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==== 5.2.4.3 Bathyal Ecosystems (200–3000 m) ==== <div id="section-5-2-4-3bathyal-ecosystems-200-3000-m-block-1"></div> Bathyal ecosystems consist of numerous geomorphic features with steep topography (Figure 5.15). These include continental slopes covering 5.2% of the seafloor, over 9400 steep-sided canyons, and >9000 conical seamounts (submarine volcanos which are mainly inactive), as well as guyots and ridges which together cover ~6% of the seafloor (Harris et al., 2014) ''.'' Seamounts and canyons support high animal densities and biomass including cold water coral, sponge and bryozoan reefs, exhibit high secondary production supported by locally enhanced primary production and intensified water flow, function as diversity hotspots and serve as stepping stones for larval dispersal (Rowden et al., 2010). Canyons transport particulate organic matter, migrating plankton and coarse material from the shelf, and are sites where intensified mixing and advection of water masses occurs (De Leo et al., 2010; Levin and Sibuet, 2012; Fernandez-Arcaya et al., 2017). Slopes, canyons and seamounts exhibit strong vertical temperature, oxygen and pH gradients generating sharp ecological zonation (Levin and Sibuet, 2012), thus changes in exposures are expected to alter the distributions of their communities (Figure 5.15, 5.16 '') (medium confidence). '' In some regions, observational records document changing conditions in bathyal ecosystems (Levin, 2018; Section 5.2.2.4). In the Northeast Pacific continental slopes associated with the California Current ecosystem, observations over the past 25 years show high variability but an overall trend of decreasing ocean oxygen and pH levels with oxygen declines of up to 40% and pH declines of 0.08 units in California . (Goericke et al., 2015) (high agreement, robust evidence, high confidence). Large oxygen declines are linked to past warming events on continental margins, over multiple time scales from 1–100 ky (Dickson et al., 2012; Moffitt et al., 2015). Studies across modern oxygen gradients on slopes reveal that suboxic (5–10 µMol kg -1 O 2 ) values lead to loss of biodiversity of fish (Gallo and Levin, 2016), invertebrates (Levin, 2003; Gallo and Levin, 2016; Sperling et al., 2016), and protozoans (Bernhard and Reimers, 1991; Gooday et al., 2000; Moffitt et al., 2014) (high confidence). Shoaling oxyclines on continental slopes have altered depth distributions of multiple co-occurring echinoid species over the past 25 years (Sato et al., 2017) and can reduce the growth rate, and change the skeletal structure and biochemical composition of a common sea urchin (Sato et al., 2018). In central Pacific oceanic canyons, fish abundance and diversity are reduced at 4 to 5 times higher oxygen concentrations than on continental slopes (<31 µMol kg -1 O 2 ) (De Leo et al., 2012). Low oxygen on continental slopes causes reductions in faunal body size and bioturbation (Diaz and Rosenberg, 1995; Levin, 2003; Middelburg and Levin, 2009; Sturdivant et al., 2012), simplification of trophic structure reducing energy flow to upper trophic levels (Sperling et al., 2013), shifts in carbon processing pathways from metazoans to protozoans (Woulds et al., 2009), and reduced colonisation potential (Levin et al., 2013). These changes are expected to lead to altered ecosystem structure and function, with lower carbon burial (Smith et al., 2000; Levin and Dayton, 2009) (medium confidence). Both carbon sequestration and nitrogen recycling are highly sensitive to small changes in oxygenation within the suboxic zone (Deutsch et al., 2011). Bathyal species adapted to OMZs where CO 2 levels are characteristically high, appear less vulnerable to the negative impacts of ocean acidification (Taylor et al., 2014). Benthic foraminifera, which are often the numerically dominant deep sea taxon, show no significant effect of short-term exposure to ocean acidification on survival of multiple species (Dissard et al., 2010; Haynert et al., 2011; Keul et al., 2013; McIntyre-Wressnig et al., 2014; Wit et al., 2016) and in fact hypoxia in combination with elevated pCO 2 favors survival of some foraminifera (Wit et al., 2016). However, lower pH exacerbates shallow foraminiferal sensitivity to warming (Webster et al., 2016). Limited evidence suggests that combined declines in pH and oxygen may lead to increase in some agglutinating taxa and a decrease in carbonate-producing foraminifera, including those using carbonate cement (van Dijk et al., 2017). Exposure to acidification (0.4 unit pH decrease) reduces fecundity and embryo development rate in a bathyal polychaete. Where both oxygen and CO 2 stress occur together on bathyal slopes, oxygen can be the primary driver of change (Taylor et al. 2014; Sato et al. 2018). Nematodes are sensitive to changes in temperature (Danovaro et al., 2001; Danovaro et al., 2004; Yodnarasri et al., 2008) and elevated CO 2 (Barry et al., 2004; Fleeger et al., 2006; Fleeger et al., 2010). There is low agreement about the direction of meiofaunal responses among studies, reflecting opposing responses in different regions. However, there is high agreement that meiofauna are sensitive to change in environment and food supply ( ''medium confidence'' ). Additional research is needed across all taxa on how hypoxia and pH interact (Gobler and Baumann, 2016). Continental slopes, seamounts and canyons (200–2500 m) are projected to experience significant warming, pH decline, oxygen loss and decline in POC flux by 2081–2100 (compared to 1951–2000) under RCP8.5 (Table 5.5). In contrast, the average changes are projected to be 30–50% less under RCP2.6 (Table 5.5) by 2081-2100. Most ocean regions at bathyal depths (200–2500 m) except the Southern and Arctic Oceans are predicted to experience on average declining export POC flux under RCP8.5 by 2081–2100 (Yool et al., 2017; FAO, 2019) with the largest declines of 0.7–8.1 mg C m -2 d -1 in the Northeast Atlantic (FAO, 2019). There is a strong macroecological relationship between depth, export POC flux, biomass and zonation of macrobenthos on continental slopes (Wei et al., 2011), such that lower POC fluxes will alter seafloor community biomass and structure ( ''medium confidence'' ) (See also Section 5.2.4.1). This is modified on the local scale by near-bottom currents, which alter sediment grain size, food availability, and larval dispersal (Wei et al., 2011). Declines in faunal biomass (6.1 ± 1.6% 95% C.I) are predicted for 96.6 ± 1.2% of seamounts under RCP8.5 by 2091–2100 relative to 2006–2015, driven by a projected 13.8 ± 3.3% drop in POC flux (Jones et al., 2014). The majority (85%) of mapped canyons are projected to experience comparable benthic biomass declines (Jones et al., 2014). By 2100 under RCP8.5, pH reductions exceeding -0.2 pH units are projected in ~ 23% of north Atlantic deep sea canyons and 8% of seamounts (Gehlen et al., 2014), with potential negative consequences for their cold water coral habitats (See Box 5.2). Mean temperature (warming) signals are projected to emerge from background variability before 2040 in canyons of the Antarctic, northwest Atlantic, and South Pacific (FAO, 2019). Enhanced stratification and change in the intensity and frequency of downwelling processes under atmospheric forcing (including storms and density-driven cascading events would alter organic matter transported through canyons (Allen and Durrieu de Madron, 2009) ( ''low confidence'' ). Changes in the quantity and quality of transferred particulate organic matter, as well as physical disturbance during extreme events cause a complex combination of positive and negative impacts at different depths along the canyon floor (Canals et al., 2006; Pusceddu et al., 2010). Canyons and slopes are recognised as hosting many methane seeps and other chemosynthetic habitats (e.g., whale and wood falls) supported by massive transport of terrestrial organic matter (Pruski et al., 2017); their climate vulnerabilities are discussed below. Seamounts have been proposed to serve as refugia for cold water corals facing shoaling aragonite saturation horizons (Tittensor et al., 2011), but could become too warm for deep-water corals in some regions (e.g., projections off Australia) (Thresher et al., 2015) ( ''one study, low confidence'' ). Seamounts are major spawning grounds for fishes; reproduction on seamounts may be disrupted by warming (Henry et al., 2016) ( ''one study, low confidence'' ). In the north Atlantic, models suggest seamounts are an important source of cold water coral larvae that maintain resilience under shifting NAO conditions (Fox et al., 2016), thus loss of suitable seamount habitat may have far-reaching consequences (Gehlen et al., 2014) ( ''low confidence'' ) (also see Box 5.2). <div id="section-5-2-4-3bathyal-ecosystems-200-3000-m-block-2"></div> <span id="table-5.5"></span> <!-- START TABLE --> '''Table 5.5''' Projected climate changes from the present to 2081–2100 given as mean (min, max) at the deep seafloor for continental slopes, canyons, seamounts and cold water corals mapped from 200–2500 m under RCP8.5 and RCP2.6 Projections are based on three 3D, fully coupled earth system models (ESMs) (as part of CMIP5): the Geophysical Fluid Dynamics Laboratory’s ESM 2G (GFDL-ESM-2G); the Institut Pierre Simon Laplace’s CM6-MR (IPSL-CM5A-MR); and (iii) the Max Planck Institute’s ESM-MR (MPI-ESM-MR). Export flux at 100 m was converted to export POC flux at the seafloor (epc) using the Martin curve following the equation: epc = epc100 (depth/ export depth)-0.858. Projections were made onto the (i) slope from a global ocean basin mask from World Ocean Atlas 2013 V2 (NOAA, 2013), (ii) global distribution of submarine canyons with canyon heads shallower than 1500 m (Harris and Whiteway, 2011); (iii) global distribution of seamounts with summits between 200–2500 m (Kim et al. 2011); and (iv) global occurrence of cold water corals between 200–2500 m (Freiwald et al. 2017). <!-- TABLE --> {| class="wikitable" |- | ''' ''' | ''' Temperature ''' ''' (''' '''o''' '''C)''' | ''' pH ''' | '''DO ''' '''(''' ''µ'' '''Mol kg''' '''-1''' ''')''' | '''POC flux''' '''(mgC m''' '''-2''' '''d''' '''-1''' ''')''' |- | ''' ''' | '''RCP2.6''' | ''' RCP2.6''' |- | Continental slopes | +0.30 (–0.44, + 2.30) | –0.06 (–0.19, –0.02) | –3.1 (–49.3, +61.7) | –0.39 (–16.0, +3.9) |- | Canyons | +0.31 (–0.27, +1.76) | -0.05 (-0.13, +0.01) | –3.5 (–44.7, +29.3) | –0.33 (–10.53, +3.53) |- | Seamounts | +0.13 (+0.01, +0.67) | -0.02 (-0.11, +0.005) | –3.46 (–18.9, +4.1) | –0.15 (–2.20, +1.33) |- | Cold water corals | +4.3 (–0.29, +1.85) | -0.07 (-0.13, 0.0) | –3.5 (–25.6, +24.7) | –0.7 (–10.5, +3.4) |- | |- | | '''RCP8.5''' |- | Continental slopes | +0.75 (–8.4, +4.4) | –0.14 (–0.44, –0.02) | –10.2 (–67.8, +53.8) | –0.66 (–33.33, +10.3) |- | Canyons | +0.19 (–0.03, +1.14) | -0.11 (-0.35, +0.02) | –0.8 (–28.8, +10.1) | –0.80 (–28.76, +10.07) |- | Seamounts | +0.66 (–0.75, +3.19) | -0.03 (-0.19, +0.001) | –0.50 (–7.2, +3.0) | –0.50 (–7.18, +2.98) |- | Cold water corals | +0.96 (–0.42, +3.84) | -0.15 (-0.39, +0.001) | –10.6 (–59.2, +11.1) | –1.69 (–20.1, +4.6) |} <!-- END TABLE --> <div id="section-5-2-4-4chemosynthetic-ecosystems"></div> <span id="chemosynthetic-ecosystems"></span>
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