Rev Fish Biol Fisheries (2014) 24:519–559 DOI 10.1007/s11160-014-9342-1

RESEARCH PAPER

Effects of climate change on Canada’s Pacific marine ecosystems: a summary of scientific knowledge

Thomas A. Okey • Hussein M. Alidina • Veronica Lo • Sabine Jessen

Received: 25 August 2013 / Accepted: 20 January 2014 / Published online: 22 February 2014 Ó Springer International Publishing Switzerland 2014

Abstract The marine life of Canada’s Pacific marine geomorphology, climate, and oceanography, includ- ecosystems, adjacent to the province of British ing a dynamic oceanographic and ecological transition Columbia, may be relatively responsive to rapid zone formed by the divergence of the North Pacific oceanographic and environmental change associated Current into the Alaskan coastal current and the with global climate change due to uniquely evolved California Current, and by currents transporting warm plasticities and resiliencies as well as particular sen- tropical waters from the south. Despite long-term sitivities and vulnerabilities, given this dynamic warming in the region, sea surface temperatures in and highly textured natural setting. These marine Canada’s Pacific have been anomalously cool since ecosystems feature complex interfaces of coastal 2007 with La Nin˜a-type conditions prevailing as we enter a cool phase of the Pacific Decadal Oscillation, possibly masking future warming. When warmer El T. A. Okey (&) Nin˜o conditions prevail, many southern species School of Environmental Studies, University of Victoria, invade, strongly impacting local species and reorga- PO Box 1700, STN CSC, Victoria, BC V8W 2Y2, Canada e-mail: [email protected] nizing biological communities. Acidification and deoxygenation are anomalously high in the region T. A. Okey due to the weakening ventilation of subsurface waters Ocean Integrity Research, 1128 Empress Avenue, resulting from increased stratification. A broad spec- Victoria, BC V8T 1P4, Canada trum of biological responses to these changes are H. M. Alidina expected. Non-climate anthropogenic stressors affect WWF-Canada, 409 Granville Street, Suite 1588, the capacity of biota to adapt to climate changes. It Vancouver, BC V6C 1T2, Canada will be challenging to forecast the responses of V. Lo Á S. Jessen particular species, and to map climate vulnerabilities Canadian Parks and Wilderness Society, 410-698 accurately enough to help prioritize and guide adap- Seymour Street, Vancouver, BC V6B 3K6, Canada tation planning. It will be more challenging to develop forecasts that account for indirect effects within V. Lo University of Bologna, Royal Netherlands Institute for biological communities and the intricate and appar- Sea Research, and Ghent University, Via S. Alberto 163, ently non-deterministic behaviours of highly complex 48123 Ravenna, Italy and variable marine ecosystems, such as those of Canada’s Pacific. We recommend and outline national S. Jessen Department of Geography, Simon Fraser University, and regional climate assessments in Canada and TASC2 Suite 8800, Burnaby, BC V5A 1S6, Canada adaptation planning and implementation including 123 520 Rev Fish Biol Fisheries (2014) 24:519–559 integrated coastal management and marine spatial assessments served as central or initial sources for planning and management. information specific to Canada’s Pacific marine eco- systems, including a series of edited compendiums of Keywords Climate change impacts Á work by Fisheries and Oceans Canada on the state of Acidification Á Deoxygenation Á Climate the physical, biological, and fishery resources of adaptation Á Cumulative impacts Á Global marine Canada’s Pacific marine ecosystems (e.g. Irvine and hotspots Crawford 2013), summaries of that work in the form of State of the Oceans advisory reports (e.g. DFO 2013), and other key publications focused on climate change impacts in the region (BCME 2007; Beamish et al. Introduction 2009; Hutchings et al. 2012; Irvine and Crawford 2011; Johannessen and MacDonald 2009; Lucas et al. Increased emissions of greenhouse gasses since the 2007; McKinnell and Dagg 2010; Perry et al. 2007; industrial revolution, and particularly during the last Walker and Sydneysmith 2007). We also examined few decades, has caused a broad spectrum of physical, regional contributions to the 2010 Canadian Marine chemical, and biological changes in the world’s Status and Trends Report series (DFO 2010) focusing oceans (Brander 2010; Doney et al. 2012; Harley on Canada’s marine ecosystems of the Pacific north et al. 2006; Hobday et al. 2006; Poloczanska et al. coast and Hecate Strait (Cummins and Haigh 2010) 2007; Poloczanska et al. 2013) and many of these the Strait of Georgia (Ianson and Flostrand 2010) and changes are expected to accelerate (IPCC 2007; the West Coast of Vancouver Island (Johannessen and Solomon et al. 2007). We use the term climate change McCarter 2010). In terms of biological changes, our to refer to marine environmental changes relating to assessment focused on three of the four identified the emissions of fossil fuels including ocean acidifi- bioregions of Pacific marine area (sensu DFO 2009)— cation, ocean de-oxygenation, sea-level rise, and other the Northern Shelf, the Southern Shelf, and the Strait physical and chemical changes that are not climate per of Georgia—and less so on Canada’s offshore Pacific se. bioregion (Fig. 1). Planning for the effects of climate change in a There is a spectrum of modelling approaches in particular region or locality, including the develop- development for characterizing and forecasting the ment of adaptation strategies, requires knowledge effects of climate change on the ecosystems of this and about physical changes, potential impacts, and vul- surrounding regions. These include, but are not limited nerabilities on regional and local scales. The first step to, development of end-to-end marine and fishery to understanding these regional and local manifesta- ecosystem modelling (e.g. Kaplan et al. 2012; Kaplan tions of global climate change impacts is to undertake and Leonard 2012; Ruzicka et al. 2012; Steele 2012; detailed examinations of particular regions of interest. Steele et al. 2012), future variations of which may We conducted such an assessment for Canada’s include outputs of global climate models downscaled Pacific marine ecosystems (Okey et al. 2012), and and enhanced as regional ocean models (ROMS) (e.g. the present paper is a synopsis of our review of the Foreman et al. 2008, 2012) as well as potential current knowledge of observed and expected climate- integration of such models with fishery-ecosystem related physical, chemical, and biological changes in modelling approaches that include whole biological this region. The update of the second part of that communities (e.g. Ainsworth et al. 2011), bioclimatic work—a preliminary spatial mapping of climate envelope modelling (Cheung et al. in press, 2009, change vulnerability for Canada’s Pacific marine 2010, 2011), fisheries models (e.g. Agostini et al. areas is forthcoming (Okey et al. unpublished data). 2008), approaches that include social and economic We reviewed existing literature and information systems (e.g. McCay et al. 2011; Miller et al. 2010), about climate change impacts in Canada’s Pacific and other modelling initiatives and approaches (e.g. marine ecosystems, the broader Northeast Pacific, and King et al. 2011; Stock et al. 2011). Most modelling globally, in order to adequately characterize all the and analytical approaches applied to this region categories of change occurring in Canada’s Pacific benefit from broad international collaborations with marine areas. Several existing reviews and long-term goals for developing useful forecasting 123 Rev Fish Biol Fisheries (2014) 24:519–559 521

Fig. 1 The four biogeographic units of the Canadian Pacific Ocean (DFO 2009) in relation to one of 17 Global Marine Hotspots, which are areas warming faster than 90 % of oceans (Hobday and Pecl 2013) (map produced by E. Gartner)

tools (Hollowed et al. 2009, 2011, 2013, Okey et al. ecosections of Canada’s Pacific marine region, and a 2008). A thorough review of knowledge about climate brief summary discussion of options and recommen- change impacts provides value that is independent of dations for climate adaptation planning. such modelling, but it is also provides key information for informing such models and tools. The information herein has already informed a preliminary spatially- The context of Canada’s Pacific ecosystems explicit climate vulnerability assessment of Canada’s marine ecosystems (Okey et al. unpublished data). Canada’s Pacific marine ecosystems extend from 48° The close proximity of Canada’s Pacific marine to 53° North latitude in the Northeastern Pacific Ocean ecosystems to one of the ocean’s hotspots of predicted and from the intertidal zone of the Province of British temperature change (Fig. 1) foreshadows rapid eco- Columbia to the 200 nautical mile boundary of the logical changes on a broad scale. The current review is Exclusive Economic Zone (Fig. 2). This area includes framed in the context of the initial work of the Global and encompasses a highly complex and dynamic Marine Hotspots Network (Hobday and Pecl 2013). coastal oceanographic transition zone where the Our present contribution includes a description of eastward-flowing North Pacific Current intersects Canada’s Pacific marine ecosystems, including the with North America and diverges into the northward three coastal bioregions, a review of various physical flowing (downwelling) Alaska Coastal Current and the and chemical changes in the region’s climate and southward flowing (upwelling) California Current. In oceanography, an overview of the impacts on the addition, the California Undercurrent extension trans- region’s marine life, a review of sensitivity and ports warm tropical waters to at least Vancouver responses of key taxonomic groups and species, a Island (Thomson and Krassovski 2010), and the review of the sensitive characteristics of the marine Davidson Current—a wind-driven coastal

123 522 Rev Fish Biol Fisheries (2014) 24:519–559

Fig. 2 Map of Canada’s Pacific marine ecosystems and the British Columbia coastline with place names that are referred to in the text (map produced by S. Agbayani)

countercurrent inshore of the California Current— 1997; Tillmann and Siemann 2011). Coastal and flows seasonally from Mexico to Canada’s Pacific. inland mountain ranges capture considerable heat and Water properties along the coast are strongly influ- a high rainfall from saturated Pacific air (Salmon enced by conditions that enhance these flows from the 1997) and they hold a seasonal snowpack, glaciers, south, such as the El Nin˜o phase of the El Nin˜o and ice caps (UNEP 2007), thereby shaping particular Southern Oscillation (ENSO), and these conditions spatial and temporal patterns of high coastal runoff. A encourage the invasion of southern species (e.g. variety of mixing processes including tidal, estuarine, Mackas et al. 2007; Mackas and Galbraith 2002). and wind-driven upwelling (Perry et al. 2007; Whitney The latitude of this complex transition zone fluctuates et al. 2005) make coastal waters nutrient-rich through- in response to oceanographic variability including the out the growing season and drives the production of relative strength of these currents, as well as the El phytoplankton and higher trophic levels in the region. Nin˜ophase of ENSO (Perry et al. 2007; Royer 1998).A Runoff through the Fraser River and the many other highly-articulated and glacially-sculpted coastline, coastal watersheds also influences the coastal ocean- which includes fjords, headlands, islands, and bea- ography (Conway and Johannessen 2007), but river ches, indicates similar complexities of the submarine inputs tend to reduce productivity since they form a geomorphology, and this affects the fine-scale vari- fresh surface layer devoid of nitrate, though large ability of the area’s coastal and offshore oceanogra- rivers do provide silicate. Generally high production phy. Thomson (1981) provided the seminal rates are indicated by moderate phytoplankton bio- description of the region’s oceanography. mass but high fishery yields (Ware and Thomson Coastal British Columbia makes up the central core 2005). of the coastal rainforest extending from south-central The interplay of the spatiotemporal variability of Alaska to Northern California (Schoonmaker et al. oceanography, geomorphology, and coastal

123 Rev Fish Biol Fisheries (2014) 24:519–559 523 hydrology, combined with high and varying primary introductory review of modelling approaches, but productivity, creates a highly complex, dynamic, and useful forecasts may take some time. Planning for rich mosaic of habitats and niches for the organisms climate change effects on Pacific Canada’s marine inhabiting the region. This has allowed the develop- ecosystems thus requires frameworks that are simpli- ment of a relatively rich fauna and flora (Lo et al. fied, generalized, and available. The present assess- 2011) with certain taxa such as sea stars (Asteriodea) ment of effects was thus conducted within a hierarchy exhibiting a very high species diversity (Lamb and of meaningful scales including broad biogeographic Hanby 2005), while other taxa have relatively low units, ecosections, and habitats. These three scales species diversity. were each feature-based and self-similar, and were These unique conditions also promote the presence used to organize observed and projected changes and of migratory and transient species including stopover to summarize vulnerability (Okey et al. unpublished seabird migrants, destination migrants such as whales, data). sea lions, and salmon, and environmental migrants The four biogeographic units of Canada’s Pacific such as zooplankton and pelagic fish species that marine areas (DFO 2009) are shown in Fig. 1 along- migrate with climate variability (Perry et al. 2007). side the Northeastern Pacific hotspot of projected Pacific salmon famously provide an important transfer temperature change identified by the Global Marine and concentration of nutrients from the open ocean to Hotspots Network (Hobday and Pecl 2013). The British Columbia’s terrestrial and freshwater ecosys- twelve ‘ecosections’ within Canada’s Pacific marine tems (Cederholm et al. 1999). The ecosystem services areas (Fig. 3) were previously identified based on the provided by this productivity and unique diversity of British Columbia Marine Ecological Classification marine life and habitats supported the flourishing of (BCMEC), which was developed by British Colum- First Nations civilizations and communities, as well as bia’s Provincial Land Use Coordination Office (BC all of Pacific Canada’s contemporary human commu- 2002). nities and economies. We thus begin our assessment of climate impacts In a summary of an extensive synthesis of ecolog- and vulnerabilities in Canada’s Pacific marine eco- ical information about the northern marine area of systems with overviews of British Columbia’s three British Columbia (Lucas et al. 2007), Perry et al. coastal biogeographic units: the Northern Shelf, the (2007) emphasized phytoplankton, nearshore macro- Southern Shelf, and the Strait of Georgia. phytes, and detritus as the three foundations of the marine food webs of British Columbia, and they Northern Shelf suggested that this trophic structure appears robust to the reduction or elimination of single food-web The Ecosystem Status and Trends report for the North components where there are other species in the same Coast and Hecate Strait ecozone (Cummins and Haigh functional group. Such groupings are, however, scale- 2010) highlights trends such as increased temperature related and the degree of functional redundancy is and decreased salinity of upper ocean waters, decreas- context-dependent and thus not well understood. Kelp ing dissolved oxygen, increasing dissolved CO2, and and eelgrass habitats do stand out as providing potential biological changes, such as shifting species important food sources and biogenic habitat for fishes distributions. Such changes are explained in more and invertebrates in most of Pacific Canada’s near- recent literature (e.g. Whitney et al. 2013) and in shore food webs. following sections herein. Productivity around the The ecological complexity of the region is high, as boundaries of the Northern Shelf area is driven by are uncertainties about climate effects on the biota. seasonal and episodic switches in upwelling and These impacts and interacting features are furthermore downwelling, variable seasonal patterns of freshwater spatially and temporally heterogeneous and fractal. runoff from watersheds, and enhancement by strong Understanding these ecosystems and their components tidal mixing in narrow passes and channels and at is thus particularly challenging. Mechanistic projec- other ecological interfaces. The banks on the conti- tions of climate effects on system components, or on nental shelf in the region are separated by large and the whole system, will emerge increasingly in the deep troughs or submarine canyons connecting to the future for this or similar regions, as indicated by the deeper continental slope. The Northern Shelf area is 123 524 Rev Fish Biol Fisheries (2014) 24:519–559

Fig. 3 Coastal marine ecosections of British Columbia (Can- transitional Pacific ecosections are not shaded in this figure. ada’s coastal Pacific) (shaded) and dynamic ecosystem features Based on figure in J. Mathias, DFO Oceans Directorate, (black circles and lines). The subarctic Pacific and the unpublished report ‘‘Rationale for the LOMA Boundary’’ 2003. less studied by western science than the southern Southern Shelf coastal bioregions, but it coincides with the Pacific North Coast Integrated Management Area (PNC- The Southern Shelf bioregion, which includes the west IMA)—one of Canada’s five featured Large Ocean coast of Vancouver Island (WCVI) and the Juan de Management Areas (LOMAs) for implementing Fuca Strait, is exposed to spatially and seasonally aspects of Canada’s Oceans Act, as well as the complex interfaces of estuarine outflow, upwelling regional study area of the Marine Planning Partnership patterns, and local and regional currents and eddies, (MaPP) for the North Pacific Coast, and so is the focus including the prominent Juan de Fuca Eddy over the of initiatives for integrated marine resource planning. shelf straddling the international border (e.g. Foreman First Nations such as the Haida Nation are taking et al. 2008; MacFadyen et al. 2005, 2008). Runoff is a prominent role in marine planning in their traditional moderate and more seasonal from the smaller water- territory (Jones et al. 2010). Supporting compendiums sheds and fjords of the WCVI, whereas a major of ecological information have been assembled (Lucas buoyancy current from the Fraser River watershed and and Jamieson 2007; Lucas et al. 2007; Perry et al. other watersheds of the Strait of Georgia and broader 2007). Trophodynamic ecosystem modeling has also Salish Sea flow through the glacially-carved Juan de been conducted in this region (e.g. Ainsworth et al. Fuca Strait, as do strong tidal currents to influence 2002; Ainsworth and Pitcher 2005a, b, c, 2006; outer coastal waters. These processes are all strongly Ainsworth et al. 2008; Gue´nette et al. 2007), including influenced by ENSO and potentially other oceano- geographically broader work specifically addressing graphic oscillations (Dallimore et al. 2005; Okey and the combined effects of different climate change Dallimore 2011) and exhibit complex interactions impacts and fisheries (Ainsworth et al. 2011; Preikshot with migratory fauna in both parts of the Southern 2007). Shelf Bioregion. Several studies have examined the

123 Rev Fish Biol Fisheries (2014) 24:519–559 525 productivity, fisheries, and other biological implica- climate conditions in Canada’s Pacific marine areas tions of climate variability and change at different including winds, currents, water temperature, salinity, time scales in this area (e.g. Beamish et al. 2009; oxygen content, and pH. Some of these changes will Dallimore and Jmieff 2010; Mackas et al. 2007; result from local effects, such as on temperature and Robinson and Ware 1999; Ware and Thomson 2000, precipitation, while others will emerge through 2005; Wright et al. 2005). Fisheries and Oceans regional and larger-scale rearrangements of atmo- Canada (DFO) recently completed its Ecosystem spheric and ocean circulation. The observed and Status and Trends report for the WCVI (Ianson and predicted changes to a number of these physical and Flostrand 2010). A Social-ecological Assessment has chemical parameters have been summarized by Jo- been conducted for social-ecological system sustain- hannessen and Macdonald (2009) for the Strait of ability planning (Okey and Loucks 2011). Some Georgia and in McKinnell and Dagg (2010) for the ecosystem modeling has been conducted in the region, North Pacific Ocean, and most recently by Whitney including some related to climate change impacts et al. (2013) regarding important changes to the North (Ainsworth et al. 2011; Cisneros-Montemayor 2010; Pacific pycnocline. Espinosa-Romero et al. 2011; Preikshot 2007). Long-term trends in physical and chemical condi- tions in Canada’s Pacific waters are expected be Strait of Georgia consistent with global trends qualitatively (i.e. war- mer, more acidic, more stratified, rising sea level), but The Strait of Georgia bioregion is the most studied of climate change may intensify the North Pacific all of Pacific Canada’s bioregions, and is considered to currents and potentially the Pacific atmospheric sub- be generally sensitive to climate change impacts given tropical anticyclone (IPCC 2007) leading to a north- its geomorphology and oceanography, strong influ- ward shift in the oceanographic transition zone and ence of rivers, estuarine and tidal circulations, and increased California Current upwelling (Snyder et al. proximity and exposure to relatively high human 2003). Even if the latter does not extend to Canada’s populations and related stressors (Johannessen and Pacific region, it may affect the composition of waters MacDonald 2009). This bioregion has been exposed to flowing northward. The El Nin˜o/Southern Oscillation the longest history of contemporary human activities, may also become more frequent and more intense (Lee and has thus been modified more than others in and McPhaden 2010; Merryfield 2006; Trenberth et al. Canada’s Pacific (e.g. Ban et al. 2010; Pauly et al. 2002), strongly affecting the position and dynamics of 1998; Pitcher 2005). Expected changes include warm- the transition zone and associated marine life. There is ing, increases in pH, decreases in dissolved O2, effects good evidence that ecosystem variability is increasing of sea-level rise and storms on low-lying and sensitive in the California Current (Bograd et al. 2010). areas, seasonal shifts in biological production and food Oceanographic changes in Canada’s Pacific stood web structure, adverse effects on Pacific salmon out from average global changes in some other unique species, and increased stress to endangered species ways as well. La Nin˜a conditions since 2008 brought such as killer whales (Johannessen and MacDonald anomalously low sea surface temperatures to Canada’s 2009). Fisheries and Oceans Canada has produced a Pacific, as measured in the region (see Irvine and broader technical document on the status and trends of Crawford 2012), interrupting a measured long-term the Strait of Georgia ecosystem (Johannessen and trend of increasing ocean water temperatures in the McCarter 2010). Various ecosystem modeling region of about 1.0 °C per century (BCME 2007; approaches are being developed for the Strait of Beamish et al. 2009; Freeland 2013; Whitney et al. Georgia, including trophodynamic modelling (Li 2007). These cool conditions may be temporarily et al. 2010; Preikshot et al. 2013). masking the underlying warming trajectory, as there are indications that warming rates are accelerating globally (IPCC 2007; Reid and Beaugrand 2012; Changes in climate and oceanography Solomon et al. 2007) and that rivers and adjacent nearshore waters may be warming twice as fast as Increased atmospheric concentrations of greenhouse offshore waters (Morrison et al. 2002; Reed et al. gases will affect a variety of oceanographic and 2011; Rodenhuis et al. 2007; van Vliet et al. 2013; 123 526 Rev Fish Biol Fisheries (2014) 24:519–559

Walker and Sydneysmith 2007; Wu et al. 2012). (Whitney et al. 2007). At this rate, it would take less Secondly, the surface waters of northeast Pacific than 100 years for current SST at the extreme coasts are among the most acidic on earth due to northwest point of the Canada’s Pacific coast (Langara upwelling of deep ocean water (Feely et al. 2008; Island, British Columbia) to be similar to those Hauri et al. 2009) and Canada’s Pacific region is currently observed off the southwest coast of Van- likewise exceptionally acidic (Ianson 2008). Thirdly, couver Island (Amphitrite Point). This rate, however, dissolved oxygen in the subsurface waters of the may accelerate, with projections between 0 and 1 °C northeastern Pacific Ocean declined steadily due to from 2015 to 2025, and between 1 and 2 °C from 2045 increased stratification in the upper water column in to 2055 (IPCC 2007; Solomon et al. 2007). These the North Pacific Ocean, measured off the West Coast trends are consistent with rises in the global sea of Vancouver Island and elsewhere, which was in turn surface temperatures. due to decreased surface salinity (Whitney et al. 2007) However, La Nin˜a oceanographic and atmospheric resulting from increased precipitation and glacial conditions in the Eastern Pacific have led to anoma- ablation in the region (BCME 2007; Larsen et al. lously cool water in the Northeastern Pacific, includ- 2007; Rodenhuis et al. 2007; Walker and Sydneysmith ing Canada’s Pacific waters, almost every year since 2007). Warming sea surface temperatures reinforces 2007 (Crawford and McKinnell 2013; Irvine and this stratification and the resulting decreased oxygen Crawford 2013). It is not known whether climate (Freeland et al. 1997; Freeland 2013; Whitney et al. change will lead to persistently cooler waters in this 2007). region thorough its influences on oceanographic Changes related to the natural variability and patterns and cycles such as the El Nin˜o Southern complex oceanography of the region can be construed Oscillation (ENSO). erroneously as broader-scale or global changes in climate and oceanography. Ecological changes in the Precipitation and hydrology oceans are being driven by changes across the temporal spectrum from short-term natural variability Precipitation increased during the last 50 years over to long-term climate change. Distinguishing natural much of British Columbia during all seasons by some variability from these directional changes is a great estimates (Rodenhuis et al. 2007) or just during challenge, and is crucially important for understanding warmer months by other estimates (BCME 2007; climate change and its impacts. In the North Pacific, Walker and Sydneysmith 2007), based on observa- the climate change signal is projected to become tions. Future projections indicate wetter winters. stronger than the climate variability signal by approx- Projections from the Canadian Regional Climate imately 2035 (Overland and Wang 2007). Certain Model, forced by the A2 emissions scenario, corre- climate-related changes are already considered to have spond well with the global models regarding the wetter emerged from the noise, such as hypoxia and north- annual conditions in British Columbia (Rodenhuis ward shifts of some biota (F. Whitney, Fisheries and et al. 2007), but they indicate increased precipitation Oceans Canada, pers. comm., 2 March 2009). Changes in both winter (?14 %) and summer (?10 %) for the in variability itself might be an important aspect or 2050s. The same model projects an increase in indicator of climate change (Alheit et al. 2010; Katz precipitation of up to 25 % for both summer and and Brown 1992). winter over the north coast region. Other projections generally agree on 6 % increases in precipitation Temperature (Rodenhuis et al. 2007). Extreme precipitation events are expected to increase during the coming century in Sea surface temperatures (SST) increased by about most regions of the world (IPCC 2007; Kharin et al. 1.0 °C in the Strait of Georgia and elsewhere around 2007; Solomon et al. 2007) and in British Columbia British Columbia during the last century based on during some seasons and in some areas (Pike et al. lighthouse records (BCME 2007; Beamish et al. 2008). 2009). This is consistent with temperature increases Increased runoff from British Columbia’s water- at isopycnal surfaces between 100 m and 400 m along sheds is expected, due mainly to wetter winters. This Line P off the west coast of Vancouver Island can increase turbidity, sedimentation rates, and 123 Rev Fish Biol Fisheries (2014) 24:519–559 527 organic material in rivers, lakes, and near-coastal Oxygen waters causing more hypoxia and other negative impacts on marine communities. However spring Dissolved oxygen is expected to decrease in the global snowpack will decrease in British Columbia by about ocean by 1–7 % this century (Keeling et al. 2010; 55 % by the 2050s (Rodenhuis et al. 2007), causing Rabalais et al. 2010; Sarmiento et al. 1998), but these the annual flow of rivers to shift to earlier in the changes are occurring much more rapidly in the North season. Glacial runoff increases and then decreases in Pacific Ocean where oxygen has already decreased by responses to warming due to melting. Most of BC’s 22 % during the last 50 years at depths of between 100 glaciers have already passed the first phase and runoff and 400 m (Batten et al. 2010; Whitney et al. 2007). is already decreasing (Stahl et al. 2006). Glaciers of This is a rate of 0.39–0.70 lM year-1 or an integrated northern British Columbia are not yet in an advanced rate of 123 mmol m-2 year-1. Similar trends have state of decay, so they will likely have more summer been measured along Canada’s Pacific coast, with loss runoff through the year 2050, while southern British rates of 0.5 to 1.0 lM per year being common below Columbia glaciers already have less summer runoff the surface mixed layer (Crawford and Pena 2013; (Irvine and Crawford 2011). These emerging changes Whitney et al. 2013). will add to the stress on stream ecosystems already Subsurface dissolved oxygen along Canada’s impacted by drier, warmer, and less-oxygenated Pacific coastlines originates largely from ventilation summer conditions, and otherwise modify the charac- in and near the Okhotsk Sea, which is dependant to ter of aquatic ecosystems and groundwater. The very some degree on the amount of ice formed each winter. southern part of the province may see net decreases in In the past few decades, less ice has resulted in less precipitation. dense water formation there, and therefore less oxygen transport into the pycnocline waters (150–500 m) of the subarctic Pacific (Whitney et al. 2013). Warming Salinity and stratification in Siberia during winter is thus affecting oxygen supply to the habitat of the groundfish, shrimp, and The salinity of the surface waters of the North Pacific hake, and other biota in Canada’s Pacific marine Ocean has decreased, as indicated by a measured ecosystems. freshening at Ocean Station P, west of Vancouver This oxygen decline in waters between 250 and Island, during the last 50 years at an average rate of 400 m depth along the British Columbia’s continental 0.0036 year-1 (Whitney et al. 2007). Increased strat- shelf affects commercial fish populations by decreas- ification of the upper ocean is reducing ventilation of ing or compressing available fish habitat. Examination the ocean interior. However, nutrients are accumulat- of 11 years of fisheries data and *25 years of ing near the base of the ocean mixed layer, resulting in oceanographic data shows that the vertical range of no loss of nutrients to the surface layer during winter both groundfish and oxygen rich waters are shrinking mixing (Whitney 2011; Whitney et al. 2013). The by about 3 m per year (F. Whitney, Pers. Comm. 8 nutrient enriched, oxygen deprived waters of the ocean March 2012). These findings are consistent with pycnocline are upwelled onto the continental margin published declines of oxygen and compression of where they can cause serious hypoxia such as has been habitat for deep water fish in California (Bograd et al. observed off the Washington and Oregon coasts in 2010; Koslow et al. 2011; McClatchie et al. 2010). recent years (Chan et al. 2008). Stratification can also Survey data also show that oxygen declines are be reinforced by warming sea surface temperatures occurring in all areas of the subarctic Pacific (Keeling (Freeland et al. 1997; Whitney et al. 2007). Nearshore et al. 2010). The oxygen depletion caused by increased biological communities may be modified strongly by precipitation and stratification and decreased ventila- changes in seasonal and episodic salinity patterns and tion often causes mobile organisms to take refuge in fluctuations in these settings. In general, the salinity in shallow water (McKinnell and Dagg 2010) where they most coastal areas of Canada’s Pacific, will decrease can succumb to physiological stress or predation. (especially during winter), especially in the north, but Hypoxic or anoxic water impinges on the North it may decrease in the southern portion of Vancouver American west coast where it has affected marine life Island due to potential decreases in rainfall and runoff. considerably in recent years, often being referred to as 123 528 Rev Fish Biol Fisheries (2014) 24:519–559 anoxic or ‘‘dead’’ zones (Chan et al. 2008; Dybas extent. Tidal mixing in the Juan de Fuca Strait may

2005; Grantham et al. 2004). Such anoxic zones elevate ocean CO2 concentrations there and in the strongly affect marine life; they are expected to Vancouver Island Coastal Current (Ianson et al. 2003). become more widespread, and will co-occur with The Northeastern Pacific Ocean has some of the acidification hot-spots. lowest pH in the world because it is at the end of the

The coastal fjords of British Columbia may be ocean conveyer belt, and respired CO2 has built up in particularly vulnerable to decreased oxygen due to these ancient upwelling waters (Feely et al. 2008; their geomorphological and chemical conditions (Dal- Hauri et al. 2009). Marine organisms of the area have limore and Jmieff 2010; Dallimore et al. 2005; Hay thus adapted to these marginal pH conditions, but et al. 2009; Johannessen and MacDonald 2009). The continued pH declines may lead to fundamental spring and summer depths of hypoxic waters in ecological shifts, which have already been docu- Saanich Inlet, a fjord of southeastern Vancouver mented in the region (Wootton et al. 2008). Island, is now 25 meters shallower than it was 50 years ago (F. Whitney, pers. comm., 2 March Sea level 2009). Saanich Inlet may be somewhat unique among coastal fjords in Canada’s Pacific, but qualitatively Global sea level is expected to rise by 10–22 cm by similar changes may occur generally. 2050, and by 21–44 cm by 2100 (IPCC 2007), and it is expected to continue rising for at least decades due to Ocean pH anthropogenic warming. Sea-level will rise at different rates along Canada’s Pacific shorelines as it is strongly Oceans have absorbed about 130 billion metric tons of influenced by variations in uplift and subsidence and

CO2 over the last 250 years—more than a third of all oceanographic factors (Barrie and Conway 2002; human-produced CO2 emissions (Feely et al. 2008)— Cherniawsky et al. 2004; IPCC 2007; Thomson et al. thereby decreasing ocean pH to the lowest levels in 2008; Thomson and Crawford 1997; Walker and 20 million years (decreased pH equals increased Sydneysmith 2007). The mass subsidence of the acidity) (Caldeira and Wickett 2003, 2005; Feely Fraser River Delta, at Vancouver, for example, will et al. 2004; Hauri et al. 2009; Orr et al. 2005; Sabine augment (accelerate) global sea-level rise there by as et al. 2004). This increased acidity impacts the ability much as 1–2 m during the coming century (Mazzotti of many organisms to produce and maintain their et al. 2009). Elsewhere along Canada’s Pacific coast, calcium carbonate structures (Feely et al. 2004, 2008), tectonic uplift and crustal rebound from the weight of and stresses other metabolic processes. The IPCC AR4 ice sheets that covered the region 12,000 years ago model mean indicates a decrease in global surface will compensate sea-level rise to varying degrees ocean pH ranging from 0.1 to 0.2 units by 2050. Other (Thomson and Crawford 1997). For instance, tectonic models suggest that the pH of surface oceans will lift related to plate collision and subduction is forcing decrease by 0.3 to 0.4 units by the end of the century Vancouver Island to tilt such that the relative sea level (Feely et al. 2008). Corrosive waters are occurring first at the west coast Vancouver Island town of Tofino is at higher latitudes, such as the Northeast Pacific, due to dropping at an average rate of 16 cm per 100 years temperature effects on CO2 absorption (Byrne et al. (Irvine and Crawford 2012), though the next Cascadia 2010; Orr et al. 2005). This can potentially cause Subduction Zone earthquake could suddenly drop the major shifts in distributions and community assem- land by a metre and cause a massive tsunami. Other blages by both latitude and depth. factors that influence sea level rise, such as erosion and The aragonite saturation depth along Canada’s deposition rates, vary on small scales. Pacific coast is presently between 200 and 400 m, and Regional thermal expansion and salinity effects in it may decrease to between 100 and 400 m during the British Columbia will also exacerbate the expansion 2040s (Guinotte and Fabry 2008), reducing habitable due to global ocean warming (IPCC 2007; Thomson space for sensitive species. Upwelling of aragonite- et al. 2008). In the Strait of Georgia, thermal unsaturated water into the continental shelf of western expansion and salinity account for 37 and 63 % North America already occurs on a seasonal basis respectively, of total steric sea-level rise (Thomson (Feely et al. 2008), but will increase in frequency and et al. 2008). The combined effect of a Cascadia 123 Rev Fish Biol Fisheries (2014) 24:519–559 529 earthquake, a tsunami, and sea level rise could raise Strengthened upwelling has indeed been observed in both sea level and the height of a tsunami by as much the California coast region over the last 30 years, and as a metre (B. Crawford, DFO, Personal communica- regional climate model simulations indicate future tion, 10 August 2012). upwelling intensification (Snyder et al. 2003). These The mean observed absolute sea-level rise (not deep waters are richer in nutrients, but poorer in oxygen accounting for steric sea-level rise) in the province is and more acidic than waters usually occupying the in agreement with estimates of global eustatic sea- shelf, and may generate significant ecosystem impacts level rise (1.5–2.0 mm/year) observed from tide gauge in the future (Chan et al. 2008). In 2005, upwelling was data, in addition to the global IPPC estimates delayed in the northern California Current, which (1.85 ± 0.5 mm year-1 between 1961 and 2003) reduced production and energy flow throughout the (Thomson et al. 2008). A 95 year dataset at Prince food web, and helps explain the anomalous nekton Rupert, British Columbia, revealed a rising rate of occurrences that year (Bograd et al. 2010; Brodeur et al. 11–12 cm per century (Abeysirigunawardena and 2005; Trudel et al. 2006;Wing2006). Walker 2008; BCME 2007), which is consistent with another estimate of 1.4 ± 0.6 mm per year between Oceanographic oscillations 1939 and 2003 (Abeysirigunawardena and Walker 2008). However, an addendum to Thomson et al. The El Nin˜o-Southern Oscillation (ENSO) strongly (2008) estimated that sea level rise along the coast influences oceanographic and biological variability in could range from 52 to 103 cm by 2100 in an the Northeast Pacific Ocean and coastal zones, with RSL2100 High Scenario. conditions being warmer than average during El Nin˜o Coastal sensitivity to sea level rise depends on and colder than average during La Nin˜a. The ocean- geology, such that most impact will be felt along sandy, ographic impacts related to changes in ocean temper- low-lying regions. An analysis of coastal sensitivity to ature, stratification, precipitation, winds, productivity, sea-level rise has been developed for all of Canada and and upwelling are all influenced by the warm and cold for British Columbia (BCME 2007; Shaw et al. 1998; phases of the ENSO. There is some indication that El Thomson et al. 2008) based on a variety of parameters. Nin˜ophases might become more frequent or change in According to these estimates, most of the mainland other ways (Merryfield 2006; Trenberth et al. 2002), fjordal systems of Pacific Canada have low sensitivity with every stage of the oscillation becoming faster so to sea-level rise, but the sensitivity is moderate around that anomalies would be of shorter duration. In such a most of Vancouver Island and in some northern areas, scenario, impacts on more sensitive ecosystem com- including the areas around Prince Rupert and Bella ponents might increase in such conditions, but impacts Bella. Sensitivities are rated as high at the Northeast on more resistant components might diminish. This is corner of Haida Gwaii and at the Fraser River Delta a topic with considerable uncertainty, in terms of both (see also Mazzotti et al. 2009). the physical and the biological implications of any effect of climate change on ENSO. The Pacific Ocean currents Decadal Oscillation (PDO) is a longer-term signal with similar warm and cold phases, but there are few The North Pacific Current, which brings warm waters hints that climate change will alter the PDO drasti- from the central Pacific, and the California Current, cally. Even if changes in these oscillations are which flows southward from British Columbia to minimal, their strong effects will be overlaid on a Mexico, may intensify with global climate change different, potentially less resilient, mean state, given (IPCC 2007). Wind regulates the latitude at which the the combined effects of multiple climate stressors, not Pacific Current impinges on the coast, and projections to mention other local and regional anthropogenic of changes in winds at Canada’s Pacific coasts are stressors. somewhat uncertain (IPCC 2007). Warming may also result in a northward shift and intensification of the Storminess and wave heights subtropical anticyclone (IPCC 2007) leading to a northward shift of the transition zone, and a strength- A recent ensemble analysis of IPCC global climate ening of the upwelling in the California Current. models (Ulbrich et al. 2008) indicated a poleward shift 123 530 Rev Fish Biol Fisheries (2014) 24:519–559 in storm activity globally (see also Easterling et al. including both structural and functional changes. This 2000), including a 7 % increase in storms near the may be especially important when the signal of long- Aleutian Islands by 2100, but it indicated no changes term directional change (i.e. non-stationarity) in Canada’s Pacific region. A recent study by Ruggiero becomes the dominant forcing over the noise of et al. (2010) measured[50 cm increases in Significant climate and oceanographic variability in the North Wave Height at buoys off Oregon since the mid- Pacific, which is expected to occur around 2035 1970s, with the very largest waves increasing by about (Overland and Wang 2007). Such rapid reorganization 2.5 m. This is consistent with predictions of the U.S. can be accelerated by problems such as match- Climate Change Science Program that increased ocean mismatch where co-evolved species become separated temperatures will lead to increased intensities of in space or time. hurricanes and extra-tropical storms (Karl et al. A conspicuous seabird example of predator–prey 2008). However, Canadian scientists have not found mismatch indicates the types of ‘production mis- a similar trend in Canada’s Pacific waters (Gemmrich match’ problems that will become increasingly appar- et al. 2011). There is uncertainty in predictions of ent for other taxa. The population peak of the copepod wave heights in Canada’s Pacific. If storminess does Neocalanus cristatus began occurring earlier in the increase in this region, it may cause considerable year as ocean temperatures increased during the 1990s coastal inundation and erosion (Thomson et al. 2008; (Bertram 2001) causing a mismatch between the peak Walker and Sydneysmith 2007), especially when timing of Cassin’s auklet nestlings on Triangle Island, combined with future sea-level rise and high tides British Columbia and the availability of this critical (Thomson et al. 2008). food, eventually leading to complete reproductive failure of the auklets in 2010 (Borstad et al. 2011; Hipfner 2008) (Table 2). This example foreshadows Overview of impacts on marine life broader productivity-related ecological changes in Canada’s Pacific given the responsiveness of plankton A unique, rich, and productive diversity of marine life in the region to climate variability and change (Batten inhabits Canada’s Pacific marine areas, and this and Mackas 2009; Bertram 2001; Mackas et al. 2007, marine life benefits human communities and econo- 1998). mies both directly and indirectly. Climate change will Harley (2011) provided a clear example from have differential effects on all of these species, thereby intertidal ecosystems of Pacific Canada’s Southern modifying species relationships and reassembling Shelf region of how climate change can alter inter- biological communities rather than simply shifting specific relationships with important implications for all species distributions uniformly (Table 2). Effects the structure, functions, and biodiversity of these on marine life will be complex and may include some marine ecosystems. He showed that warming condi- increases in values due to the positive effects of tions favour the predatory sea star, Pisaster ochraceus, potential increases in upwelling and nutrients, and thereby reducing the cover and vertical extent of some rapid negative impacts given changes in oxygen mussel beds, with cascading effects on other associ- and pH (Chan et al. 2008; Ianson 2008; Kleypas et al. ated species. This example provides useful informa- 2005; Miller et al. 2009; Widdicombe and Spicer tion about the effects of one aspect of climate change 2008). These differential effects will affect ecosystem on one community type, but less is known about the structure and function, and thus ecosystem services, in effects of climate change on the structure and dynam- ways that are very challenging to predict or forecast. ics of most other community types in Canada’s Pacific. Resiliency and responsiveness vary among species Sudden rapid changes in the structure and function due to natural partitioning of function and life-history of biological communities, i.e. threshold effects or because species limit their functional similarities shifts to alternate states (Connell and Sousa 1983; (sensu MacArthur and Levins 1967; Okey 2003) when Okey 2004b; Scheffer et al. 2001), may occur much confronted with natural variability in climate and sooner than predicted by oceanographic patterns alone oceanography. However, this non-uniform respon- due to the strong effects that local and regional non- siveness aspect of biological communities to direc- climate stressors can have in reducing the natural tional change may lead to their rapid reorganization resiliency of biological communities (Gunderson and 123 Rev Fish Biol Fisheries (2014) 24:519–559 531

Holling 2002; Holling 1973; Ludwig et al. 1997). top-down) control (e.g. Ainsworth and Pitcher 2005b, Most coastal marine ecosystems, including those in 2006; Ainsworth et al. 2008). If nutrient upwelling, Canada’s Pacific, are exposed to broad suites of local vertical mixing, and river discharge are highly vari- and regional non-climate anthropogenic stressors (Ban able and subject to multiple overlapping cycles, then et al. 2010), and alleviating these local and regional stronger bottom-up control is likely, but if grazers and stressors will be key to increasing the resiliency of predators are adapted to rapidly exploit predictable Canada’s Pacific marine ecosystems to climate change resources, then top-down control may also be strong. impacts. Expert or Delphic approaches can be used to Some mixture of trophic control is inevitable, and so rank the relative importance of these various stressors forecasts will have limited accuracy and usefulness (e.g. Okey and Loucks 2011) in order to prioritize when they do not account for trophic interactions. attention, management, and limited resources, but In the present contribution, we can summarize how quantitative approaches (e.g. Ban et al. 2010) can also climate change might affect Canada’s Pacific marine be used on their own, or in combination with expert ecosystems and biological communities based on approaches, as well as within analyses such as observations of changes, knowledge of these settings, vulnerability assessment, discussed later. ecological theory, and results of some initial model- Meanwhile, observed ecological changes—such as ling. We organize this summary of examples into four increased abundances of southern species, declines of observable themes of changes to marine life: more northern species, and other unusual sightings and changes (Table 2)—provide insights into past, current, Shifts in species distributions, community and future changes, even if these changes are more related composition, and structure to the ENSO and other cycles of climate and ocean variability. Examination of the biological responses to Climate change may shift the distributions, abundances, such variability is indeed a useful analogue for under- timing, and ecological presence of species, thereby standing directional climate change, with the caveat that changing structure and functions of Pacific Canada’s the character of these shorter-term fluctuations and events marine biological communities. Some species are may differ somewhat in both quality and quantity from extremely responsive to changes in oceanographic the physical, chemical, and biological changes caused by conditions including Pacific salmon species, sardines, longer-term climate change. anchovies, and Pacific hake (e.g. Robinson and Ware These complex and uncertain changes in population 1999; Ware and Thomson 2000;Wrightetal.2005) abundances and distributions, mismatches, indirect (Table 2), while others are more adaptable to changing effects, and shifting biotic assemblages are driven not conditions. Changes in phenology, or the timing of life just by changes in temperature, salinity, and produc- stages, or vigor and vulnerability to predation, will also tivity, but also by rapid acidification, which is variably vary among species. Simple physiological stress, or affecting both calcifying and non-calcifying organ- variations in tolerance, can shift the balance of compe- isms alike, and by declines in the dissolved oxygen tition or predation thereby considerably shifting species (Whitney et al. 2007) which is leading to shrinkage in compositions (e.g. Harley 2011). These non-uniform the vertical range of both groundfish and the oxygen responses lead to the spatial and temporal mis-matches rich layer they inhabit by about 3 m per year (F. of co-evolved species discussed previously, and poten- Whitney, Pers. Comm. 8 March 2012), similar to tially non-linear or disproportionately large shifts in trends documented in California (Koslow et al. 2011; communities and assemblages. McClatchie et al. 2010). Spatial distribution shifts will be complex (Schiel Still, the switching between cold and warm regimes et al. 2004) involving vertical (Harley et al. 2006)and emerges as the most conspicuous drivers of climate- shoreward (Brodeur et al. 2006) movements, as well as a related change at higher trophic levels, as mediated by general poleward movement of species in the northeast- primary and secondary productivity. Ware and Thom- ern Pacific (Brodeur et al. 2003, 2005, 2006; Cheung son (2005) suggested that the trophic dynamics in the et al. in press; Harding et al. 2011;Orsietal.2007; Northeast Pacific are strongly bottom-up controlled, Rogers-Bennett 2007a;Trudeletal.2006;Wing2006; but trophodynamic modelling of Canada’s Pacific Zacherletal.2003). Observations from Northeastern ecosystem dynamics indicates mixed (bottom-up and Pacific pelagic fish species (Orsi et al. 2007) have 123 532 Rev Fish Biol Fisheries (2014) 24:519–559 informed a modelling effort in Pacific Canada and overlap of northern and southern biotas. A number of adjacent areas of the Northeastern Pacific that forecasted examples of novel species occurrences during the an average poleward shift of these pelagic species’ 1982–1983 El Nin˜owere documented by Fulton (1985) distribution centroids of 20.9 ± 3.54 km decade-1 and summarized by Okey et al. (2012), and a variety of under the SRES A2 scenario from 2000 to 2050 (Cheung other anomalous occurrences have been recorded more et al. in press). A slower observed mean northward trend recently as well (Brodeur et al. 2006; Trudel et al. 2006; of 12 km decade-1 has been documented for temperate Wing 2006) (Table 2). Impressions of increased bio- demersal species on the Eastern Bering Sea Shelf diversity from these observations, however, may be (Mueter et al. 2009; Mueter and Litzow 2008)as misleading because arrivals of new species are more consistent with other studies (Murawski 1993; Perry conspicuous and thus more reported than are extirpa- et al. 2005), though this varies greatly among species. A tions of endemic species (also discussed later). recent global meta-analysis of marine species responses Climate change can affect the occurrence of inva- indicated a leading edge poleward rate of 72.0 ± sive species in Canada’s Pacific waters by increasing 13.5 km decade-1 and a trailing edge poleward rate of introduction pathways, facilitating establishment suc- 15.4 ± 8.7 km decade-1 (Poloczanska et al. 2013). cess, and promoting species range expansions (Lo et al. Some species are migrants or transient in Canada’s 2010). The occurrence and impacts of marine disease Pacific marine ecosystems, and are affected by changes are likely to increase, such as withering syndrome in throughout their ranges that interact with important life Pacific coast abalone species (Moore et al. 2000; van history stages. Pacific hake come to Canada’s Pacific to Blaricom et al. 1993) or oyster parasites such as in the feed and return to home waters to spawn, but home Northeastern United States (Ford 1996). Hosts already spawning habitat can deteriorate due to changing stressed due to climate change will be more susceptible conditions such as reduced oxygen. Relocating to other to infection (Harvell et al. 2002). Canada’s Pacific will areas where young can survive may be difficult since likely see rapid increase in invasive species such as successful spawning likely requires ocean circulation those from ballast water and on ship hulls, drilling patterns that retain early stages near the coast. Failures in muds, and other vectors. The risk of establishment may cod recruitment in Canada’s Pacific have, for example, be amplified by climate variability and change because been linked to outflow of Hecate Strait waters into Haida invasive species tend to be tolerant to large changes in Eddies during El Nin˜o events, thereby reducing larval temperature and salinity (Levings et al. 2004). The retention (Sinclair and Crawford 2005). European green crab (Carcinus maenas), the golden It is possible that hypoxia could drive southern star tunicate (Botryllus schlosseri), and the violet groundfish communities poleward much faster than tunicate (Botrylloides violaceus) have invaded Van- warming. Fisheries managers in Alaska could errone- couver Island coastlines and are expected to continue ously assume stocks are doing well due to biomasses expanding their range throughout Pacific Canada that are stabilized by poleward range shifts from the (Epelbaum et al. 2009; Kelley et al. 2011). Other south. Establishing quotas in such a circumstance impacts of climate change include changing zonation could lead to overfishing and stock collapse when patterns of seagrasses and salt marshes, possible migration slows or stops. facilitation of harmful algal blooms, and range expan- sions of other taxa (Lo et al. 2010). Increased occurrence and establishment of new species Changes in favorable conditions and biodiversity

Pacific Canada’s relatively rich fauna and flora within Patterns of primary production and toxic algal blooms certain taxa, e.g. sea stars (Asteriodea) (Lamb and may change due to changes in oceanography and Hanby 2005), compared with similar latitudes, may be nutrient availability, with potential increases in pro- explained by high productivity and habitat diversity, ductivity due to increased upwelling in the region (e.g. including the influences of the dynamic transition Irvine and Crawford 2013). However, uncertainty of zone. It thus seems reasonable to postulate that an future predictions about productivity in this region is increasingly dynamic transition zone in the future high since there are also a variety of reasons that might increase biodiversity by effectively increasing productivity might decrease in at least some areas of 123 Rev Fish Biol Fisheries (2014) 24:519–559 533

Canada’s Pacific, including increased silt inputs from oxygen, and other habitat factors (Ianson 2008; Perry erosion, decreased nitrate supply if denitrification et al. 2007; Whitney et al. 2007) combine with the effects increases, decreased energy transfer to higher trophic of temperature changes to influence species distribution, levels if phytoplankton size decreases, etc. vigor, and presence, it is not clear that future temperature- In contrast, ocean acidification is likely to have driven range shifts in marine ecosystems would generally disproportionately large effects on Northeast Pacific increase biodiversity. coastal marine ecosystems due to upwelling of It is equally plausible that all the above factors increasingly corrosive waters there, leading to serious together will have increasingly negative consequences physiological impacts on calcifying biota and other for biodiversity in Canada’s Pacific marine ecosys- pH-sensitive biota in Canada’s Pacific (e.g. Ianson tems, especially when combined with non-climate 2008). There are also indications that the physiological stressors such as fisheries and aquaculture (e.g. effects of increased CO2 concentrations (= lowered Hutchings et al. 2012), forestry, pollution, and all of pH) are more pronounced for invertebrates than for the other non-climate anthropogenic stressors of the fish (Harley et al. 2006), though the mechanistic coastal marine ecosystems of this region (e.g. Ban understanding of the physiological effects of CO2 on et al. 2010). This is because climate change factors ocean biota is incomplete (Portner et al. 2005). Also, would tend to shift species away from the functional deoxygenation will constrain the movements and space that they adapted to within the multidimensional generally decrease habitable space for marine organ- hypervolumes of their resource axes (Hutchinson isms, as discussed previously. Such range shifts may 1957; Okey 2003), or the shape of those functional be vertical, forcing deep-water-adapted species to spaces would change, for instance elongating and shallower waters, where they may be more vulnerable thinning. Canada’s marine biodiversity is thought to to both natural and fishing mortality. There could be be on the verge of a biodiversity crisis due to a variety horizontal displacements in fish due to hypoxia/ of local and regional stressors, and climate change will acidification, resulting in migration towards Alaska. likely exacerbate this crisis (Archambault et al. 2010). Other climate-related factors that will also variably affect biodiversity in Canada’s Pacific include highly Changes due to interactions with other stressors textured changes in temperature and salinity (Perry et al. 2007), an increased prevalence or virulence of The combined or cumulative effects of multiple marine diseases (Ford 1996; Harvell et al. 2002; stressors, both climate-related and otherwise, reduces Moore et al. 2000; van Blaricom et al. 1993), toxic the resilience of biological communities to climate algal blooms (Mudie et al. 2002), invasive species change impacts (e.g. Holling 1973). Initial ecosystem (e.g. Epelbaum et al. 2009; Kelley et al. 2011), modeling efforts have indicated that the combined increased exposure and sensitivities to other toxic effects of multiple climate related changes and substances (Coelho et al. 2013; Hallegraeff 2010), fisheries will be greater than the sum of the effects changes in storm regimes (Easterling et al. 2000; of each separately (e.g. Ainsworth et al. 2011), but Gemmrich et al. 2011; Ruggiero et al. 2010; Ulbrich modeling that incorporates broader suites of climate et al. 2008), and regional and local changes in sea level and non-climate stressors in this region are yet to (Mazzotti et al. 2009; Thomson et al. 2008). emerge. Spatial (habitat-based) assessments of vul- As species generally shift poleward with warming nerability to climate change (Okey et al. unpublished conditions, biodiversity would also increase if the data), which incorporate the cumulative impacts of ‘trailing edge’ of a species’ range shift is slower or less stressors (Ban et al. 2010) can be used as a screening- responsive than the ‘leading edge’, as in terrestrial level approach for prioritizing limited resources. ecosystems. One recent meta-analysis of distribution Commercially exploited species tend to be top shifts in marine ecosystems (Sunday et al. 2012) indicated predators in their communities. Removal of top that trailing-edges of species distributions appear to be predators reduces the local species richness, simplifies generally as responsive as leading-edges whereas another the structure and trophic relations of the community, indicated that trailing edges are shifting much slower than and leads to an increase in shorter-lived prey popu- leading edges (Poloczanska et al. 2013). Given that lations. Exploitation of forage fishes (e.g. herring) also factors such as decreases in pH, salinity, dissolved reduces top predators by removing available 123 534 Rev Fish Biol Fisheries (2014) 24:519–559 production. Both types of fisheries tend to make the variables and stressors. In general, the warming along community more vulnerable to environmental vari- the coast and intensification of the Aleutian Low ability, including climate change (Perry et al. 2007). associated with the warm (positive) phase of the PDO Community response will be largely determined by (Hare and Mantua 2000; Hare et al. 1999; Mantua et al. the impacts on key species (Sanford 1999; Schiel et al. 1997) tends to reduce the fisheries landings in southern 2004), taxa, or habitats. These key components must British Columbia and the U.S. Pacific northwest (in be identified and closely monitored. Examples of key the California Current), while increasing landings species in Canada’s Pacific areas include subtidal from northern British Columbia and Alaska (Beamish kelps, eelgrass, and cold water corals and sponges et al. 2009). (biogenic habitat forming components), the purple sea Sensitivities of fish species to climate change can be star and other predators from salmon to seabirds, masked by their life-history characteristics. Shorter- orcas, pinnipeds (predation pressure), forage fishes, lived species are known as sensitive sentinels for shifting benthic invertebrates, and plankton such as copepods climate and oceanographic conditions (Chavez et al. and krill (prey availability). New approaches are 2003) as they respond to short-term variability, but they available to rank species importance in whole ecosys- also have inherently high capacities to adapt. Longer- tems (Jorda´n et al. 2008; Okey 2004a) with respect to lived species are more resistant to short-term climate interaction strength and keystoneness, and this could variability because they can afford long periods of low or allow conservation prioritization. no recruitment, but these species may be more vulner- Despite the importance and practical efficiency of able to longer-term directional change in a given location focusing on key species, marine and coastal biological because they cannot adapt as fast as shorter-lived communities with stable, genetically diverse popula- species. Thus, sablefish, which can live to 100 years, tions and higher biodiversity are more resistant to will have more resistance and thus a greater lag time environmental changes and hence should be more before the impacts of climate change manifest (Beamish resilient to the impacts of global warming (Duffy and et al. 2009), though they will be less capable, ultimately, Stachowicz 2006; Ehlers et al. 2008; Hughes et al. of coping with changed conditions. Warming may result 2003; Steneck et al. 2002). Adaptation strategies in a shift in the makeup of fishery resources in the area should thus promote, protect, and restore biodiversity west of Vancouver Island, including declines in the in general (e.g. Wilson 1992, 2002). availability of salmon, herring, and resident hake and There are many examples ofclimate interactions with increases in the importance of migratory hake, mackerel, non-extractive stressors as well. One of many is the and tuna (Phillips et al. 2007; Robinson and Ware 1999; interaction of climate change with the effects of Walker and Sydneysmith 2007; Ware and Thomson anthropogenic pollution (Coelho et al. 2013) and its 2000; Wright et al. 2005). exacerbation of harmful algal blooms (Hallegraeff Fishing frequently results in the selective removal of 2010). Also, such interactive effects operate multi- older and larger individuals with effects including a directionally. Coastal marine and aquatic ecosystems shortening of the spawning season, a decrease of age-at- stressed by climate change will thus be more vulnerable maturity, and shorter generation replacement times to the effects of local and regional non-climate stressors. (Jennings et al. 1999; Pauly et al. 2001). All these factors tend to make the exploited population less resilient and more sensitive to disturbances and changes in climate, Sensitivities and responses of key taxa and species and visa versa (Perry et al. 2007, 2010). Global bioclimatic envelope modelling was The effects of climate cycles and variations in the recently conducted for commercial fish species (Che- northeast Pacific is a useful starting point for under- ung et al. 2009), and this approach has now been standing the potential impacts of climate change, but applied at the scale of the Northeast Pacific shelf seas surprises are inevitable (Doak et al. 2008; NRC 2002) (Cheung et al. in press) focusing on the 28 pelagic because long-term climate change will shift ocean nekton species that are captured in pelagic trawl climate conditions beyond historical boundaries of monitoring there (Brodeur et al. 2003, 2005; Harding variability, and because effects are multidimensional et al. 2011; Orsi et al. 2007), as discussed previously. and interactive with both climate and non-climate Earlier estimates from these modelling outputs were 123 Rev Fish Biol Fisheries (2014) 24:519–559 535

Table 1 Summary of the summary by Beamish et al. (2009) species and other species and taxa in our review of potential climate impacts on B.C. fisheries within 50 years (Table 2). Species Potential impact

Sablefish Stocks in the south may be reduced, but the Climate change effects in B.C. marine ecosections northern stock in Canada’s Pacific may benefit. Overfishing may mediate climate effects We summarized some potential manifestations of Pacific Stocks in the Strait of Georgia should remain climate change in British Columbia’s 12 marine herring high, but offshore stocks may decline ecosections (Table 3). Additional descriptions of the Pacific hake The Strait of Georgia stock should remain potential sensitivities of British Columbia ecosections high. The offshore stock may also if not are provided in Okey et al. (2012). Although, both of overfished. Range could extend northward, to Haida Gwaii these summaries inevitably miss many subtle and even Pacific halibut The abundance within the population should conspicuous implications of climate change sensitiv- remain at high levels as a consequence of a ities and effects within whole ecosections, it is a useful stormier North Pacific in the winter. The starting point for developing a prioritization of ana- abundance in Canada’s Pacific may be lytical or management resources for understanding reduced slightly as fewer juveniles migrate south into Canada and planning for climate change in Pacific Canada. Pacific ocean The major impact will be increases in the A companion paper (Okey et al. unpublished data) perch frequency of strong year classes, which will provides an example of spatial mapping of the improve abundance vulnerability of Canada’s Pacific marine ecosystems Pacific Pacific sardine will generally increase in to climate change by combining estimates of the sardine Pacific Canada, with continuing natural sensitivity of habitats to climate change variables with fluctuations will continue a spatial analyses of exposure to climate variables and Pacific cod Pacific cod will gradually disappear from the then with spatially mapped cumulative impacts of Strait of Georgia and off the west coast of Vancouver Island as bottom temperatures non-climate stressors, used as an inverse proxy for warm adaptive capacity. Vulnerability estimates are sum- Pacific All Fraser River stocks will decline, with marized by ecosection, by habitat type, and at finer salmon sockeye, pink, and chum declining more resolution. That analysis complements and builds on than coho and chinook. Skeena and Nass the foundation of basic information provided here to Rivers salmon stocks to the north will increase due to improved ocean set the stage for prioritization, planning, and adapta- productivity. Pacific salmon will begin tion strategy development. Okey et al. (2012) provide reproducing in Arctic rivers descriptions of sensitive characteristics of Canada’s Pacific marine habitats. incorporated into simulations by Ainsworth et al. (2011) to explore the combined ecological impacts of Summary and recommendations different climate variables, fisheries, and indirect effects on Northeast Pacific biological communities, The geophysical, oceanographic, and hydrological based on further reviews of the sensitivities of species setting of Canada’s Pacific region has produced an and functional groups to different climate variables. ecological context with highly complex and variable These efforts together advance the articulated vision texture, dynamics, and multidimensionality. The biota of integrated ecological modelling of climate impacts in this region has naturally developed particular on marine ecosystems (Hollowed et al. 2009, 2011; combinations of resiliencies and responsiveness to Okey et al. 2008), here for the Northeast Pacific natural fluctuations and variability, but it is too early to Ocean. accurately characterize how whole biological com- Beamish et al. (2009) reviewed the impacts of munities will respond to long-term changes in climate climate change on British Columbia’s commercial fish stressors beyond the bounds of natural variability. stocks. We have summarized their summary in Local and regional manifestations of climate change, Table 1. We examined these commercially important or global change, are themselves multidimensional, 123 536 Rev Fish Biol Fisheries (2014) 24:519–559

Table 2 Evidence of sensitivities and responses of key taxa and species to climate change in Canada’s Pacific marine ecosystems Observed or predicted responses References

Phytoplankton Warm conditions reduce primary production, and cold Beamish et al. (2009), Goes et al. (2001), Kudela et al. (2006), conditions increase it Mackas et al. (2007), Mackas et al. (2006), Mundy et al. (2010), Ware and Thomson (2005), Whitney et al. (1998), Wong et al. (1998) Spring bloom also responsive to variations in warm/cold Mundy et al. (2010) conditions Phytoplankton assemblages will shift toward warmer water Hare et al. (2007), Richardson (2008) forms Macrophytes A variety of effects are expected on seagrasses and Harley et al. (2012), Koch et al. (2013), Lucas et al. (2007), macroalgae, but uncertainty is high Okey et al. (2006), Poloczanska (2006), Poloczanska et al. (2007) Increased ocean carbon concentrations may facilitate Harley et al. (2006) macrophytes Calcareous macroalgae may be replaced by fleshy macroalgae Koch et al. (2013) Zooplankton Responsive to the cycles of cool-water/high production versus Mackas et al. (2007), Mundy et al. (2010) warm-water/low production Biomass decreased with shift toward a positive PDO in the Beamish et al. (2009), Ware and Thomson (2005) mid-1970 s Seasonal peaks of Neocalanus cristatus and N. plumchrus are Batten et al. (2010), Batten and Mackas (2009), Bertram earlier and shorter during warm regimes and later during cold (2001), Bograd et al. (2010), Mackas et al. (1998) regimes Timing of copepod seasonal migration shifts in response to Batten et al. (2010), Mackas et al. (1998), Mackas and Tsuda ocean regime shifts (1999) Poleward shift in many species Archambault et al. (2010), Batten et al. (2010), Mackas et al. (2007), Richardson (2008) Many subtropical mesozooplankton (copepods) in 2008 Bograd et al. (2010) Euphausiid, chaetognath, hyperiid, and thaliacean species Lavaniegos and Ohman (2003, 2007), Mackas and Galbraith expand and contract their ranges with climate fluctuations (2002) Warm El Nin˜o conditions lowers krill, Euphasia pacifica, DFO (2006b) abundance Strait of Georgia zooplankton communities are strongly Li et al. (2013) correlated with climate forcing indices and weakly correlated with local factors Shrimp and prawns Warm conditions from 2003 to 2005 allowed smooth pink DFO (2008b) (J. Boutillier, Fisheries and Oceans Canada, pers. shrimp (Pandalus jordani), a southern species, to expand comm., 7 April 2009) northward into Pacific Canada and occupy part of the range of its northern counterpart, P. borealis Bivalves Expected to be strongly affected by acidification DFO (2008b), Wootton et al. (2008) Northern abalone (Haliotis kamtschatkana) Sensitive to acidification in addition to emerging diseases, Jamieson (2001), Moore et al. (2000), Rogers-Bennett (2007b), historical overfishing and poaching, and changes in food, Tomascik and Holmes (2003), van Blaricom et al. (1993), habitats, and predators Wallace (1999), Zhang et al. (2007)

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Table 2 continued Observed or predicted responses References

Commercial squids and octopods Landings of L. opalescens increased in northern British DFO (2006a), Pellegrin et al. (2007) Columbia during the late 1990 s for unclear reasons Climate driven changes in commercial squid populations may Pecl and Jackson (2008) have large ecosystem effects Crabs Recruitment of tanner crabs is known to be sensitive to climate Lucas and Jamieson (2007), Zheng and Kruise (2006) fluctuations, and their continental slope distribution expose them to acidification and deoxygenation Dungeness crab are vulnerable to climate-related changes in Yunker and Cretney (1995) currents, waves, precipitation, temperature, runoff, and non- climate stressors Sea cucumbers and sea urchins Sea cucumbers (holothurians) and sea urchins (echinoids) are Doney et al. (2009) calcifying organisms, and thus are vulnerable to acidification Pacific salmon (Oncorhynchus spp.) Canada’s Pacific salmon are influenced by changes in climate e.g. Salmon (1997) and oceanography across the whole North Pacific Changes in climate and oceanography influence biological Brodeur and Ware (1992), Roemmich and McGowan (1995), productivity Venrick et al. (1987) Salmon are responsive to changes and variations in biological Beamish (1995), Beamish and Bouillon (1993), Beamish et al. production (1999a), Beamish et al. (1999b), Mantua et al. (1997), McFarlane et al. (2000), Zabel et al. (2006) Salmon ranges and distributions are contracting and shifting Abdul-Aziz et al. (2011), Battin et al. (2007), BCME (2007), northward due to changes in climate, oceanography, and BCMWALP (2002), Beamish et al. (1997), Brodeur et al. productivity (2006), Crozier et al. (2008), Lackey et al. (2006) A variety of non-climate stressors affect salmon including Battin et al. (2007), Nehlsen et al. (1991), Slaney et al. (1996), overfishing and habitat degradation, increased aquaculture, Krkosek et al. (2007), Noakes et al. (2000), Saksida (2006), possible changes in depredation, and even increased hatchery Ford et al. (1998), Okey et al. (2007), Preikshot (2007), production Williams et al. (2010), Hilborn (1992), NRC (1996), Waples (1999) The Pacific Decadal Oscillation (PDO) corresponds to Mantua et al. (1997), Hare and Mantua (2000), Beamish and dramatic shifts in salmon productivity regimes with higher Bouillon (1993) catches of chum, pink, coho, and sockeye in Canada’s Pacific and Alaska The survival of pink, chum, and sockeye salmon decreased Hare et al. (1999), Mueter et al. (2002) with increasing ocean temperatures in Pacific Canada and Washington State while increasing with increasing temperatures in Alaska—showing opposite effects in Northern and Southern areas Rivers and lakes will warm faster than the ocean, and this may Battin et al. (2007), BCME (2007), Beamish et al. (2009), strongly affect salmon reproduction and survival in Canada’s IPCC (2007), McKinnell (2008) Pacific region High uncertainty of climate effects on Pacific salmon relates to Martins et al. (2012) the multidimensionality of climate impacts on them and the range of sensitivities by life stage and habitat Arrowtooth flounder (Atheresthes stomias) Arrowtooth flounder has increased its biomass considerably Anderson and Piatt (1999), Hunt et al. (2002), Spencer (2008), since the mid-1970 s in the Gulf of Alaska, while forage Wilderbuer et al. (2002) fishes have declined, resulting from a shift to a warmer regime

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Table 2 continued Observed or predicted responses References

Sablefish (Anoplopoma fimbria) Canada’s Pacific stocks favored by positive PDO conditions King et al. (2000) California stocks unfavored by positive PDO Schirripa (2008) Recent overfishing reduces resilience Beamish et al. (2009) Pacific hake (Merluccius productus) The migratory stock and the spawning area expanded Beamish et al. (2009), Benson et al. (2002), Brodeur et al. northward rapidly starting in the mid-1990 s, resulting in a (2006), McFarlane et al. (2000), Phillips et al. (2007) higher proportion of the stock in Canadian waters, but this diminished in recent years Production of the Strait of Georgia population has increased Beamish et al. (2009) with rising local temperatures The biomass index of Pacific hake has declined since 1987, Beamish et al. (2009) possibly due to overfishing Walleye Pollock (Theragra chalcogramma) Sensitive to climate fluctuations due to effects on interactions Anderson and Piatt (1999), Ciannelli et al. (2005), Criddle of recruitment dynamics, predation, and cannibalism et al. (1998), Stabeno et al. (1995), Wespestad et al. (2000) Pacific cod (Gadus macrocephalus) Future warming may eliminate populations in the Strait of Fu and Beamish (2008) Georgia and off the west coast of Vancouver Island Good climate change indicator, but confounded by depletion Beamish et al. (2009) by overfishing and low recruitment Pacific Ocean perch (Sebastes alutus) Recruitment favored during positive PDO conditions such as Schnute et al. (2001) the 1977–1988 warm regime Lingcod (Ophiodon elongatus) Recruitment influenced by ocean climate variability Haggarty et al. (2004) Pacific halibut (Hippoglossus stenolepis) Recruitment favoured by positive PDO conditions and by Beamish et al. (2009) general warming, but warming might prevent migration southward from Alaska Interannual and decadal-scale variability of ocean conditions is Clark and Hare (2002) the major source of recruitment variability Sole Positively affected by higher winter and summer temperatures Teal et al. (2008) in the North Sea, but further temperature increases could harm nurseries Reduced precipitation has negatively affected sole in Portugal Vinagre et al. (2007) Pacific herring (Clupea pallasii) Strongly influenced by climate-ocean fluctuations and Ware (1991) oscillations with cooler water leading to higher abundances Warm ocean water associated with poor recruitment Schweigert (2007) Overfished in Canada’s Pacific during a period of climate- Hourston and Haegele (1980), Schweigert et al. (2010) related poor recruitment, and remains depleted Stocks on the west coast of Vancouver Island and the Northern Boldt et al. (2013), Schweigert et al. (2010) Shelf remain at very low levels Outer coast stocks may move north and be replaced by sardines McFarlane et al. (2005), Schweigert et al. (2010) from the south with long term warming May be impacted by increased predation with recent increases Beamish et al. (2009) in Pacific hake in Canada’s Pacific waters

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Table 2 continued Observed or predicted responses References

Pacific sardine (Sardinops sagax) The famous collapse of the Pacific sardine fishery in the mid- Clark and Marr (1955), Jacobson and Maccall (1995), MacCall twentieth century was likely due to the combined effects of (1979), Ware and Thomson (1991) climate-ocean variability and overfishing Large-scale ocean-climate conditions strongly influence Pacific Kawasaki and Omori (1986) sardine distributions and abundances through their food More positive PDO states and warmer coastal waters generally Beamish et al. (2009), Jacobson and Maccall (1995), increase sardine populations in Canada’s Pacific through both McFarlane et al. (2005), Zwolinski et al. (2012) increased production and migration Sardines were widespread throughout the southern Hecate Flostrand et al. (2013) Strait and Queen Charlotte Sound in 2006 and 2007, thanks to prior warm conditions and a very strong 2003 year-class, but they have declined since with the arrival of cool conditions Eulachon (Thaleichthys pacificus) Sensitive to climate-related changes such as shifts in the timing Schweigert (2007) of spring freshet events, which affect their spawning Mortality associated with climate changes may limit recovery, Schweigert et al. (2012) along with fishing and marine predation Long-term declines from California to Alaska over the past Beacham et al. (2005), Schweigert et al. (2013) 20 years, particularly in Pacific Canada’s central and southern areas where abundance remains low Anomalous fish and squid occurrences Humboldt squid (Dosidicus gigas) increased in abundance in Brodeur et al. (2006), Cosgrove (2005), DFO (2008b), Field the California Current, including the southern section of et al. (2007), Trudel et al. (2006) Canada’s Pacific waters, between 2002 and 2006 D. gigas may have extended its range northward in response to Watters et al. (2008) changes in temperature, and such squid appear more strongly controlled by bottom-up forcing than by predators Pacific mackerel (Scomber japonicas) invaded Canada’s Hargreaves and Hungar (1995), Trudel et al. (2006), Ware and Pacific in the early 1990s and in 2005 and may have Hargreaves (1993) contributed to the very poor year-classes of salmon stocks Pacific jack mackerel (Trachurus symmetricus) also shifted Brodeur et al. (2006), Orsi et al. (2007) *1,600 km northward from southern California to the Oregon-B.C. coastline during the warm summers of 2004 and 2005 Ocean sunfish (Mola mola) were particularly abundant from at Brodeur et al. (2006), Trudel et al. (2006) R. Williams, least Oregon through British Columbia during the warm unpublished data years of 2004 and 2005 with a concentration in northern Queen Charlotte Sound and the adjacent continental slope Other southern and oceanic species - Pacific butterfish Brodeur et al. (2006), Trudel et al. (2006), Wing (2006) (Peprilus simillimus), Pacific pomfret (Brama japonicus), yellowtail (Seriola lalandi), and opah (Lampris guttatus) were all unusually abundant in the region in warm 2005 Salmon shark (Lamna ditropis) have developed a summertime Healey and Hennessey (1998), Okey et al. (2007), Thomson concentration in the Queen Charlotte Sound, possibly due to et al. (1994), Williams et al. (2010) returning salmon taking the northern route past Vancouver Island more consistently due to warming Anomalous occurrences of many species were documented in Fulton (1985) Canada’s Pacific during the 1982–1983 El Nin˜o event

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Table 2 continued Observed or predicted responses References

Marine and coastal birds Warming conditions and associated shifts in plankton Atkinson et al. (2004), Frederiksen et al. (2006), Napp and communities adversely affect seabirds and other higher Hunt Jr. (2001), Parrish et al. (2007), Peery et al. (2007), trophic level species Springer et al. (2007) Cassin’s auklet (Ptychoramphus aleuticus) survival at Triangle Bertram (2001), Bertram et al. (2005), Hipfner (2008) Island declined with increased ocean temperature that led to a mismatch between the timing of breeding and their main prey, the copepod Neocalanus cristatus Rhinoceros auklet (Cerorhinca monocerata) were similarly Bertram et al. (2001), Hedd et al. (2006) affected by declines in their fish prey with increasing SST Brown Pelican (Pelecanus occidentalis) invaded Clayoquot Palm (2006) (R. Palm, personal communication, 21 August Sound on the west coast of Vancouver Island and increased 2013) there during the last warm phase of the PDO in the mid- 2000 s, but virtually disappeared there after the 2007 shift back to cool Marine mammals Impacts of climate change to marine mammals are indirect, Learmonth et al. (2006), Simmonds and Isaac (2007) mainly through changes in prey communities Marine mammal population declines in the Northeastern Francis et al. (1998), Schell (2000) Pacific have been linked to decreases in primary production Other indirect effects of climate change include susceptibility Gilmartin and Forcada (2002), Lahaye et al. (2007), Learmonth to disease and contaminants and increased competition et al. (2006) Long-beaked Common Dolphins (Delphinus capensis) were Ford (2005) sighted in Canadian waters for the first time during 1993–2003, during warm-water oceanographic events Grey whales (Eschrichtius robustus) may encounter less Bluhm and Gradinger (2008) benthic food due to increases in river runoff and turbidity Killer whales (Orcinus orca) in Canada’s Pacific are sensitive Lusseau et al. (2004) to large-scale variations in ocean climate Reductions in quantity and quality of salmonids will adversely DFO (2008a), Ford (2006), Ford et al. (2005) affect Pacific Canada’s resident killer whale populations Pinnipeds—The decrease in production brought about by El Crocker et al. (2006), Le Boeuf and Crocker (2005), Nin˜oconditions disperses prey and adversely affects pinniped Learmonth et al. (2006), Melin et al. (2012) populations mainly through lowered reproductive success Sea otters (Enhydra lutris) populations are likely shaped Byrnes et al. (2011), Koch et al. (2013), Wootton et al. (2008) mainly by prey availability, and climate change can affect kelp forest communities and their non-kelp forest prey resources E. lutris can strongly mediate or cascade climate change effects Paine (1969, 1995) given its disproportionately strong keystone role Sea turtles Increased temperatures will cause poleward shifts in species Booth and Astill (2001), Glen and Mrosovsky (2004), foraging ranges, earlier breeding, changes in food MacLeod et al. (2005), McMahon and Hays (2006), availability, and sex ratios skewed toward females. Warmer Mrosovsky et al. (1999), Poloczanska et al. (2007), temperatures will bring more sea turtles to Pacific Canada, Poloczanska et al. (2009), Robinson et al. (2009), but higher overall turtle mortality might be expected with Weishampel et al. (2004), Yntema and Mrosovsky (1982) changes in the variability of temperature and productivity, and eventual inundation of nesting beaches

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Table 3 Summary of some potential climate change manifestations in British Columbia marine ecosections, added to previous ecosection descriptions (BC 2002), organized by biogeographic unit (DFO 2009) Marine Physiographic Oceanographic Biological features Boundary rationale Potential climate ecosection features features change manifestations

Northern Shelf Dixon Across-shelf trough Strong freshwater Mixture of neritic Distinguished from Increased runoff and Entrance with depths mostly influence from and sub-polar area to south by stratification. \300 m; mainland river plankton species; strong freshwater Reduced salinity surrounded by low- runoff drives north- migratory corridor discharge influence and associated lying coastal plains westward flowing for Pacific salmon; changes to (Hecate coastal buoyancy some productive buoyancy flow. Depression) current and and protected areas Ocean warming. estuarine-like for juvenile fishes Sea-level rise circulation and invertebrates Hecate Very shallow strait Semi-protected Neritic plankton Marine in nature but Ocean warming. Strait dominated by waters with strong communities much shallower, Sea-level rise. coarse bottom tidal currents that w/oceanic with associated Changes in runoff sediments; promote mixing; intrusion; salmon greater mixing than and salinity surrounding coastal dominantly and herring nursery areas to the south lowlands ‘‘marine’’ waters area; abundant benthic invertebrates; forage for marine mammals and birds North Coast Deep, narrow fjords Very protected Low species diversity Unique physiography Ocean and Fjords cutting into high waters with and productivity and stratification continental coastal relief restricted due to poor water compared with warming. Increase circulation; often exchange and bordering regions in runoff. Decrease strongly stratified nutrient depletion; in salinity. unique species Increased assemblages in stratification; benthic and intensified anoxia plankton communities Queen Wide shelf Ocean wave Mixture of neritic More oceanic (deep) Oceanic warming. Charlotte characterized by exposures with and oceanic and marine than Intrusion of low Sound several large banks depths plankton Vancouver Island anoxic and acidic and inter-bank mostly [ 200 m communities; Shelf and Hecate waters into channels and dominated by northern limit for Strait canyons oceanic water many temperate intrusions fish species Queen Predominantly High current and Very important for More marine than Ocean and Charlotte shallow (\200 m); high relief area; marine mammals; Johnstone Strait; continental Strait high relief area very well mixed; migratory corridor much more shallow warming. Increase with deeper fjord moderate to high for anadromous with high relief and in runoff. Decrease areas salinities with some fishes; moderate high currents than in salinity freshwater inputs in shellfish habitat Queen Charlotte the inlets and fjords Sound Johnstone Narrow, constricted Protected coastal Migratory corridor Johnstone Strait has Changes in currents, Strait channels waters with strong for anadromous greater mixing and temperature, currents; well fishes; rich sessile, more channels than salinity, and mixed; poorly hard substrate areas to south; productivity stratified invertebrate Queen Charlotte community; diverse Strait more marine species assemblage of benthic fishes

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Table 3 continued Marine Physiographic Oceanographic Biological features Boundary rationale Potential climate ecosection features features change manifestations

Continental Steep sloping shelf Strong across slope Upwelling zone; Transitional area Acidification and slope and downslope; productive coastal between anoxia in the deep turbidity; currents plankton continental slope layers. Ocean communities and and abyssal plane warming. Changes unique assemblages to ocean currents of benthic species Southern Shelf Vancouver Narrow, gently Open coast with Highly productive More open shelf than Oceanic warming. Island sloping shelf oceanic wave with neritic Juan de Fuca Strait; Changes to Shelf exposures; plankton more freshwater nearshore northward, coast- community; influence (coastal buoyancy-driven hugging buoyancy northern limit for buoyancy current) flow and offshore current due to hake, sardine, than Queen ocean circulation freshwater northern anchovy, Charlotte Sound influence; seasonal and Pacific upwelling at outer mackerel; margin productive benthic community; rich fishing grounds for benthic fish and invertebrates Juan de Deep trough; major Semi-protected Migratory corridor Much more marine Changes in currents, Fuca Strait structural feature coastal waters with for anadromous than Strait of temperature, accentuated by strong ‘‘estuarine- fish; moderately Georgia; less ‘‘open productivity, glacial scour like’’ outflow productive; mixture shelf’’ than salinity, oxygen, current (coast- of neritic and Vancouver Island and acidity hugging buoyancy oceanic plankton Shelf current to north); species major water exchange conduit with ‘‘inland sea’’ Continental Steep sloping shelf Strong across slope Upwelling zone; Transitional area Acidification and Slope and downslope; productive coastal between anoxia in deep turbidity; currents plankton continental slope layers. Ocean communities and and abyssal plane warming. Changes unique assemblages to ocean currents of benthic species Strait of Georgia Strait of Broad shallow basin Protected coastal Nursery area for Stronger Fraser River Increased Georgia surrounded by waters with salmon, herring; signature than areas temperature. coastal lowlands significant abundant shellfish to north or west Changes in runoff (Georgia freshwater input; habitat; neritic and currents. Depression) high turbidity and plankton Acidification and seasonally community dissolved oxygen stratified; very warm in summer

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Table 3 continued Marine Physiographic Oceanographic Biological features Boundary rationale Potential climate ecosection features features change manifestations

Offshore Pacific Subarctic Includes abyssal Eastward-flowing Summer feeding Northern and western Freshening of Pacific plain and subarctic current ground for Pacific boundaries surface waters, continental rise; bifurcates at coast salmon stocks; undefined; eastern increased major fault occurs with northerly abundance of boundary stratification, along west margin; flowing Alaska pomfret, Pacific coincident with decreased oxygen seamount chain Current; current saury, albacore shelf break; concentrations, trends NW/SE flow generally tuna, and jack southern boundary change in northward mackerel in indistinct productivity throughout year summer; boreal plankton community Transitional Includes abyssal Area of variable Transition zone Northern boundary Freshening of Pacific plain, and currents; southerly between southerly, indistinct and surface waters. continental rise; areas may be temperate, and approximately Increased also includes affected by northerly boreal coincident with stratification. spreading ridges, southward-flowing plankton southern limit of Decreased oxygen transform faults, California Current communities; Alaskan Current concentrations. triple junction, and in summer, but mixing of oceanic (winter); eastern Change in plate subduction remainder of area and coastal boundary at shelf productivity zone characterized by plankton break. Western weak and variable communities boundary undefined currents; Davidson adjacent to coastal Current along shelf shelf edge flows north in winter, south in summer and the responses of species to each climate dimension that relate to the complexities of Canada’s Pacific are differential. As these multiple dimensions of long- marine ecosystems in both time and space. The present term change emerge from the natural or background assessment, as with other regional assessments (Hob- bounds of variability, varied responses among species day et al. 2006; Poloczanska et al. 2007) demonstrates will lead to mis-matches of co-evolved species the utility of regional summaries of global change (Gaston et al. 2009; Hipfner 2008), shuffling of effects as a starting point for the development of species assemblages (Darling et al. 2013; Hobday climate change prioritization, planning, policies, and 2011) including invasions of new species, non-linear adaptation strategies in this region, and their imple- threshold-type community changes and thus dispro- mentation. Such assessments help identify ecological portionately rapid shifts to different community states and social issues and analytical and management with different functions and ecosystem services (Okey capacities requiring further attention or investment. 2004b), and possible reductions in biodiversity. This Additional steps include screening assessments such may affect human society strongly given both direct as spatial mapping of ecological vulnerability (Okey and indirect dependencies on coastal and marine et al. unpublished data) and assessments of socio- resources and functions. economic and cultural vulnerabilities and social The manifestations of global change in a given adaptive capacities (Allison et al. 2009; Cinner et al. region, and the biological and ecological effects of 2009; Hughes et al. 2012; Lin and Morefield 2011; those changes, thus depend on context and scale of McClanahan et al. 2008; Moser et al. 2012). examination. The present assessment uncovered par- All of the biota in Canada’s Pacific marine ecosys- ticular biological responses of key taxa and species tems will be affected by how global changes are

123 544 Rev Fish Biol Fisheries (2014) 24:519–559 manifested in this region. The present analysis sum- effects are likely to occur with the next warm phase marizes available knowledge of responses of many (positive PDO) or El Nin˜o event, or combinations key species and taxa based on our sampling of the thereof. A variety of future changes in precipitation, available literature from this region and the broader hydrology, salinity, and stratification, sea-level, and Northeast Pacific. It builds a deductive synthesis of storminess will also add to the overall novel stress that examples of how climate change is affecting, and may climate change exerts on Canada’s Pacific marine continue to affect, Canada’s Pacific marine ecosys- ecosystems. We refrained from reviewing the physi- tems and biological communities, based on: observa- ological tolerances and sensitivities of biota to the tions of physical, chemical, and biological changes; various climate change stressors, but that body of knowledge of Pacific Canada’s ecological settings; knowledge is developing rapidly, and is critically ecological theory; and results of some initial quanti- important for ultimately projecting or forecasting tative analysis and modelling. It also summarizes responses. potential climate change manifestations by ecosection Given that our examination of changes in climate and habitat. This synthesis allowed us to broadly and and oceanography, and the responses of marine generally identify the potential impacts of climate species, are based on historical observations and changes in Canada’s Pacific, and to begin to identify climate projections to mid-century or the end of the what may be required in terms of adaptive actions or century, we fully expect our resulting summary to policy responses. underestimate the physical, chemical, and biological Our findings were consistent with more general changes that we should expect in the next century and literature about climate change impacts on marine beyond, when there will be considerably larger ecosystems, fish, and fisheries (Brander 2010; Harley consequences of these changes. Furthermore, the et al. 2006; Hobday et al. 2006; Hutchings et al. 2012; climate manifestations and the biological responses IPCC 2007; Perry et al. 2010; Poloczanska et al. 2007; described in this review, even examined in synthesis, Poloczanska et al. 2013), including recent reviews that likely considerably underestimate the overall impacts highlight California Current ecosystems (Doney et al. because although we do suggest that the overall effects 2012) and the North Pacific generally (Hollowed et al. will result from the cumulative impacts of both 2013), but the overall character of climate-related climate-related and non-climate-related stressors, we changes in Canada’s Pacific marine ecosystems are have no way of expressing those overall effects or unique, given its unique setting and biota. Our findings even estimating them in the form of a review—without were also consistent with another literature review of integrative quantitative analysis. climate change effects and adaptation approaches in For millennia, the rich biological communities marine and coastal (terrestrial) ecosystems of the within the coastal marine ecosystems of Canada’s ‘North Pacific Landscape Conservation Cooperative Pacific region have enabled the development of (NPLCC) region’ (extending from northern California likewise rich and highly complex human societies of to Kodiak Island, Alaska) (e.g. Tillmann and Siemann the sovereign first and subsequent nations of Canada’s 2011). Pacific region through intimate and critical social- Acidification and de-oxygenation stand out as the ecological connections. These human communities most prominent and urgent threats to the marine life and broader ecological communities were both shaped and human communities of Canada’s Pacific marine by the particular complexity of the region’s climate, ecosystems. Variations and changes in temperature oceanography, geomorphology, and productivity. All and productivity also emerged as a principal driver of the climate-related and non-climate-related changes shaping and regulating marine life of the region, as it is to the region’s ecology are inexorably connected to associated with strong effects on productivity, shifts in these various nations, communities, and the people species distributions, changes in phenology, and within them, through to the present. Although climate- mismatch of co-evolved species, all of which have related changes to Canada’s Pacific marine ecosys- been documented. These effects on marine life, tems has global significance in various ways, affecting however, may be masked or delayed by the cool people throughout the world, the local and regional regime (negative phase of the PDO) that has predom- implications of climate change effects take precedence inated in Canada’s Pacific since 2007. Sudden, large in our examination, because this is the working and 123 Rev Fish Biol Fisheries (2014) 24:519–559 545 effective scale of manifested impacts and adaptation ecological vulnerabilities to climate change; (5) basic planning. Although collective action on the global biological and social research on tolerances and scale is required to reduce exposure to global stressors, sensitivities of organisms, habitats and human systems the people of Canada’s Pacific can potentially do much to climate-related stressors, and (6) modelling of to manage the impacts of climate-related stressors by climate change and impacts including all needed developing adaptation strategies that adjust the local scales, components, systems, and integration. and regional non-climate stressors, which may Some of these could be incorporated into an overall strongly affect the ecological and social-ecological adaptation framework that could feature a tiered resistance and resilience to climate impacts. approach to assessment and management consisting One person’s stressor is, however, another person’s of (1) routinely including climate change consider- livelihood, and so adaptation strategies to cope with ations into existing coastal and marine resource the effects of climate change on Canada’s Pacific assessment and planning; (2) conducting national marine ecosystems will need to be integrative, opti- and regional assessments of the effects of climate mized, and negotiated, but intergenerational equity change on marine ecosystems (as described in the must be a guiding principle so that the future is not previous paragraph); (3) developing cost-benefit and discounted for only temporary prosperity (Sumaila feasibility analyses to further identify and prioritize and Walters 2005). The inevitable limitations and sets of strategies for management sectors, and (4) failures of these adaptation strategies will also need to developing an integrative and cross-sectoral regional be addressed through precautionary strategies. We climate adaptations implementation plan for the must also meet the challenge of preserving the non- marine and coastal zone including prioritized actions stationary foundations of future ecosystem values and for reducing climate vulnerability, monitoring and services. Even before considering broader values, adaptive feedback, and approaches for internalizing human activities such as fisheries and most other costs and revealing trade-offs among disparate human coastal and marine and freshwater activities will need activities and sectors. It is critical that the objectives of to adjust to non-stationary ecosystems and resources. adaptation strategies and frameworks be identified and The planning of such adaptive shifts should begin clarified. We recommend that adaptation strategies be immediately given the rate of historically observed focused on promoting, protecting, and restoring bio- change and variability in the region and around the diversity in general, as presumably the most certain world. Industries and resource managers dealing with way of banking ecosystem functions and services as high value trade-offs will need continually refined we attempt to cope with non-stationary dynamics and forecasts or other adaptive or dynamic management non-linear changes and surprises. approaches (e.g. Hobday et al. 2011). The strategic protection of key species and habitats From our analysis, we can derive recommendations is a central and key element to climate adaptation that for key analytical capabilities that would provide protects biodiversity, other values, and ecosystem prioritization for assessment and management needs services. In the context of climate change, this related to climate change in Canada’s Pacific. These protection must be approached with the assumption would comprise National and Regional Climate that biological communities are non-stationary, rather Assessments, akin to regional and ocean technical than assuming that some elusive historical baseline or regional reports of the United States National Climate equilibrium can be restored, especially given that Assessment (Griffis and Howard 2013; Markon et al. marine species are moving and shuffling so much faster 2012) and here would include (1) regional summary than terrestrial species (Hobday 2011; Poloczanska assessments of climate change effects, such as the et al. 2013). The goal of such protection should thus present analysis; (2) summaries of effects by mean- relate to biodiversity and ecosystem function rather ingful ecological and social units such as ecosections, than stasis. Natural biological refuges that are less habitats, and institutions; (3) the development of exposed to climate changes, or are somehow buffered ecological and human well-being indicators and from them, may help limit or slow down rapid associated monitoring programs that can detect envi- functional or biotic degradation from climate changes ronmental changes and ecological effects; (4) quanti- and allow for more biological adaption (e.g. Jessen and tative spatial assessments of ecological and social- Patton 2008). Identifying and protecting such natural 123 546 Rev Fish Biol Fisheries (2014) 24:519–559 marine climate refugia is feasible in Canada’s Pacific implemented in a timely manner, in all of our marine given the spatial heterogeneity of exposure to climate and coastal activities. stressors due to the region’s complex geomorphology, That scientific foundation includes the develop- climate, and oceanography. Local and regional non- ment of monitoring programs that can detect climate climate stressors such as fisheries and other marine changes and impacts through the development of uses would be excluded or adjusted in climate refugia useful sets of indicators of the state of ecological and as a precautionary buffer to account for the additional human systems, and of the pressures on them, both stress that climate changes exert on valued or repre- climate and non-climate related (Jessen and Patton sentative ecosystem components. 2008). This includes improving and integrating the The manifested effects of climate change on a monitoring of key climate change variables and species, population, or biological community are indicators at finer temporal and spatial scales, includ- exacerbated, and thus mediated, by non-climate local ing temperature, salinity, pH, dissolved oxygen, as and regional stressors. These regional and local well as key sentinel species and other indicators of stressors decrease the resistance and resilience of biological communities and ecological features and these natural systems to the stress exerted by climate human communities. An expert-based approach to the change variables, and visa-versa. It is the cumulative development of ecological indicators has been devel- impacts of all these stressors that determine the overall oped for Canada’s Pacific marine ecosystems (Okey vigor and health of any given component or the overall et al. unpublished data). ecosystem, and it is often the local and regional We have summarized a broad spectrum of scientific stressors that have stronger adverse effects on the knowledge to gain an overall synthesis of the effects of system than climate change stressors do, at least at the climate change in Canada’s Pacific marine ecosys- present time. An assessment of the effects of climate tems. Our goal was to stimulate and guide the change on a given system, such as Canada’s Pacific, is development and implementation of climate adapta- thus incomplete without accounting for non-climate tion strategies and policy. Science is entering a new anthropogenic stressors (Jessen and Patton 2008). The era of predictive and integrative modelling for cumulative impacts of these non-climate local and addressing climate change impacts, and adaptation regional stressors in Canada’s Pacific have been planning will require such advancement in capability. examined spatially (Ban et al. 2010) and are being This is illustrated by the latest generations of model- incorporated into a spatial vulnerability assessment as ling in the IPCC process (http://www.ipcc.ch/), as well an inverse proxy for the adaptive capacity component of as by advancements in addressing climate change ecological vulnerability to climate change (Okey et al. impacts with trophodynamic modelling (Ainsworth unpublished data). Such vulnerability assessments can et al. 2011) and end-to-end ecosystem models (Rose provide habitat-based or other types of screening-level et al. 2010), and efforts to continue developing inter- assessments to prioritize particular areas or components national collaborations in the science of forecasting for assessment or management attention. Such analyses the effects of climate change on marine species and provide explicit information about adjusting non- ecosystems (Ainsworth et al. 2011; Hollowed et al. climate stressors to increase the adaptive capacity of 2011, 2013). As reflected in these advancements, it is the system to climate change impacts. useful to approach this wicked problem by continuing Although the development of adaptation and man- to develop and refine projections of how individual agement strategies is crucial and should be developed species and communities will respond in the future, immediately, we conclude by emphasizing that scien- but we also must invest in making the transition to tific, local, and traditional knowledge is the foundation projecting the changes in whole biological commu- upon which we improve our understanding of the nities by accounting for cumulative impacts, the wicked problem of global change effects on Canada’s multidimensionality of stressors, differential respon- ecosystems. Smart adaptation actions emerge only ses of species, indirect effects within communities, from such a foundation. Continued development of the and the apparently non-deterministic behaviors within underlying science must occur in tandem with the highly complex and variable marine ecosystems, development of climate adaptation and management especially in complex settings such as Canada’s regimes. Most importantly, such regimes must be Pacific. 123 Rev Fish Biol Fisheries (2014) 24:519–559 547

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