How has climate change impacted marine food-webs in the past, and how might we predict changes in the future?

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How has climate change impacted marine food-webs in the past, and how might we predict changes in the future? John K. Pinnegar, Georg Engelhard, Julia Blanchard, Joe Scutt-Phillips, William Cheung International Symposium: Climate Change Effects on Fish & Fisheries, Sendai, Japan. Session A2 Monday 26th April, 2010

Programme 1. The North Sea ecosystem 2. Changes in fish distribution (across Europe) 3. Climate vs fishing vs habitat modification 4. Consequences for fisheries 5. Consequences for food-webs 6. Predicting the future 7. Some conclusions

The North Sea Ecosystem: Water Depth (m) ICES Stock-Assessment Areas E6 E7 E8 E9 F0 F1 F2 F3 F4 F5 F6 F7 F8 62 61 52 51 50 Latitude 60 59 58 57 56 55 54 53 52 51-4 -3-2 -1 0 1 2 3 4 5 6 7 8 9 Longitude 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 280 270 260 250 240 230 220 210 200 190 180 170 160 150 140 130 120 110 100 90 80 70 60 50 40 30 20 10 0 The North Sea is a semi-enclosed basin with a depth ranging from 30m 200m The ecosystem is dominated by soft-bottom habitats (sand, mud, gravel) The north Sea harbours a wide range of fish stocks exploited mainly by: France, Germany, Belgium, Netherlands, Norway, England, Scotland, Denmark.

Changes in fish distribution (across Europe) Thermal performance Plaice Flounder Cod Seabass From Freitas et al. (2007) Temperature is one of the primary factors, together with food availability and suitable spawning grounds that determine the largescale distribution patterns of fish. Because most fish species prefer a specific temperature range, an expansion or contraction of the distribution range often coincides with long-term changes in temperature and/or climate. The recent warming trend in the northeast Atlantic has coincided with an apparent northward shift in the distribution of fish species from southerly latitudes

Perry et al. (2005) demonstrated that distributions of both exploited and non-exploited North Sea fishes have changed markedly over the last 25 years. Centres of distribution generally shifted by distances ranging from 48 to 403 km during the period 1977 2001. These authors concluded that further temperature rises are likely to have a profound impact on commercial fisheries. A = cod B = monkfish C = snake blenny [see Science 308: 1912-1915]

Fish have also moved to deep waters.. Deeper Shallower pogge wolffish scaldfish solenette grey gurnard cod witch long rough dab megrim dab angler haddock hake whiting lemon sole ling plaice saithe cuckoo ray starry ray four-bearded rockling lesser spotted dogfish sole spurdog lesser weaver Norway pout bib poor cod -15-10 -5 0 5 10 Decadal change in depth anomaly (m per decade) [see Dulvy et al. 2008 ] [See presentation B2-6054] From 1980-2004, the North Sea fish assemblage deepened by ~3.6 m per decade The deepening response was more dramatic in comparison with the latitudinal response that had previously been reported Coldwater species, like megrim and anglerfish, are deepening fastest with warm-water species (e.g. sole and bib) shallowing over time.

Distribution shifts: climate versus fishing? There is much controversy around distribution shifts of North sea fishes over past 3 decades: Climate change hypothesis : warming climate causes warm-adapted species to expand northward, and/or cold-adapted species to contract at south-end of range Fishing pressure hypothesis : fishing pressure has been consistently higher in the southern compared to northern North Sea, causing higher mortality in the south and hence, an apparent distribution shift Other possible drivers include eutrophication, habitat modification

So far studies on North Sea fish distribution shifts have been based on survey data limited to most recent 3 decades: Here, 9 decades of cod, haddock, sole and plaice distribution data were analysed 1913-2008: Statistical Charts with cod, haddock, sole & plaice cpue by rectangle, for British steam and motor otter trawlers Analyses of Scottish and English commercial catch data spanning the period 1913-2007, by Engelhard et al. (Cefas, Lowestoft) has revealed that the peak catches of target species such as cod, haddock, plaice and sole, have all shifted

Cod distribution seems to have shifted steadily north-eastward, towards deeper water Haddock catches have moved very little in terms of centre of distribution, but their southern boundary has shifted northwards by approximately 130 km Plaice 1920s 1950s 1930s 1960s Late 1940s 1970s Plaice have moved offshore towards the central North Sea (i.e. north-westwards) 1980s 1990s 2000s

Changes in the centre of gravity of plaice distribution North South Latitudinal Longitudinal Depth West East Deeper Since WWII, near-continuous northward shift (and eastward up to 1990s), depth shift mimics N-S depth gradient Shallower

1920s 1930s Late 1940s 1950s 1960s 1970s 1980s 1990s 2000s 1920s-2000s: sole distribution (cpue normalised by year) 1920s: very inshore distribution in SW 1930s 1960s: shift/expansion more offshore and more NE (esp. German Bight) 1980s 2000s: contraction away from NE and again more inshore, but more limited to SW

Changes in the centre of gravity of sole distribution Latitudinal Longitudinal Depth North South West East Pre-1980s, shifts in CoG of sole appear linked to warmer and cooler climate regimes Cold 1910s 1920s: sole limited to (shallow) SW, then during warm 1930s 1950s expansion N- and E, then during cold 1960s 1970s contraction to shallower SW Deeper Shallower [ Also see presentation A2-6080, ter Hofstede & Rijnsdorp]

There have been big changes in sea surface temperature (SST) Note the generally close correlation between time series & the overall warming trend during the 20 th Century Marsdiep SST Dover SST 14 13 Central England Air Torungen SST HADISST North Sea Helgoland SST Warm period in 1930s Cool period in the early 1970s Dramatic warming after 1988/89 12 ºC 11 10 9 8 7 1890 1910 1930 1950 1970 1990 2010

However there have also been big changes fishing pressure.. Fishing mortality rates have been higher in the southern North Sea than in the north (Heath et al., 2003, Heath et al., 2007). Apparent changes in distribution (as indicated by Perry et al. 2005) could simply be a consequence of local patterns of fishing pressure and different rates of depletion in spatially segregated sub-stocks. (MEFEPO)

.. and significant modification of habitats. e.g. Maasvlakte is part of the harbour and industrial area of Rotterdam, the Netherlands. It was created in the 1960s by reclaiming land from the North Sea. Maasvlakte 2 in the next few years will cover 1000 hectares. There has been considerable public concern that this development will impact the transport and retention of flatfish and herring larvae. Erftemeijer et al (2009) attempted real-time modelling of hydrodynamic forcing (with wind, air pressure and river discharge) and larval behaviour to establish whether this development is likely to have an effect. In this case the impact will be negligible [See presentation A2-6258]

What will distribution shifts mean for fisheries? Populations may move away from (or towards) the area where fishing fleets operate. Distribution changes may have significant consequences for the distance that must be travelled by fishing boats to reach the target resources with implications for fuel usage and time at sea. Also species distributions may migrate across the boundaries where quotas belong to different nations. Species may move outside the boundaries of marine protected areas / fishery closure areas Incoming species may be commercially exploitable and therefore offer new opportunities for fisheries.

Species may move outside MPA boundaries The Plaice Box was first established in 1989 to reduce discards of juvenile plaice (i.e. to protect nursery grounds). Recent surveys in the Wadden Sea have shown that 1-group plaice is almost absent from the area where it once was very abundant. The Plaice Box is now much less effective as a management measure in comparison with the situation 10 or 15 years ago. MPA boundaries may need to be adaptive in the future. Latitude E6 E7 E8 E9 F0 F1 F2 F3 F4 F5 F6 F7 F8 62 52 51 61 50 49 60 48 47 59 46 45 58 44 43 57 42 41 56 40 39 55 38 37 54 36 35 53 34 33 52 32 31 51-4 -3-2 -1 0 1 2 3 4 5 6 7 8 9 Longitude -Closure Area - Plaice Box - 12 Mile Limit - 200m Depth [see van Keeken et al., 2007; and presentation A2-6258]

The current debate about North Atlantic Mackerel In 2009 mackerel appeared to have moved away from the Norwegian Sector, resulting in disagreements over permissible catches by Norwegian boats in EU waters. Norwegian vessels were forcibly evicted from Scottish waters by UK fishery patrol vessels once they had caught their allotted quota Both Iceland and the Faroe Islands unilaterally claimed additional quota, because mackerel had moved into their waters Such disagreements may become more common place in the future 9 th October, 2009

7000 6000 5000 4000 3000 2000 1000 Incoming species & new fisheries.. 0 (a) 1973 1975 1977 1979 1981 1983 1985 1987 1989 1991 1993 1995 1997 1999 2001 2003 2005 Portugal Netherlands France Spain UK Other 7000 6000 5000 4000 3000 2000 1000 0 (b) 1973 1976 1979 1982 1985 1988 1991 1994 1997 2000 2003 Azores Biscay West of Scotland International fishery landings of red mullet Mullus surmuletus between 1973 and 2005. Portuguese coast Celtic Seas North Sea European fishermen have witnessed and responded to a number of new opportunities in recent years, as warm-water species have moved further North and/or their exploitation has become commercially viable for the first time. Notable examples include new and/or expanding fisheries for seabass, red mullet, john dory, anchovy and squid in the Channel and southern North Sea

Anchovy in the Channel 31 st October 2009

Will fish become harder to catch? Perhaps yes! It is known that gear geometry and hence catchability of certain fish species can be greatly influenced by water depth (Godø and Engås 1989). Given that Dulvy et al. (2008) found that the whole North Sea fish assemblage has deepened by ~3.6 m per decade since 1981, we might anticipate that traditional target fish will become more difficult to catch.

Consequences for food-webs? Considerable effort has been dedicated to the digitization of fish stomach content data spanning the period 1884-2010 to look for major changes in food-webs over the past 100 years The DAPSTOM database (www.cefas.co.uk/dapstom) now contains 175,000 records for 135 fish species and is searchable online. Using this data it has been possible to compare stomach contents of fish (of similar size) in the Dogger Bank region in 1902-1909, with those in 1950-1959 and 2004-2010 Have North Sea food webs changed????

Sandeels represent a greater proportion of the diet now compared to 100 years ago Plaice Cod Mobile prey (e.g. crabs, hermit crabs) are now more important prey items Bivalves (in particular Solen spp. and Mactra spp.) were more important in the past Callaway et al. (2007) demonstrated that crabs have dramatically increased in abundance since 1902, whereas many slow-growing bivalves have declined. Haddock Dab Were these changes driven by fishing pressure, habitat modification or climate????

Predicting the future. Modelling strategies for predicting the potential impacts of climate change on the distribution of species have often focused on the characterization of a species' bioclimate envelope. Original (static) cod distribution In other words, by looking at the current range of temperatures tolerated by a species, it is possible to predict future distribution, if we know how the physical environment in an area will likely change in the future. A world-wide analysis has been carried out (Cheung et al. 2009) using this technique, based on 1066 commercial fish and invertebrate species. [See presentation P1-D1-6105] Relative abundance 0 0 > > > > > > > > > Low High Cod distribution after 50 years (Climate projection from NOAA/GFDL CM 2.1)

Predicting how food-webs might look in the future. Ecopath with Ecosim (EwE) has emerged as one of the most popular and widely applied ecosystem modelling approaches in the marine environment. Seals Megrim Cod Macrocarnivores Polychaete - feeders Echinod- Plaice feeders Sole Molluscfeeders Polychaetes Molluscs Echinoderms Whales & Dolphins Whiting Hake Horse Monk Mackerel Piscivores Haddock Mackerel Blue Cephalopods Whiting Planktivores Larvae Macro- Smallplanktocrustacean feeders Krill & Mysids Crabs, anomurans Shrimps & lobsters Herbivorous- Zooplankton Hydroids, Small-benthic sponges, Micro- crustaceans bryozoans etc. Zooplankton Meio-fauna Pelagic- Micro-benthos Bacteria At it s core Ecopath is essentially a food-web model, and includes all fluxes between biological components of the system, from detritus and bacteria up to whales. Macro-Algae Detritus Phytoplankton Ecosim is a time-dynamic version of Ecopath and can be used simulate the long-term or historic impacts of different fishing practices. Ecosim can be tuned to fit real time-series data or forced using assumptions about fishing and long-term (and seasonal) climate or plankton productivity.

The impact of climate change and fishing on food-webs in Lake Victoria.. Approaches: Ecosystem modelling Top down pressure Bottom up climate forcing What data are available to us? [See poster P1-D1-6199]

The impact of climate change and fishing on food-webs in Lake Victoria.. Results vary dramatically depending on the demographic (human population) and climate scenario selected Outcomes were chiefly dependant on future fishing pressure in Lake Victoria High fishing effort results in return of Haplochromine sp. biomass Greater understanding of the relationship between primary production and climate is needed urgently

Conclusions & future plans.. 1. There have been major changes in the distribution of North Sea fish species and the prey that they consume 2. It is very difficult to separate the influence of long-term climate change from the effects of fishing and habitat modification 3. Fisheries in the North Sea are already being affected by changes in fish distribution (for better or worse) 4. Techniques are being developed to predict what North Sea fish communities might look like in the future and how food-webs might change

Volcanic Ash cloud - 1530GMT 15 April Acknowledgements EU-RECLAIM, EU-INCOFISH, UK Defra-MF1202, MF1109