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Introductory comments - Global change in marine ecosystems: Patterns, processesand interactions with regional and local scale impacts

1. Background to volume

The world is changing rapidly, ultimately due to the pressure ofhuman population growth driving change at global, regional and localscales. There is convincing and widely accepted evidence that theclimate is changing as a result of anthropogenic forcing due togreenhouse gas emissions (Mitchell et al., 1995; Lee et al., 2006; IPCC,2007). Increased concentrations of one greenhouse gas — carbondioxide is causing a reduction in the pH of the oceans (Caldeira andWickett;, 2003; The Royal Society, 2005; Doney et al., 2009). Globaltrade is also leading to homogenization of floras and faunas as speciesare deliberately and/or accidentally transported around the world(Mack et al., 2000; Kolar and Lodge, 2002; Ruiz and Carlton, 2003;Drake and Lodge, 2004; Rahel, 2007).

Overfishing is occurringglobally for large pelagic species at the topoffood webs (Jackson et al., 2001; Myers and Worm, 2003; Worm andMyers, 2003; Heithaus et al., 2008; Baum and Worm, 2009), and at aregional scale for benthic species in most shallow seas (Solan et al.,2004; Steneck, 2006; Kaiser et al., 2007; Genner et al., 2010). Pollution,whilst being more regulated, especially from point sources, is allpervasive (Thompson et al., 2002) and plastic litter is a global problem(Fig. 1; Thompson et al., 2004; Thompson et al., 2009). Despitepreventative efforts, oil spills can still be a major threat to the marineenvironment (Southward and Southward, 1978; Gundlach et al., 1983;Hawkins and Southward, 1992),with the 2010 sinking of theDeepwaterHorizon oilrig in the Gulf of Mexico representing one of the worst oildisasters in United States history. The coasts are becoming increasinglydeveloped leading tohabitat loss at local and regional scales (Lotze et al.,2006; Airoldi and Beck, 2007). Responses to rising and stormier seaswillinevitably lead to intervention in coastal processes as people andinfrastructure will need to be protected, leading to even more coastalhabitat modification and loss (Fig. 2; Airoldi et al., 2005; Martin et al.,2005; Moschella et al., 2005; Bulleri and Chapman, 2010; Chapman andUnderwood, 2011-this issue).

2. Detecting change

Marine and coastal ecosystems also tend tofluctuate naturally, due toboth deterministic processes (e.g., ocean circulation and ENSO: Nicolet al., 2000; McPhaden et al., 2006) and stochastic events (e.g.,hurricanes: Paerl et al., 2006; Hughes et al., 2009). It is often extremelydifficult to distinguish human-induced change from natural spatial andtemporal variation (Underwood, 1991; Underwood, 1992; Underwood,1994; Hughes et al., 2009)— unless there are long-term and broad-scaledata available (Southward, 1980; Southward et al., 1995, 2005; Hawkins

et al., 2003;Mieszkowska et al., 2005; Poloczanska et al., 2011;Wethey etal., 2011-this issue). This is particularly the case with fluctuations inresponse to climate change where broad-scale low amplitude, longwave-length signals have to be distinguished from local and regionalhigher frequency noise (Genner et al., 2004; Harley and Connell, 2009).Furthermore, there can be regional scale differences in response toclimate change (McGinty et al., 2011; Philippart et al., 2011).

There is also the perennial challenge of distinguishing betweenweather and climate (see Helmuth et al., 2006; Firth et al., 2011-thisissue) — especially recently in northern Europe where the winters of2009/10 and 2010/11 have been the coldest for 30 years (Fig. 3; seeWethey et al., 2011-this issue). These were not as extreme, however,as in 1962/63 (Crisp, 1964) which was one of the coldest on record(Wethey et al., 2011-this issue).

3. Linking levels of organisation

A further challenge is linking responses at different levels ofbiological organisation. Metabolism and individual performance istemperature dependent in most marine organisms, other thanmammals and birds. Temperature will influence growth and repro-ductive output (Pauly, 1980; Vilchis et al., 2005; Moore et al., 2010)and extremes will cause stress and ultimately mortality (Fig. 4;Williams and Morritt, 1995; Helmuth and Hofmann, 2001; Firth andWilliams, 2009; Denny et al., 2011-this issue; Firth et al., 2011-thisissue; Seabra et al., 2011-this issue; Sorte et al., 2011-this issue;Wethey et al., 2011-this issue), but other factors such as extremerainfall will also influence survival (Morritt et al., 2007; Firth andWilliams, 2009). Reproductive and recruitment success seem to drivemost changes, especially at biogeographic boundaries leading to rangeextensions (Herbert et al., 2007; Mieszkowska et al., 2007; Menge etal., 2011;Wethey et al., 2011-this issue). Whilst temperature is clearlyan ultimate factor underlying change (Beaugrand et al., 2002; Harley,2008), its influence can be modified by local proximate factors(Zacherl et al., 2003) and often distributional boundaries are set bycombinations of propagule output, connectivity influenced by larvalsupply, hydrography (Gaylord and Gaines, 2000; Lima et al., 2006;Lima et al., 2007) and habitat quality (Herbert et al., 2009). Whilstpopulation level responses are relatively easy to measure and detect,community level responses are often more subtle and modified byboth direct and indirect effects (Harley, 2006; Morelissen and Harley,2007; Moore et al., 2007a, 2007b; Hawkins et al., 2008; Poloczanskaet al., 2008; Firth et al., 2009; Kordas et al., 2011) with implications forecosystem functioning (Hawkins et al., 2009; Johnson et al., 2011;Wernberg et al., 2011a).

Journal of Experimental Marine Biology and Ecology 400 (2011) 1–6

0022-0981/$ – see front matter © 2011 Elsevier B.V. All rights reserved.doi:10.1016/j.jembe.2011.02.001

Contents lists available at ScienceDirect

Journal of Experimental Marine Biology and Ecology

j ourna l homepage: www.e lsev ie r.com/ locate / jembe

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4. Adapting to change

Responses to climate change at all levels of biological organisation,from individuals through to populations and via communities toecosystems, have become very apparent over the last 15–20 years andare clearly happening now with the potential to accelerate into thefuture.

Climate change will continue until measures to mitigate it byreduction of greenhouse gases takes effect. Inertia in the climatesystem suggests a time lag of at least 50 years. Meanwhile society hasto adapt to climate change (Adger et al., 2005; Parry et al., 2007; Stern,2007). This can only be achieved by managing the interactions withclimate change of those global, regional and local scale impacts we cancontrol: such as spread of non-natives (Fig. 5; Dukes and Mooney,1999; Stachowicz et al., 2002; Stachowicz and Byrnes, 2006; Sax et al.,2007) over-fishing (e.g. Hawkins et al., 2003; Genner et al., 2004,2010; Dulvy et al., 2008; Cheung et al., 2009), eutrophication(Rosenberg, 1985; Smith et al., 1999; Burkholder et al., 2007; Atalah

and Crowe, 2010; Fitch and Crowe, 2011), pollution (Hardman-Mountford et al., 2005; Halpern, 2007) and habitat loss (Reise, 2005;Lotze et al., 2006; Airoldi and Beck, 2007).

Oceanacidificationwas initially viewedas a long-termbut inevitableand predictable consequence of elevated atmospheric CO2 (Ridgwelland Zeebe, 2005; The Royal Society, 2005), with effects likely to bemanifested in around 50 years time. However, more recent work hasindicated that such effects may already be occurring (Wootton et al.,2008; Hofmann et al., 2010; Kroeker et al., 2010). Effects are likely to bemanifested at early life history stages (Findlay et al., 2010; Crim et al.,2011-this issue; Yuet al., 2011-this issue) aswell as adults (Findlayet al.,2009; Gooding et al., 2009) including reproductive potential (Reuteret al., 2011), especially in species with considerable calcification.

5. Structure of the volume

This special volume was by invitation — some people declined,pleasingly most accepted. We also encouraged recommendations for

Fig. 1. Plastic debris floating on the surface of the water in Semporna, Borneo (Thompson et al., 2004; Thompson et al., 2009).

Fig. 2. Artificial coastal defence structure near Llandudno, North Wales (Moschella et al., 2005).

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input from younger colleagues; thus we have a nice mix of papers byestablished experts and longstanding contributors to JEMBE and earlyand mid-career scientists. The combination of reviews, synopticpapers and new data papers has hopefully provided a balance ofaccumulated wisdom and fresh perspectives from around the world.We did reject some papers and as a result, some areas are more thinlyrepresented than others.

The volume is ordered into themes, the front end comprisingreview papers with a regional focus. Johnson et al. (2011) review howchanging physical conditions have precipitated cascading effects ofecological change in benthic rocky reef and pelagic systems inTasmania. Wernberg et al. (2011a) review the observed impacts ofclimate change on subtidal temperate coasts in Australia and assesshow these systems are likely to respond to future warming. Schiel(2011) provides a broad-scale perspective of the east and west coastsof New Zealand to categorise changes in intertidal communitystructure, diversity, and biogeographic differences in dominantspecies and experiments reveal that disturbance to the dominantcanopy results in cascading responses in the mid-shore community.Philippart et al. (2011) discuss the impacts of climate change on theregional seas of Europe highlighting observations, expectations andindicators of change. In addition to climate change, interactions withother regional- and local- scale impacts are considered.

The next section comprises long-term and broad-scale studies,beginningwith twoEuropean synoptic reviewpapers. Henderson et al.(2011) discuss the results of a 30-year study of fish and crustacean

abundance at a site in SW England. Whiteley et al. (2011) review thelatitudinal variation in the physiology of marine gammarid amphipodsbetween Portugal and Svalbard. Merzouk and Johnson (2011) comparethe historic and recent patterns of distribution and abundance of kelpspecies in eastern Canada. Geffen et al. (2011) discuss the populationdynamics of plaice in relation to temperature-dependent nurseryground processes in the Irish Sea. McGinty et al. (2011) discuss thevariation in the responses of zooplankton to climate change in the NEAtlantic. Wethey et al. (2011-this issue) assess the effects of extremeweather during winter 2009/10 on the processes determining thebiogeographic limits of intertidal faunal species in northern Europe.Poloczanska et al. (2011) compare contemporary data with historicaldata to assess whether there were latitudinal changes in distributionand abundance consistent with global climate change along Australia'seast coast. Finally, Mieszkowska and Lundquist (2011) document thebroad-scale geographic distributions, abundances and assemblagepatterns of intertidal limpets around the New Zealand coast.

The next section focuses on the individual level responses to globalenvironmental change. Lockwood and Somero (2011) review theliterature and present new data that document physiologicaldifferences between native and invasive mussel species on theCalifornia coast. Denny et al. (2011-this issue) use the inter-individualvariation in body temperature among intertidal organisms tocharacterisemicro-scale variation in potential thermal stress. Helmuthet al. (2011) explore the sensitivity of aerial mussel body temperaturein relation to variation in environmental conditions in California.Seabra et al. (2011-this issue) examine the relative magnitudes ofsmall-scale versus large-scale latitudinal patterns of intertidal limpetbody temperatures. In a trans-continental comparison, Sorte et al.(2011-this issue) discuss differences in distribution and physiologicaltolerances of intertidal and subtidal species.

We then look at population and community level responses, startingwith a reviewpaper on the effect of temperature on species interactions(Kordas et al., 2011). Giménez (2011) explores how increasedtemperatures can lead to trophic mismatch through an alteration ofthe variance of the abundance distributions of consumers or resourcesover time. Menge et al. (2011) discuss the broad-scale and long-termvariation in recruitment in Oregon. Firth et al. (2011-this issue) presentevidence that thermal stress relating to exposure to cold air tempera-tures caused a mortality event of an invasive mussel in Florida; if suchevents become less frequent in a warmer world then invasive speciesare likely to spread more as they will not be held in check (Thieltges etal., 2004). Wernberg et al. (2011b) examine the relationship betweenocean temperature and patch characteristics of subtidal habitat-formingalgal species along a latitudinal gradient in Australia. Fitch and Crowe(2011) investigate the individual and combined effects of increased

Fig. 3. Settlement of the northern barnacle species Semibalanus balanoides (whiteindividuals) interspersed among adults of the southern species Chthamalus stellatus andChthamalusmontagui (darker individuals) ona rocky shore atNewquay, England followingcold winter 2009/10 (Wethey et al., 2011).

Fig. 4. The intertidal limpet Cellana grata exhibiting “mushrooming behaviour” inresponse to extreme high temperature of rocky substratum, Hong Kong (Williams andMorritt, 1995; Williams et al., 2005).

Fig. 5. The invasive mussel Perna viridis fouling a pontoon in Tampa Bay, Florida (Firthet al., 2011).

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temperature and nutrient and organic enrichment on ecosystemprocesses in intertidal sedimentary systems. Frid (2011) discusses thetemporal variability in the functioning of the benthos over a 30-yeartime-scale in the North Sea. This section focuses primarily on responsesto climate change, particularly temperature, but also includes interac-tions with other impacts such as non-native species, nutrient enrich-ment and fishing pressure.

We then move on to ocean acidification. Crim et al. (2011-thisissue) discuss the effects of elevated seawater CO2 concentrations onlarval development and survival in an endangered abalone, while Yuet al. (2011-this issue) discuss the effects on urchin larval develop-ment. Porzio et al. (2011) describe the effects of increasing CO2 levelson macroalgal communities along a pH gradient caused by volcanicvents in the Mediterrenean Sea.

The last section focuses on anticipating and adapting to globalchange. Connell et al. (2011) focus on a key contingency affectingindirect effects (consumer and producer) and consider their inclusionin ecological forecasts. Chapman and Underwood (2011-this issue)review the major impacts and the experimental research that hasbeen and is being done to build coastal infrastructure in a morebiodiversity-friendly manner. Jackson and McIlvenny (2011) discussthe effects of coastal squeeze from rising sea level on rocky shores inScotland. Finally, Thomsen et al. (2011) present a framework forassessing the impacts of invasive species.

6. Dedication

This volume is dedicated to Dennis Crisp, Alan Southward andEdouard Fischer-Piette — pioneering scientists, who undertook broad-scale surveys (e.g., Fischer-Piette, 1932; Fischer-Piette, 1936; SouthwardandCrisp, 1954a; Fischer-Piette, 1955; Fischer-Piette and Prenant, 1956;Fischer-Piette and Prenant, 1957; Crisp and Southward, 1958; Fischer-Piette, 1963), started long-term time series (Southward, 1967;Southward, 1991) and experimented to find out the processes drivingsuch patterns in the field and in the laboratory. These scientists realisedquite early on the importance of geographic gradients and fluctuationsof climate acting over time in shaping the distributions of species andcommunity structure including modulation of biological interactions(e.g., Southward and Crisp, 1954b). Much of their work has provided abaseline against which to measure change over the last few decades(Herbert et al., 2003;Mieszkowska et al., 2005; Lima et al., 2006; Lima etal., 2007). They have also influenced many of the contributors to thevolume. Their papers are cited throughout the volume despite beingover 50 years old in some cases and some of their data has been used toinform predictions of future states by modelling (e.g., Poloczanska et al.,2008) — truly lasting impact!

Acknowledgments

Compiling this special edition has been a rewarding challenge andto conclude we would like to thank all of those who have beeninvolved in its preparation and production. Firstly, we are verygrateful to all of the authors for accepting the invitation to participateand for adhering to such demanding deadlines. Each contribution hasbeen formally peer reviewed and then edited to draw the contribu-tions together. We wish to thank all of the reviewers who very kindlyprovided thoughtful input for which we are very grateful. We alsowish to thank Kirsteen McKenzie for carrying out much of the copyediting on this volume. Finallywewish to thank the staff at the Journalof Experimental Marine Biology and Ecology for all of their help withthe production of this special volume. Louise Firth and Steve Hawkinshave been supported by the THESEUS project (EU FP7, contractnumber 244104: Innovative technologies for safer European coasts ina changing climate) and the URBANE project (Urban research onbiodiversity on artificial and natural coastal environments: enhancingbiodiversity by intelligent design) funded by the Esmée Fairbairn

Foundation. Several papers in this volume draw on historical data setsrelevant to the NAGISA History of the Near Shore project from whichSJH was supported in the past. S. Hawkins was also supported by aNERC grant in aid for the Marine Biological Association of the UK, plussmall grant NE/E010482/1. We also wish to thank Sandy Shumwayand Roger Hughes for the invitation to edit this volume. [SS]

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Louise B. FirthSchool of Ocean Sciences,

Bangor University, Menai Bridge, Anglesey,LL59 5AB, United Kingdom

Corresponding author.E-mail address: l.firth@bangor.ac.uk.

Stephen J. HawkinsSchool of Ocean and Earth Science,

National Oceanography Centre Southampton,Waterfront Campus, University of Southampton, European Way,

Southampton, Hampshire SO14 3ZH, United KingdomSchool of Ocean Sciences,

Bangor University, Menai Bridge, Anglesey,LL59 5AB, United Kingdom

Marine Biological Association of the UK, The Laboratory,Citadel Hill, Plymouth PL1 2PB, United Kingdom

6 L.B. Firth, S.J. Hawkins / Journal of Experimental Marine Biology and Ecology 400 (2011) 1–6