Trophic amplification: A model intercomparison of climate driven changes in marine food webs

and 15% under GFDL and IPSL, respectively, for both EcoTroph, FEISTY, and BOATS and about 15% and 5% under GFDL and IPSL, respectively for FEISTY (S8 Fig). Thus, an approach based on the productivity of ecosystems and not their biomass results in less trophic amplification, which suggests a smaller impact on fisheries potential catches. 4.6 Low trophic level, key MEMs driver, estimation uncertainty and consequences In our study, we find that depending on low trophic level forcing in each MEM (low trophic level biomass vs. low trophic level production), the magnitude of biomass changes relative to primary productivity is very different (S9 Fig), which is consistent with the results highlighted in Heneghan et al (2021). The magnitude of the biomass response of MEMs is, on average, larger for MEMs driven at low trophic level by production than those driven by biomass forc- ing (S1 Table and S9 Fig). The nature of this difference, thus, could firstly lie in the inconsis- tency that arises from 1-way forcing with biomass or production [19]. This could dampen the consumer decline when forcing with biomass. For example, the comparison of 1-way and 2-way forcing with APECOSM supports this assumption, although the small difference in terms of global biomass decline might not explain all [50]. This study highlighted the relative changes in consumer biomass compared to changes in NPP to investigate trophic amplification. Hence, we compared a production (NPP) to a bio- mass (consumer biomass). In a supplementary analysis, we explored the changes of low tro-

phic level drivers of each MEM (other than NPP; i.e., phytoplankton and zooplankton biomass) which have a very different spatial structure and temporal trend (S9 Fig) compared to NPP [5]. At the global scale, low trophic level (i.e., phytoplankton and zooplankton) biomass is projected to decrease under IPSL while NPP is projected to increase.. One of the mecha- nisms driving this inversion (increase in NPP, decrease in phytoplankton biomass) is when temperature-driven phytoplankton metabolic costs exceed what can be met with the increase in NPP. Under GFDL, the decrease in phytoplankton and zooplankton biomass is projected to be slightly stronger than that of NPP. PLOS ONE Trophic amplification revealed PLOS ONE | https://doi.org/10.1371/journal.pone.0287570 August 23, 2023 17 / 23

As shown in S9 Fig, trophic amplification based on low trophic level driver of each MEMis reduced but consistent with to trophic amplification based on NPP under GFDL.On the con- trary, the signal inversion between NPP and other MEMlow trophic level driver, under IPSL, lead to a different perception of trophic amplification (compared to trophic amplification rela- tive to NPP) for APECOSM; EcoOcean, FEISTY and ZooMSS. The inversion in trend under IPSL (Fig 3, changes in consumer biomass compared to changes in NPP) becomes an amplifi- cation for the four aforementioned models when considering the changes in the individual model’s low trophic level drivers (S9 Fig). 5. Conclusion The main response to the global decrease in NPP is a negative amplification, resulting in a more pronounced decrease in consumer biomass. Our study suggests that this response may be due to the way temperature influences different physiological and ecological processes in each ecosystem model. When an increase in NPP is projected, the temperature can offset the positive impact by attenuating or reversing the changes in biomass, leading to smaller increases or even decreases in biomass corresponding to positive attenuation or negative inversion respectively. Disaggregation of trophic amplification across different sizes within the food web highlighted a greater decrease in biomass of larger, high trophic levels on average. As a result, there are believable parts of the ocean for which we may see a large climate-induced decline in fishable biomass despite no change or increasing NPP.

Supporting information S1 Table. A taxonomy of marine ecosystem models taking part in the Fish-MIP project (modified from (Heneghan et al., 2021; Lotze et al., 2019; Tittensor et al., 2021)). (TIF) S2 Table. Conversion between trophic levels-based and size-based bins. (TIF) S1 Fig. Spatial patterns of projected total consumer biomass relative changes in percent. Shown are global ensemble projections at a 1 × 1 degree resolution. mean 2090s individual model projections of total consumer biomass relative change over the reference period (1995_2014) under: (a) GFDL-SSP5-8.5; and (b) IPSL-SSP5-8.5. (TIF) S2 Fig. Main outputs of the earth system models considered in the current study. mean changes in Sea surface temperature (a) and net primary production (b) in the 2090s relative to the reference period 1995_2014, under SSP5-8.5 for IPSL. (TIF) S3 Fig. Percentage of surface covered by each total consumer biomass response type, for the seven considered MEMS, under GFDL-SSP5-8.5 modelling. (TIF) S4 Fig. (a) Spatial distribution of the total consumer biomass response types, for the nine con- sidered MEMs, forced with IPSL-SSP5-8.5 modelling and (b) magnitude of the different types of change for the seven considered MEMS forced with IPSL-SSP5-8.5 Each response type is estimated at a 1 × 1 degree resolution, from the change in consumer biomass and NPP expected at the end of the century, relatively to the reference period. (TIF) PLOS ONE Trophic amplification revealed PLOS ONE | https://doi.org/10.1371/journal.pone.0287570 August 23, 2023 18 / 23

S5 Fig. The percentage of surface covered by each total consumer biomass response type, for the nine considered MEMS, under IPSL-SSP5-8.5 modelling. (TIF) S6 Fig. Model agreement on biomass response types projected in the 2090s, under IPSL-SSP5-8.5 configuration. Response types correspond to at least seven out of nine models in agreement for the coloured cells while grey cells indicate where less than seven models proj- ect the same type of response and white cells where no data were available. The percentage numbers in the pie chart correspond to the relative surface areas. (TIF) S7 Fig. Evolution of R ratio (total consumer biomass change divided by NPP change) in regards to SST increase relative to reference period. Where NPP is expected to decrease (a), or increase (b) under IPSL-SSP5-8.5 combination. Grey lines separate biomass response types. (1), (2) and (3) refer to positive inversion, negative attenuation and negative amplification, respectively, while (4), (5) and (6) refer to positive inversion, negative attenuation and negative amplification, respectively. (TIF) S8 Fig. Comparison between change over time in the total consumer production and change in total consumer biomass. Under EcoTroph simulation (first row) FEISTY simula- tion (second row) and BOATS simulation (third row). The Left and the right plots correspond to GFDL-SSP5-8.5 and IPSL-SSP5-8.5 forcing, respectively. Blue, and red lines correspond to the total consumer biomass change and total consumer production change, respectively. (TIF)

S9 Fig. Ensemble projections of low trophic level drivers change and total consumer biomass changes, relative to 1995–2014, under SSP5-8.5 and for all the considered MEMs with the GFDL-SSP5-8.5 forcing (a) and with the IPSL-SSP5-8.5 forcing (b). For temporal trends, all values are relative to the standardised reference period of 1995–2014. Vertical grey line indi- cates the last year of the historical period. Full lines correspond to MEMs’ total consumer bio- mass change and lines with diamonds to MEMs’ low trophic level changes. (TIF) Acknowledgments We thank ISIMIP for assistance with processing ESM inputs and outputs and the ESM/MEM community for enabling these impact analyses. This work has been conducted as part of a col- laboration between L’Institut Agro, Rennes, and the University of British Columbia, Vancou- ver. We further thank Jerome Guitton for his technical support. Author Contributions Conceptualization: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Data curation: Vianney Guibourd de Luzinais, Hubert du Pontavice, Nicolas Barrier, Julia L. Blanchard, Matthias Bu¨chner, Jason D. Everett, Jerome Guiet, Olivier Maury, Camilla Novaglio, Colleen M. Petrik, Jeroen Steenbeek. Formal analysis: Vianney Guibourd de Luzinais. Funding acquisition: William W. L. Cheung, Didier Gascuel. PLOS ONE Trophic amplification revealed PLOS ONE | https://doi.org/10.1371/journal.pone.0287570 August 23, 2023 19 / 23

Investigation: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Methodology: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Project administration: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Resources: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Software: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Supervision: Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Validation: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Visualization: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Writing – original draft: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, William W. L. Cheung, Didier Gascuel. Writing – review & editing: Vianney Guibourd de Luzinais, Hubert du Pontavice, Gabriel Reygondeau, Nicolas Barrier, Julia L. Blanchard, Virginie Bornarel, Matthias Bu¨chner, William W. L. Cheung, Tyler D. Eddy, Jason D. Everett, Jerome Guiet, Cheryl S. Harrison, Olivier Maury, Camilla Novaglio, Colleen M. Petrik, Jeroen Steenbeek, Derek P. Tittensor, Didier Gascuel. References 1. IPCC. Observations: Ocean Pages. Climate Change 2013—The Physical Science Basis, Cambridge:

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