and on Atlantic water inflow into the North Sea; a positive NAO is generally linked with strong wind circulation and higher atmospheric and sea temperatures in western Europe (Hurrell 1995; Jones et al.