the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Temporal and Spatial Influences of Environmental Factors on the Distribution of Mesopelagic organism in the North Atlantic Ocean
Abstract. Mesopelagic organisms play a critical role in marine ecosystems and the global carbon cycle, acting as key intermediaries between trophic levels through diel (DVM) and seasonal vertical migrations (SVM). However, the seasonal vertical migration patterns of these organisms, and the environmental drivers influencing them, remain insufficiently understood. Here, we analyzed 83,603 backscattering coefficient (bbp) profiles obtained from 720 BGC-Argo floats deployed in the North Atlantic Ocean from 2010 to 2021. This extensive dataset enabled the identification of spiking layer signals, allowing us to investigate the diurnal and seasonal vertical distributions of mesopelagic organisms, as indicated by these bbp spikes. Additionally, we examined the horizontal heterogeneity in these distributions and their correlations with key environmental variables. Our findings reveal distinct diurnal migrations, characterized by multilayered aggregations predominantly in the mid-ocean during daylight, with prominent signals at depths around 150 m, 330 m, 650 m, and 780 m. At night, a strong scattering layer forms in the upper ocean, with signals concentrated at depths shallower than 350 m, particularly in the top 100 m. Seasonal analyses shows that in spring and winter, the average bbp spike intensity is lower in the upper ocean than in the mid-ocean, although the frequency of bbp spikes is higher in the upper ocean. In contrast, summer and autumn—especially summer—exhibit both higher mean bbp spike intensity and frequency near the surface. Spatially, mesopelagic organisms migrate deeper in the northeast and remain shallower in the southwest, correlating with higher temperatures and shallower distributions. Random forest analysis identified temperature as the most influential environmental factor affecting the distribution of mesopelagic organisms year-round, with the temperature gradient being particularly critical. Other critical factors include seawater salinity, dissolved oxygen, surface chlorophyll concentration, and latitude, with relative importance of 29.44 %, 15.49 %, 14.85 %, 13.46 %, and 12.35 %, respectively. This study enhances our understanding of the mechanisms driving carbon transfer to the deep ocean and the energy and material cycles within marine ecosystems, providing a basis for future fisheries management in mesopelagic environments.
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RC1: 'Comment on egusphere-2024-2991', Anonymous Referee #1, 17 Nov 2024
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The authors of Temporal and spatial influences of environmental factors on the distribution of mesopelagic organisms in the North Atlantic Ocean have conducted an interesting and ambitious study extending the analysis carried out by Haentjens et al., 2020 to address crucial questions about the spatiotemporal distribution of mesopelagic organisms in the North Atlantic.
I have found a few challenges understanding what Jie Yang and colleagues have done so I hope my comments can help them clarify their work. I recommend engaging with major revisions of this work that I believe has good potential. I wish the best to the authors and I am looking forward to learning about their findings from the suggested further analyses.
I have a few major concerns:
- The methods need more detail. The Material and methods part of the study is very brief which makes it hard to assess whether the used methods are appropriate or not. For example:
- Why did the authors include remotely sensed data of variables that are measured by BGC-Argo anyways? Wouldn’t the in situ data be more accurate and surely match the location of the floats compared to 4km resolution satellite data?
- The calculation of the smad needs to be better explained with a description of all the terms included in equation (1).
- Why did the authors calculate vertical temperature gradients? How do they expect they would impact the distribution of mesopelagic organisms? Is this an assessment to measure stratification (e.g., Fernandez et al., 2017)? If so, why not using MLD instead? Or are they horizontal gradients?
- In Figure 2 - what are the profiles ‘Profile 1’ and ‘Profile 2’ that are used in the analysis? How can they be concurrent?
- The authors mentioned dividing profiles by whether they are from the day or night. What about profiles collected at dawn and dusk? Most BGC-Argo profiles will likely measure profiles around noon or midnight so I suspect there won’t be many of those, but profiles at dawn and dusk could confuse the analysis so I believe should be excluded or treated separately.
- The random forest model needs to be described in more detail. For example, what is the response variable that is then depicted in Figure 7- is it the anomaly of the depth of the main spike? Also, how was the random forest parameterized (e.g., number of trees, etc.?)
- The authors mention (and I think it’s a good idea) normalised profiles. Yet, I couldn’t find a very clear explanation of how profiles are normalised (including in the Figure 2 diagram). Please explain this important step of your analysis.
- Some of the results appear to be a bit confusing to me. In particular:
- Something that I find challenging is how to disentangle the spatial variability from the temporal one. The area of study includes regions with complex and diverse oceanographic regimes (e.g., Della Penna and Gaube, 2019 for differences within the western side of the domain, but also strong differences between the eastern and western side) so if the profiles for a specific season are mostly from a specific region there is a risk of associating the typical pattern observed with the season rather than the region. There are a few extra analyses that could help with this, for example: Figure 1. What is not clear to me is - are these the used validated profiles or the total profiles available for the regions? Why is this picture so different from Figure 6?. I recommend including in the supplementary material the equivalent map plotted for each individual season to identify if there is a spatial bias associated with each season.
- Some results are described in the text but are not clearly backed up by figures or tables. For example, in lines 193-195 the authors talk about light conditions, but I wonder if these are included in the modelling? Even just having the solar angle could be a way to incorporate season and location in a single descriptor.
- Figure 7 is from my perspective where a lot of valuable results should be but it’s not as clear as I think it should be: what are the explained variables on the y axis (e.g., what are the numbers -150,200 meaning on the axis?) and it doesn’t include any units.
- The interpretation feels a bit overstretched to me in a few points:
- The authors compare the patterns they observed with those identified by Klevjer et al., 2020a, yet both the Klevjer et al., papers in 2020 are focused on a small portion of the domain of this study. I suggest the authors be cautious with how they are connecting their findings with the Klevjer et al., 2020 papers and potentially use other references to compare their findings with. For example, Klevjer et al., 2016 show results from the southern part of the North Atlantic and Della Penna and Gaube, 2020, Wiebe et al., 2023, and Fennell and Rose, 2015, show results from the Western side of the North Atlantic. I’m sure there are more studies the authors could consider to compare the distributions of spike layers they observed.
- A few pretty important statements are not backed up by references. For example the statement in lines 209-210 needs references. In addition, I think that in some cases the references used are not really backing up the statements made in the discussion. For example, the Contrerar-Catala et al., 2016 paper to my knowledge deals with the larvae of mesopelagic fish and not mesopelagic fish in general (I’m also not sure they used the same metrics of signal abundance and frequency so it’s hard to make a comparison).
- The relationship between latitude, light levels, and temperature is not explored very clearly. In line 256 the authors discuss the fact that the bbp spikes appear shallower in warmer regions, gradually deepening with increasing latitudes. They attribute this to a change in light conditions, but then they back their statement up with an explanation based on temperature. I think the mechanisms associated with light levels and temperature need to be disentangled (or discussed more clearly at least). To my knowledge, underwater light levels are strongly related with the distribution of DSL with a variability that occurs at scales ranging from the basins (Asknes et al., 2017) to the small differences in cloud coverage (Omand et al., 2021). In general, many species seem to ‘follow an isolume’ (see example from Della Penna et al., 2022 or Asknes et al., 2017). According to this framework, we expect the animals inhabiting DSL to be deeper in clearer waters and shallower in waters with higher light attenuation coefficients (see for example Braun et al., 2023). This is quite the opposite of what the authors are describing in their observations. Perhaps there is another mechanism that is dominating here. Could it be a different mesopelagic community (see the work by Proud et al., 2017 or the recent paper by Chawarski et al., 2022). I’m also wondering, could this pattern be explained by the larger abundance of sinking aggregates at high latitudes (smaller phytoplankton could dominate the lower latitudes with particles that just don’t make it that deep)?
Minor points
Throughout the text: I suggest having a space between text and references. For example, at the end of page 1: “nutrient cycling(Klevjer et al., 2016)” would be more readable if written as “nutrient cycling (Klevjer et al., 2016)”
Title: I suggest editing the title adding an ‘s’ to ‘organism’ as I think the authors are interested in more than a specific one.
Abstract:
Line 5: “spiking layer signals”: this is not a concept that is obvious - I suggest using a more commonly used term, maybe ‘bbp spikes’?
Line 14: ‘shallower distributions’ of what?
Line 16: Please specify that you are talking about vertical temperature gradient (is this the case?)
Introduction
Line 25: “mesopelagic zone is diel vertical” -> “mesopelagic zone is the diel vertical”
Line 30: The statement about the SVM needs a reference or two to back it up.
Line 40: Why is there a specific reference to ADCP? The statement made here is about all active acoustics approaches (ADCP but also scientific echosounders).
Line 46: “renderiimportanceerful” is not a word. Please edit.
Line 56: In what way the Behrenfeld et al., 2019 paper discussed evolutionary patterns?
Material and methods
Line 72: I don’t think it is correct to say in any way that pelagic fish are largely untapped resource for fisheries, especially in the North Atlantic! Maybe the authors refer to ‘mesopelagic’ here? I suggest clarifying or removing this bit of text.
Line 124: I suggest removing the use of the word ‘pinnacle’ and sticking with ‘spikes’.
Figure 3 - I think this figure could be a great opportunity to provide a visual reference to the reader of how a profile of the metrics discussed in the results look like (e.g., frequency and density or spikes, etc.). I suggest adding them as a subfigure so that the reader has an immediate sense of how these metrics relate to the original ‘spike’ data. I also suggest using 2.0 as the maximum value of the bbp ratio as there are no points about 2.0 and going to 2.5 is a bit of a waste of precious space.
Results - Lines 168-176: I think this content belongs to the discussion and not in the results.
Through the results: I think you are using here only daytime profiles but I’m not sure I could find easily this piece of information anywhere. If this is the case, please make sure it’s explicit.
Line 180: Why is there a reference to the Mediterranean Sea here?
Line following 185: Are the environmental variables used here from remote sensing or those measured from BGC floats?
Discussion
Line ~205: When referring to the sources of variability of the position of DSL I recommend including the presence of mesoscale eddies as there is a good amount of work that showed they play quite a role in structuring DSL in the area: Fennel and Rose, 2015; Della Penna and Gaube, 2020; Devine et al., 2021.
Line 221: Why is there a reference to a ‘mesocosm’? Please rephrase.
Line 228: What is the ‘mesopelagic acropora signal’?
Line 272 and following: The impact of fronts on aggregations are not limited to downwelling and upwelling, so I suggest including here a mention as well of the horizontal mechanisms described in the next few lines. A very interesting discussion of light and fronts, water masses and zooplankton can also be found in Powell and Ohman, 2015.
References (not including those already in the paper)
Aksnes, D. L., Røstad, A., Kaartvedt, S., Martinez, U., Duarte, C. M., & Irigoien, X. (2017). Light penetration structures the deep acoustic scattering layers in the global ocean. Science advances, 3(5), e1602468.
Braun, C. D., Della Penna, A., Arostegui, M. C., Afonso, P., Berumen, M. L., Block, B. A., ... & Thorrold, S. R. (2023). Linking vertical movements of large pelagic predators with distribution patterns of biomass in the open ocean. Proceedings of the National Academy of Sciences, 120(47), e2306357120.
Della Penna, A., & Gaube, P. (2019). Overview of (sub) mesoscale ocean dynamics for the NAAMES field program. Frontiers in Marine Science, 6, 384.
Della Penna, A., & Gaube, P. (2020). Mesoscale eddies structure mesopelagic communities. Frontiers in Marine Science, 7, 454.
Della Penna, A., Llort, J., Moreau, S., Patel, R., Kloser, R., Gaube, P., ... & Boyd, P. W. (2022). The impact of a Southern Ocean cyclonic eddy on mesopelagic micronekton. Journal of Geophysical Research: Oceans, 127(11), e2022JC018893.
Devine, B., Fennell, S., Themelis, D., & Fisher, J. A. (2021). Influence of anticyclonic, warm-core eddies on mesopelagic fish assemblages in the Northwest Atlantic Ocean. Deep Sea Research Part I: Oceanographic Research Papers, 173, 103555.
Klevjer, T. A., Irigoien, X., Røstad, A., Fraile-Nuez, E., Benítez-Barrios, V. M., & Kaartvedt, S. (2016). Large scale patterns in vertical distribution and behaviour of mesopelagic scattering layers. Scientific reports, 6(1), 19873.
Fennell, S., & Rose, G. (2015). Oceanographic influences on deep scattering layers across the North Atlantic. Deep Sea Research Part I: Oceanographic Research Papers, 105, 132-141.
Fernandez, D., Sutton, P., & Bowen, M. (2017). Variability of the subtropical mode water in the Southwest Pacific. Journal of Geophysical Research: Oceans, 122(9), 7163-7180.
Omand, M. M., Steinberg, D. K., & Stamieszkin, K. (2021). Cloud shadows drive vertical migrations of deep-dwelling marine life. Proceedings of the National Academy of Sciences, 118(32), e2022977118.
Powell, J. R., & Ohman, M. D. (2015). Changes in zooplankton habitat, behavior, and acoustic scattering characteristics across glider-resolved fronts in the Southern California Current System. Progress in Oceanography, 134, 77-92.
Wiebe, P. H., Lavery, A. C., & Lawson, G. L. (2023). Biogeographic variations in diel vertical migration determined from acoustic backscattering in the northwest Atlantic Ocean. Deep Sea Research Part I: Oceanographic Research Papers, 193, 103887.
Citation: https://doi.org/10.5194/egusphere-2024-2991-RC1
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