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Biogeography of the Global Ocean's Mesopelagic Zone

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Biogeography of the Global Ocean's Mesopelagic Zone Report Biogeography of the Global Ocean’s Mesopelagic Zone Graphical Abstract Authors Roland Proud, Martin J. Cox, Andrew S. Brierley Correspondence [email protected] In Brief Proud et al. show that the global ocean can be partitioned into spatially distinct mesopelagic (200–1,000 m) provinces based on the depth of, and echo intensity from, acoustic DSLs. They reveal environmental drivers of DSL variability and infer a significant increase in mesopelagic biomass and trophic efficiency by 2100. Highlights d Deep scattering layer (DSL) variability demarks a global mesopelagic biogeography d DSL backscattering intensity is predictable from primary productivity and temperature d DSL depth is predictable from primary productivity and surface wind stress d DSL shallowing and ocean warming will give rise to an increase in DSL biomass by 2100 Proud et al., 2017, Current Biology 27, 113–119 January 9, 2017 ª 2017 Elsevier Ltd. http://dx.doi.org/10.1016/j.cub.2016.11.003 Current Biology Report Biogeography of the Global Ocean’s Mesopelagic Zone Roland Proud,1,3,* Martin J. Cox,2 and Andrew S. Brierley1 1Pelagic Ecology Research Group, Gatty Marine Laboratory, Scottish Oceans Institute, University of St Andrews, East Sands, St Andrews KY16 8LB, Scotland, UK 2Australian Antarctic Division, 203 Channel Highway, Kingston, TAS 7050, Australia 3Lead Contact *Correspondence: [email protected] http://dx.doi.org/10.1016/j.cub.2016.11.003 SUMMARY RESULTS The global ocean’s near surface can be partitioned Deep Scattering Layers and Acoustic Sampling into distinct provinces on the basis of regional pri- Deep scattering layers (DSLs) are ubiquitous features of the global mary productivity and oceanography [1]. This ocean that comprise biomass-rich communities of zooplankton ecological geography provides a valuable frame- and fish. They are so dense (biomass per unit volume) that in early work for understanding spatial variability in acoustic surveys echoes from DSLs were mistaken for seabed ecosystem function but has relevance only partway echoes, hence the common name ‘‘false bottom.’’ The mesope- lagic is defined as the 200 to 1,000 m depth horizon (e.g., [8]). into the epipelagic zone (the top 200 m). The meso- The physics of sound propagation enables this zone to be pelagic (200–1,000 m) makes up approximately sampled effectively from the surface with commonly employed 20% of the global ocean volume, plays important 38-kHz echosounders. Previous studies from tropical to sub-polar roles in biogeochemical cycling [2], and holds seas suggest that DSLs are rare beneath 1,000 m (e.g., [9, 10]). potentially huge fish resources [3–5]. It is, however, hidden from satellite observation, and a lack of General Characteristics of Regional-Scale DSLs globally consistent data has prevented develop- We used an automated, reproducible technique [11] to identify ment of a global-scale understanding. Acoustic and characterize DSLs in 38-kHz acoustic data collected from deep scattering layers (DSLs) are prominent fea- the top 1,000 m by numerous research and fishing vessels tures of the mesopelagic. These vertically narrow around the world. We collated data from survey transects (tens to hundreds of m) but horizontally exten- totaling 104,688 km in length (see Figure S1). Together these sive (continuous for tens to thousands of km) contained 26,474 DSLs >10 km long. layers comprise fish and zooplankton and are Inspection of the global DSL dataset revealed pronounced geographic differences in DSL depth, vertical extent (thickness), readily detectable using echosounders. We have and acoustic backscattering intensity (quantified as area back- compiled a database of DSL characteristics glob- 2 2 scattering coefficient [ABC], m mÀ [12]). ABC can be a linear ally. We show that DSL depth and acoustic back- proxy for biomass [3]. In this case, ABC is the total acoustic scattering intensity (a measure of biomass) can backscatter per m2 from DSLs in the mesopelagic zone: hence- be modeled accurately using just surface primary forth, we use the term ‘‘backscatter’’ for simplicity. Although it is productivity, temperature, and wind stress. Spatial tempting to convert backscatter to a measure of actual biomass variability in these environmental factors leads [3], we lack the data on species composition and size, and also to a natural partition of the mesopelagic into on acoustic target strength, to do this [13]. Our analysis hence- ten distinct classes. These classes demark a forth is therefore relative rather than absolute. more complex biogeography than the latitudinally Generally speaking, during the daytime, the mesopelagic zone banded schemes proposed before [6, 7]. Knowl- contained a principle DSL that was vertically broad (extending over >200 m vertically), relatively dense (backscatter approxi- edge of how environmental factors influence 5 2 2 mately 1.59 3 10À m mÀ ), and commonly (>66% chance) the mesopelagic enables future change to be centered at a depth of approximately 525 m (Figure 1). There explored: we predict that by 2100 there will be was also sometimes (<20% chance) a secondary, less dense 6 2 2 widespread homogenization of mesopelagic com- DSL (backscatter approximately 1.26 3 10À m mÀ ) approxi- munities and that mesopelagic biomass could mately 300 m deeper. increase by approximately 17%. The biomass increase requires increased trophic efficiency, Environmental Drivers of DSL Variability which could arise because of ocean warming and Differences in DSL characteristics across oceanographic frontal DSL shallowing. boundaries have been reported previously [15], but variability at Current Biology 27, 113–119, January 9, 2017 ª 2017 Elsevier Ltd. 113 Figure 1. Scattering Layer Daytime Vertical Distribution and Acoustic Backscattering Intensity A typical daytime water-column acoustic profile (an echogram), showing a ‘‘surface’’ scattering layer in the epipelagic zone (0–200 m), a principal deep scattering layer (DSL) at around 525 m (the global mean), and a secondary DSL at around 825 m, both in the mesopelagic (200–1,000 m). Data were recorded using a 38-kHz echosounder from the fishing vessel Will Watch [14] on May 30, 2012 in the southwest Indian Ocean (28.8S, 47.3E). The color bar is mean volume backscattering 1 strength (MVBS, dB re 1 mÀ ;[12]). the global scale has not been quantified. The spatial coverage of lagic provinces, a number similar to the 32 surface provinces our data spanned 14 of Longhurst’s [1] 32 pelagic surface prov- advocated by Longhurst [1] (see Supplemental Information, Fig- inces (excluding his coastal biome; see Figure S1). We binned ure S2). By choosing to focus on this scale, we were able to daytime DSL data by these surface provinces (there can be ma- compare Longhurst’s surface biogeography and our mesope- jor differences between daytime and nighttime depths of DSLs lagic biogeography: they do not overlap directly (Figure 3A). due to diel vertical migration [16], so we separated daytime Our ten-class mesopelagic biogeographic structure is more and nighttime data to avoid introducing temporal artifacts to complex and heterogeneous than the simple latitudinal banding our spatial analysis). Variability in depth of the principle daytime that pervades previous surface [6] and abyssal [7] schemes. DSL (ZPDSL, m; see Figure 1) was explained well at this spatial Although the Southern Ocean is latitudinally banded in our scheme scale (n = 14, R2 = 0.68, root-mean-square error [RMSE] = (reflecting the quasi-parallel oceanographic frontal structure in 28 m) by a simple multilinear model with mean annual primary that ocean [18]), a markedly different arrangement is evident else- 2 1 production (PP, g C mÀ dayÀ , p = 0.01) and surface wind stress where. For example, the central tropical gyres of the north and 2 (t,NmÀ , p = 0.001) as explanatory variables (Figure 2A). The south Pacific Ocean both cluster into the same class. Classes variability in backscatter from DSLs was explained well (n = 14, with high backscatter values (high mesopelagic biomass) are 2 6 2 2 R = 0.65, RMSE = 9.11 3 10À m mÀ ) by a simple multilinear found across the north Atlantic and within frontal zones at mid-lat- model incorporating PP (p = 0.017) and the temperature at the itudes, with the exception of the south Pacific sector of the South- depth of the principal DSL (TPDSL, C, p = 0.0001; Figure 2B). ern Ocean. Classes with lower backscatter values (low mesope- lagic biomass) include the polar oceans and the south Atlantic. Mesopelagic Biogeography We used a clustering approach to explore the likely geographic Present-Day Backscatter and Trophic Efficiency distribution of distinct DSL types across the global ocean (areas We estimated total global backscatter by summing together the 2 2 where total depth R1,000 m). We gridded (at 300 3 300 km products of the predicted mean backscatter value (m mÀ ) and scale) PP and TPDSL (estimated from predicted values of ZPDSL, surface area of each mesopelagic class. The present-day value which is a function of PP and t; see Figure 2A) and used K-means was 6.02 3 109 m2 ± 1.4 3 109 m2 (error limits from regression clustering (see Supplemental Information) of the normalized vari- model RMSE value; see Figure 2C). ables to identify coherent mesopelagic classes across a range of Biological production (the increase in biomass per unit time) is spatial scales (from n = 3 to 35 classes globally, classes having a function of biomass, temperature, and trophic level (TL) [19]. characteristic backscatter, PP and TPDSL values; see Supple- The mesopelagic community is made up of organisms operating mental Information, Figure S3). at a range of TLs between 2 and 4. Myctophid fish (TL = 3.2; www. The ability to model regional variability in backscatter was best fishbase.org) are a major component of mesopelagic biomass at the scale of 22 mesopelagic classes (n = 17, R2 = 0.93, [20, 21].
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