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The biotic induction of OAE-2The rapid proliferation of select microbial communities at 427.54 mcd likely represents a pre-OAE biotic perturbation (pre-OAE BP) presaging the protracted period of widespread marine deoxygenation during OAE-2, and progressive deoxygenation predating the +CIE7 (Fig. 4). At the beginning of the pre-OAE BP (427.54 mcd), abruptly elevated tetrapyrroles and crenarchaeol concentrations signify an abrupt increase in primary production by photoautotrophs and chemoautotrophs residing above the chemocline. Increased volumes of precipitating biogenic snow concordantly consumed oxygen, expanding the preexisting OMZ as anaerobic bacteria thrived based on accelerated obGDGTs synthesis. Euxinia did not penetrate the photic zone at the outset of the productivity bloom as isorenieratane was not detected and heightened rates of microbial sulfate reduction were seemingly transient, inferred from the DAGEs profile, and limited to pre-OAE BP initiation. The lack of a well-stratified water column, evinced by absent to low concentrations of halophilic archaeal lipids (i.e., extended archaeols), relatively low rates of microbial sulfate reduction, and a dense oxygenic microbial plate likely precluded the development of PZE initially.Establishing a definitive causal mechanism for the pre-OAE BP is difficult, but the concomitance of LIP activity with the productivity spike is intriguing. Application of a linear sedimentation rate from OAE-2 to the pre-OAE BP interval following previous works6,7 approximated the pre-OAE BP occurring 220 ± 4 kyr before OAE-2, lasting for ~100 kyr (427.54–426.88 mcd; see Estimating the duration of the pre-OAE BP in Supplementary Information for rationale and calculation). Significantly, this was roughly coincident with the onset of LIP activity (~200–300 kyr before OAE-2) inferred from marine osmium isotope stratigraphy27. Similarities in the modern planktonic community response, such as elevated productivity and compositional changes, between the 2018 Kilauea eruption28 and the pre-OAE BP reinforce inference of a potential magmatic trigger for this event (see Evidence for LIP trigger of the pre-OAE biotic perturbation in Supplementary Information for additional details).A constant, yet overall lower, nutrient and trace metal inventory6 (Fig. S4) combined with a redox-driven shift in fixed N species (from NO3− to NH4+)15, potentially leading to a fixed N shortage29 via intensified denitrification and annamox reactions30, were probable culprits in the failure to sustain prolific rates of primary production beyond 100 kyr at the Demerara Rise. The gradual decline in biomass production, indicated by decreasing tetrapyrrole and crenarchaeol profiles (Fig. 4), was accompanied by a notable shift in deep water communities. Sulfate-reducing bacteria exerted increasing predominance over methanogenic archaea, a trend coeval with the primary productivity spike and extending well into the OAE (Fig. 3). A collapse of autotrophic communities to pre-perturbation levels was concordant with the progressive shoaling of H2S-laden waters. Continued vertical migration of the chemocline intruded the photic zone, producing PZE that enabled anoxygenic photosynthesis by Chlorobiaceae (Fig. 4). Unlike the overall oscillatory character of PZE throughout the studied section, this protracted phase of PZE was sustained until the onset of OAE-2 (426.43–426.00 mcd, Figs. 3 and 4) and is approximately contemporaneous with a thallium (Tl) isotope excursion7 (426.40–426.30 mcd).The positive Tl isotope excursion represents the progressive expansion of bottom water anoxia predating OAE-2 by 43 ± 11 kyr6,7. However, evidence for a causal mechanism of pre-OAE deoxygenation remains indeterminate. Our comprehensive biomarker inventory provides an interpreted sequence of events culminating in the regional to global expansion of anoxia predating OAE-2. A protracted phase of enhanced primary productivity began ~220 ± 4 kyr prior to OAE-2, increasing localized production and export of organic carbon at Demerara Rise. Similar productivity spikes likely occurred in settings of comparable paleogeographic configuration (e.g., equatorial, continental margins/shelves), seeding the oceans with fixed carbon. Continued scavenging of marine oxygen via organic carbon remineralization resulted in OMZ expansion locally, and likely initiated oxygen drawdown in much of the proto-North Atlantic Ocean. Stratigraphic records of sulfur isotopes of pyrite (δ34Spyrite) from the proto-North Atlantic and Tethys Oceans11 validate the areal extrapolation of our interpretations. A gradual decline in δ34Spyrite values at Demerara Rise begins at 427.50 mcd, nearly identical to the onset of the pre-OAE BP (427.54 mcd, Fig. 4). Correlation of δ34Spyrite in a global transect (Western Interior Seaway, proto-North Atlantic, Tethys) revealed consistent behavior in δ34Spyrite prior to the +CIE, indicating increasingly expansive marine deoxygenation on a global scale11. Over ~100 kyr, increased regional biomass production induced pervasive marine anoxia, inhibiting Mn-oxide formation, producing the observed positive Tl isotope excursion, and ultimately, the globally observed +CIE reflecting enhanced organic carbon burial signaling the onset of OAE-2. Thus, the local biotic signal recorded at ODP Site 1258 underlines the crucial role the Demerara Rise, and similar undocumented settings, served in initiating deoxygenation of the global ocean.Microbial ecological dynamics during and after OAE-2Changes in microbial community compositions during OAE-2 were apparent, signified by a shift in the normalized total biomarker pool (Fig. 3) and variations in the absolute concentrations of individual biomarkers (Fig. 4). In general, OAE-2 was defined by an expansion and diversification of intermediate and deep water communities (426.00–423.07 mcd), followed by a period of instability leading to the termination of the OAE (423.07–422.00 mcd). Photo- and chemoautotrophs residing above the chemocline were adversely affected, evinced by relatively low, invariant tetrapyrrole and crenarchaeol profiles (Fig. 4). Based on these observations, we divided OAE-2 into two periods defined by contrasting paleoenvironmental conditions modulating the microbial inhabitants of Demerara Rise.The first period of OAE-2 (426.00–423.07 mcd, Fig. 4) was marked by the intrusion of a euxinic OMZ into the photic zone. Elevated, yet fluctuating isorenieratane concentrations suggest relatively persistent PZE of varying vertical extent, in agreement with previous investigations using biomarkers and nitrogen isotopes at nearby sites12,13,31. During this interval, microbial sulfate reduction was likely active as DAGEs continually increased, aligning with estimates of expanded seafloor euxinia32. The co-occurrence of abundant extended archaeols and isorenieratane intimates the role that density stratification served in maintaining the protracted PZE of OAE-2, substantiating concurrent findings based on neodymium33 and oxygen isotopes34. Vertical nutrient advection via upwelling35 led to preferential exposure to expanding intermediate water communities tolerant to sulfidic conditions in the OMZ. Scavenging of a potentially limited fixed N inventory30, depleted in NO3− and predominated by NH4+[ 15,29, and inhibition of efficient nutrient transfer by pronounced density stratification likely induced severe N deficiency in surface water communities, explaining the relatively muted productivity of oxygenic photoautotrophs (i.e., tetrapyrroles) and chemoautotrophs (i.e., crenarchaeol) observed (Fig. 4). The concentration and predominant utilization of fixed N in the OMZ led to the proliferation and diversification of intermediate and deep water microbial taxa, while a shoaling chemocline led to increased nutrient (i.e., fixed N) competition between photoautotrophs and retreating Thaumarchaeota as highlighted by our biomarker inventory and the nitrogen isotopic record31. These findings challenge previous interpretations of highly productive, predominantly eukaryotic primary producers reliant on the upwelling of isotopically depleted NH4+ during OAE-215. Instead, the decline of C30-17-nor-DPEP (Fig. S5; Supplementary Data 3), a source-specific tetrapyrrole diagenetically derived from algal chlorophyll-c36, and reconstructed water column conditions during OAE-2 indirectly support a rise in cyanobacteria, diazotrophs able to fix N2, in oxygenated, nutrient-depleted shallow waters. Increased cyanobacterial contribution is further supported by C and N stable isotopes16,37, as well as the prominence of potentially phylum-specific biomarkers across OAE-2 (e.g., 2-methylhopanoids6,14).Fig. 5: Contrasting biogeochemical conditions between the pre-OAE BP and OAE-2.a, b Microbial ecology and water column conditions during the pre-OAE BP, reflecting high primary production of organic carbon (a) and OAE-2, characterized by relatively lower organic carbon production, but substantially enhanced biomass preservation (b). c, d Averaged fractional abundances of individual biomarkers throughout the pre-OAE BP (c) and OAE-2 (d). Biomarker source organisms are abbreviated as follows: phytoplankton (P), ammonia oxidizing archaea (AOA), sulfur oxidizing bacteria (SOB), unknown anaerobic bacteria (UAB), sulfate reducing bacteria (SRB), halophilic archaea (HA), methanogenic archaea (MA).Full size imageA reversal from the formerly outlined conditions typified the second period of OAE-2 (423.07–421.99 mcd, Fig. 4). Destabilization of the stratified water column and reduced production of H2S led to deepening and contraction of the euxinic OMZ. The observed decline in halophilic archaea, coincident with an overall decline in Chlorobiaceae populations, is roughly coeval with positive neodymium isotopic excursions observed across the proto-North Atlantic33 attributed to the enhanced latitudinal commingling of proto-North Atlantic water masses38. Although detrimental to sustained PZE, the persistence of a well-developed anaerobic bacterial community (i.e., obGDGTs) suggests the lasting presence of a non-euxinic OMZ despite improved bottom water circulation. A premature recovery of the chemoautotrophic Thaumarchaeota, inhabiting the base of the photic zone, relative to the shallower dwelling obligately oxygenic phototrophs (Fig. 3) likely reflects reduced toxicity associated with retreating euxinic waters, lessened resource competition with [primarily] Chlorobiaceae, and a competitive advantage tied to preferential exposure to upwelled nutrients and tolerance to low O2 conditions.The termination of OAE-2 was marked by the temporary re-establishment of microbial community compositions mirroring those observed prior to the pre-OAE BP (Figs. 3 and 4). Contraction of the OMZ led to a deep chemocline, with PZE restricted to the basal photic zone as the production of reduced sulfide species diminished. The Thaumarchaeota continued the recovery initiated towards the latter half of OAE-2, accompanied by the rebounding oxygenic photoautotrophs. However, the recovery of shallow autotrophic communities was halted by an episode of PZE (421.19–421.04 mcd) based on abrupt increases in isorenieratane concentrations (Fig. 4). Temporary development of pronounced density stratification likely facilitated the accumulation of H2S in the lower to intermediate photic zone, producing the short-lived PZE episode. Interestingly, covariant responses observed in additional biomarker profiles (e.g., obGDGTs) to PZE during OAE-2 were not evident across this post-OAE interval, possibly due to the transient nature of PZE at this time. For example, the initial increase in isorenieratane concentrations at the onset of OAE-2 was not immediately accompanied by shifts in other biomarker classes (e.g., obGDGTs; Fig. 4), suggesting frequent recurrences of PZE may be required to illicit a major microbial ecological response as observed later during the OAE. Still, this brief episode of post-OAE PZE (421.19–421.04 mcd) coincides with a positive organic carbon isotope excursion9 (Fig. S5), trace metal drawdown6 (Fig. S4), and minor positive Tl isotope excursion7 at the Demerara Rise. Prior study7 tentatively attributed this interval to enhanced carbon burial during a post-OAE deoxygenation event of smaller magnitude, with subsequent work revealing continued pyrite burial post-OAE 211. Our biomarker inventory revealed some environmental consistencies (e.g., PZE) between this interval and OAE-2, but the overall biotic response to this post-OAE geochemical perturbation was relatively subdued and requires additional sampling and investigation to properly constrain.Broader implicationsThe recognition of the pre-OAE BP and evolving water column conditions at Demerara Rise highlights additional complexities of a dynamic ocean relevant to interpretations of OAE-2 and the +CIE. Enhanced, sustained, and widespread carbon burial is required to produce the +CIE used to define OAE-28,10. Still, the principal forcing, productivity or preservation, remains enigmatic as evidence for the former mounts12,39.Based on the tetrapyrrole profiles (Fig. 4) primary production was greatest during the pre-OAE BP and relatively muted throughout OAE-2 at Demerara Rise, assuming minimal alteration to the genetic tetrapyrrole stratigraphic signal. Biomass preservation was presumedly enhanced during OAE-2 through sulfurization11, as the OMZ transitioned from anoxic to euxinic and penetrated the photic zone, yet low tetrapyrrole concentrations persist. Previous work noted a similar discrepancy between preservation potential and porphyrin abundance, postulating a paucity of trace metals to chelate with the free-base porphyrins induced poor preservation as desulfurization did not reveal additional porphyrin content16. However, both the pre-OAE BP and OAE-2 were characterized by relatively depleted trace metal inventories6 (Fig. S4), yet exhibit contrasting tetrapyrrole profiles, suggesting relative changes in primary production were the predominate control on the stratigraphic distribution of tetrapyrroles across the studied interval at the Demerara Rise. The strong covariance between tetrapyrrole and crenarchaeol concentrations reinforces the interpretation tetrapyrroles faithfully reflect primary production (Fig. S6). Crenarchaeol, a biosynthetic product of chemoautotrophic archaea (Thaumarchaeota) comprising up to 20% of all archaea and bacteria in the modern ocean40, is structurally distinct from the tetrapyrroles making it likely that diagenetic alteration of the two biomarkers is not consistent in rate or form. Thus, the positive correlation between key proxies for major contributors to primary production, the photoautotrophs and chemoautotrophs, minimizes concern for the integrity of the biotic signal at Demerara Rise (see Tetrapyrroles as a record of primary production in Supplementary Information for additional details).These findings provide direct evidence for a causal mechanism resulting in both the Tl isotope excursion and +CIE as previously described. It is highly probable the pre-OAE BP was not exclusive to the Demerara Rise based on the immense and presently unconstrained organic carbon burial required to produce the +CIE10. Further characterization of comparable localities to Demerara Rise may reveal similar high productivity events, as primed, highly productive settings likely capitalized on exogenous nutrient delivery via efficient upwelling to the photic zone prior to stratification during OAE-2. Hence, OAE-2 and the +CIE were not coincident with heightened surface water productivity relative to the pre-OAE BP at the Demerara Rise. Rather, antecedent increases in primary production locally facilitated the initiation of the OAE as a mechanism to consume marine oxygen and subsequently enhance organic carbon preservation globally. This highlights how OAE-2, and perhaps other OAEs in the geologic record, were not instantaneously induced but rather a gradual transition stemming from sustained forcing(s). In addition, the occurrence of the pre-OAE BP well before the established onset of OAE-2 reveals how fluctuations in primary production can be linked to marine deoxygenation but may not necessarily be concurrent. As shown here, OAE-2 at the Demerara Rise was preceded by elevated primary production that progressively attenuated towards event onset. While the hallmark features of an OAE are well-established, further identification and refinement of trends preceding widespread anoxia in the past will improve our understanding of how marine deoxygenation develops, as well as our ability to assess planetary health today.A shift from a productivity- to preservation-dominant system during OAE-2 at Demerara Rise, and possibly similar paleogeographic settings experiencing the pre-OAE BP, facilitated substantial organic carbon burial producing the +CIE. Distinct shifts in water column chemistry and structure from the pre-OAE BP to OAE-2 imparted considerable changes on microbial life, which altered the primary driver governing biomass sequestration (Fig. 5). Yet, both intervals reveal relatively comparable carbonate-corrected total organic carbon values6 (Fig. S5), signifying enhanced preservation as a critical component of organic carbon burial during OAE-2 at Demerara Rise. Consequently, this work suggests that sustained increases in primary production prior to OAE-2 initiated and regulated pre-OAE deoxygenation, resulting in a progressive shift to preservation as the primary control on organic carbon accumulation in sediments. Expanding euxinia and attendant changes to biogeochemical cycling adversely affected primary producers while simultaneously enhancing organic matter preservation via sulfurization11. Flourishment of Thaumarchaeota in oligotrophic settings in the modern open ocean41, and lack thereof during OAE-2 based on diminished crenarchaeol concentrations, underscores the scarcity of bioessential elements (e.g., fixed N) caused by microbial utilization of electron acceptors further down the redox ladder due to intensified marine anoxia, ultimately limiting primary production. The switch from a productivity to preservation model, reconstructed using biomarkers (Fig. 5) and initially suggested based on drawdown of the trace metal inventory6, was also concomitant with relative warming4. Simulated projections of the marine microbial response to continued global warming in the future revealed similar biotic trends (e.g., decreased primary productivity) to warming-induced oceanographic changes42 (e.g., intensified stratification) observed during OAE-2. Thus, an abundance of proxy- and model-based results paired with conceptual evidence suggest relatively low production and enhanced preservation of organic carbon throughout OAE-2 at the equatorial Demerara Rise.The pre-OAE BP may foreshadow greater regional trends observed during OAE-2. Equatorial upwelling centers, like Demerara Rise, are spatially restricted and represent regions of already high primary production before OAE-2. Climatic shifts concurrent with OAE-2 may have produced favorable conditions for elevated primary productivity in regions unable to capitalize on or exposed to allochthonous nutrient delivery prior to the +CIE. While the pre-OAE BP offers a causal mechanism for the Tl isotope excursion and +CIE initiation, areal expansion of organic carbon preservation and production is necessary to sustain enhanced organic carbon burial for the duration of the +CIE.Continued development of preexisting proxies is critical to extract and clarify current understandings of major climatic events in Earth history. Although reliant on excellent preservation of the microbial signal, the analytical and interpretative approach used here enables simultaneous examination of a wide array of biomarkers, producing a more holistic reconstruction of oceanographic changes inferred from microbial ecological variations spanning the surface to the sediment. This is timely, as investigations of the sedimentary archives become increasingly valuable analogs to understand the response of modern oceans to natural and anthropogenic forcings. Similarities between the pre-OAE BP and modern, climate-driven marine deoxygenation are concerning, while particular attention to preexisting highly productive settings may hold the key to forecasting the geologically rapid transition to a global OAE. Even though natural processes are currently beyond our control, stifling anthropogenic catalysts of climate change may decelerate the unfortunate, progressive suitability of OAEs as climate analogs in the future. More

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We took as a starting point the hake – sole – Norway lobster Bay of Biscay ISIS-Fish database used for COSELMAR project16,20 (see http://isis-fish.org/download.html section “Bay of Biscay scenario dataset”, Database V0 in Fig. 1). This database was built using 2010 data, and was not calibrated, as it was designed for a geo-foresight study. Since our aim was to describe the system over a decade and simulate realistic dynamics close to available observations to assess management measures, we needed to update the parametrisation and calibrate the database. We took 2010–2012 as the calibration period, and 2013–2020 as the simulation period (grey arrow Fig. 1). The database has a monthly temporal resolution (constrained by the ISIS-Fish framework) and the spatial scale was set to match ICES statistical rectangles (0.5° latitude by 1° longitude rectangles, defined by the International Council for the Exploration of the Sea (ICES) https://www.ices.dk/data/maps/Pages/ICES-statistical-rectangles.aspx), consistent with available knowledge and data.In this section, we firstly describe all the data sources used to update and calibrate the database. Then, for each main component of an ISIS-Fish database – i.e. populations, exploitation and management – we describe this paper’s database parameters and assumptions. We finally describe the calibration procedure (inspired by previous work21,22), of which some results are shown in the Technical Validation section. We summarized this workflow in Fig. 1.Data sourcesData sources, estimates, and literature (including grey literature) were needed to update and calibrate the model. They are marked in Fig. 1 with salmon (data sources and estimates) and mustard (literature) blocks:

SACROIS23: French landings and effort logbook declarations for 2010 were made available at the log-event*commercial category*ICES statistical rectangle*population scale. It was used to design exploitation features of the database, as well as populations spatial structure.

LANGOLF survey: 2006–2010 LANGOLF surveys observations for 2006–2010 were made available for Norway lobster. They were used to work on Norway lobster abundance per length class and sex.

Intercatch: catch observations for 2010–2020 in the Bay of Biscay for hake, at the quarter-métier group scale, and catch observations per class for sole on 2010–2012, and 2010 Norway lobster catch observations per sex and length class24, used to describe the inter-annual effort dynamics, to calibrate and validate the model.

Estimates of hake abundance per size class in 2010, and hake quarterly estimates of recruitment on 2010–2012 from a northern hake spatial stock assessment model21, used to inform hake biology assumptions (named Other 1 in Fig. 1).

ICES WGBIE24 2010 estimates of abundance per class (sole and Norway lobster), to inform their abundance at the initial time step; 2010–2012 yearly fishing mortality estimates per age class (sole) to calibrate the database (named Other 2 in Fig. 1).

Other population, exploitation and management assumptions were informed with scientific literature25 and grey literature26,27 (Literature block in Fig. 1).

Management assumptions were informed with legal texts2,4,28,29,30,31,32,33,34 and reported quota values in working group reports24.

About populationsThis section describes for each species the assumptions and parameters values, except for accessibility, which has been calibrated, as described in section Calibration procedure. For all assumptions and values, more details are provided in Supplementary Information’s section 2.2.HakeThe stock size structure was defined with 1 cm size bins for [1;40[cm individuals, 2 cm for [40;100[cm individuals, and 10 cm for [100;130+] cm individuals35. Areas of presence were defined based on 2010 SACROIS French landings data per commercial category and statistical rectangle23, leading to the definition of a presence, a recruitment, an interim recruitment and a spawning area25 (see Supplementary Information’s section 2.2 and Figure S1). These areas allow for the description of intra-Bay of Biscay migrations related to spawning and recruitment processes: mature individuals aggregate at the beginning of the year on the shelf break to spawn, and then disperse on the shelf36,37,38,39,40 (at the beginning of April and July in the model). Also, from age 1 (around 20 cm), individuals in recruitment zone spread in interim recruitment zone, to model a diffusion towards areas neighbouring the nursery area, at the beginning of each time step (see Supplementary Information’s section 2.2 and Table S11). Maturity-at-size and weight-at-length relationships were the same functions as used by ICES working group35,41. Natural mortality was fixed at 0.5, basing on preliminar runs, instead of the commonly used 0.442. Recruitment values were defined prior to the simulation for 2010–2020 using available estimates on the 2010–2015 time series21,27. Deterministic estimates from these sources were allocated to the recruitment area in the Bay of Biscay and the beginning of each month in January-September on the whole time series, of which values are provided in the Supplementary Information’s section 2.2 and Table S3. Growth is modelled through monthly growth increments5,25. However, given the different widths of size bins in the implemented size structure, a correction was provided to values in the transition matrix to eliminate artifacts when growing to a size bin wider than the size bin of origin, as detailed in Supplementary Information’s section 2.2. Abundance at the initial step in each zone was estimated from Bay of Biscay abundance estimates for 201021. Mature individuals over 20 cm were allocated to the spawning area, all individuals strictly shorter than 20 cm were allocated to the recruitment area (as they were assumed to be less than 1 year old), and remaining individuals were allocated to the interim recruitment area. None were allocated to the presence area, in which individuals will go later in the time series, after disaggregating from the spawning area25 (Table S13).SoleThe stock is age structured, with 7 classes going from ages 2 to 7+43 (Table S2). No seasonal variations were implemented. Only a single presence zone was defined (see Supplementary Information’s section 2.2 and Figure S1), as in preliminary runs defining more presence areas for sole did not yield more knowledge in this study. We implemented ICES working group values for natural mortality, weight-at-age (Table S1) and maturity-at-age43. Recruitment occurs at the beginning of each year, individuals being recruited at age 2 (ages 0–1 were not modelled; Table S4). We implemented ICES working group estimates27 for abundance at initial time step (Table S14).Norway lobsterThe stock has a sex-size structure, with 1 mixed recruitment class at 0 cm; 33 length classes for males at 2 carapace length mm intervals, from [10;12[to [72;74[carapace length mm; 23 length classes for females at 2 carapace length mm intervals, from [10;12[to [52;54[carapace length mm. A single presence area was defined: the Great Mudbank21 (see Supplementary Information’s section 2.2 and Figure S1). Several seasonal processes occur for this stock, impacting recruitment, accessibility and growth: 1/ January, begins with the annual recruitment. Females are inside their burrows, less accessible; 2/ February-March females are inside their burrows, less accessible; 3/ April: Spring moulting, females are more accessible; 4–5/ May-August females are more accessible; 6/ September, females are inside their burrows, less accessible; 7/ October: Autumn moulting only for immature females and all males, females are inside their burrows, less accessible; 8/ November-December, females are inside their burrows, less accessible44. We implemented ICES working group values for natural mortality, weight-at-class and maturity-at-class45,46,47. Growth occurs twice a year, when moulting in April and October, and is modelled with growth increments. Recruitment occurs at the beginning of each year, modelled with a Beverton-Holt relationship26, and was assumed to have the same spatial distribution as spawning stock biomass. Abundance at initial step was derived from LANGOLF survey observations and ICES WGBIE estimates25,26 (Table S16).About exploitationThe fishing exploitation structure (fleets, strategies, métiers and gears) were derived following a classification method on SACROIS 2010 landings and effort data13,23 from French fleets, and taken from a TECTAC project (https://cordis.europa.eu/project/id/Q5RS-2002-01291) database for Spanish trawlers. More details on their definition are provided in Supplementary Information’s section 2.3, Tables S5–S9 and S20–S21 and Figure S3. Spanish longliners and gillnetters fleets exploitation was described based on catch (observations from Intercatch48) rather than effort.Hake selectivity and discarding functions (one for each gear) were taken from estimates of a spatial hake stock assessment model21. Parameters values and formulæ are provided in Supplementary Information’s section 2.3 and Tables S6-S7. On top of this, inter-annual fleet dynamics factors were included in equation (21) of ISIS-Fish documentation8 in order to account for observed catch temporal variations. These factors are therefore multiplicative parameters of the target factor of each species for each métier. They are computed using observed catch27 and differ according to the period and targeted species:

over 2010–2016, it is a ratio of observed catch in weight per year over catch observations for 2010: for hake, one per métier *season*year (left(frac{ObservedCatc{h}_{metier,season,year}}{ObservedCatc{h}_{metier,season,2010}}right)), for sole, one per métier *year (left(frac{ObservedCatc{h}_{metier,year}}{ObservedCatc{h}_{metier,2010}}right)), and for Norway lobster, one per year (identical for each métier catching Norway lobster) (left(frac{ObservedCatc{h}_{year}}{ObservedCatc{h}_{2010}}right));

over 2017–2020: at the time of writing these assumptions, more recent data was not available, and ratios were deduced from trends on 2014–2016. A linear model was fitted on ratios deduced earlier on 2014–2016. If a significant trend was identified (hake: whitefish trawlers quarters 2 and 4, longliners and gillnetters seasons 2–3; sole and Norway lobster: all métiers), the slope was used to deduce 2017–2020 ratios (the slope was halved for hake whitefish trawlers and sole and Norway lobster values to avoid unrealistic high values of effort). Otherwise, 2016 ratios were used.

All values are provided in Supplementary Information’s section A.2 Tables S22–S24, and the final values of target factors are derived from the Calibration procedure.About managementWe implemented a set of management rules close to what is currently implemented in the Bay of Biscay.All stocks are managed by TALs (Total Allowable Landings) until 2015 and then by TACs (Total Allowable Catch), except for Norway lobster, managed by TALs on the whole time series, not being under the landings obligation. To favour a better parametrisation, allowing for more reliable dynamics on the following years of the time series, no TALs were implemented during the calibration period (2010–2012; Fig. 1). These regulations were implemented from 2013 using historically TALs and TACs values24.Landings of the three stocks are also constrained by a Minimum Conservation Reference Size regulation that was implemented for all stocks using values currently enforced in the studied fishery28. Likewise, from 2016, the Landings Obligation was implemented, with de minimis exemptions for hake and sole, depending on the year and the gear used to fish them2,31,32,33,34. See Supplementary Information’s sections 2.4 and A.3, Figure S2 and Table S10 for further details on these restrictions.In response to the above management rules, a fishers’ behaviour algorithm has been developed to describe fishermen adaptation. Some métiers may be forbidden, depending on some conditions – the catch quota has been reached, the landings obligation is enforced – but also some values – the proportion of discarded catch, and also catch on previous years. Therefore fishermen change métiers within their strategy métiers set through a re-allocation of fishing effort to the latter set. This re-allocation aims to avoid quota overshooting. Further details about this algorithm are provided in the Supplementary Information’s sections 2.4 and A.3 and Figure S2.Calibration procedureThe model has been calibrated using two parameters (population accessibility and fishing target factor) involved in the catchability process (equation (21) in ISIS-Fish documentation8). The objective of the calibration is to reproduce the dynamics of catch over 2010–2012 at the species*métiers group scale, for each year or quarter depending on available data’s granularity. Calibration is sequentially performed: accessibility parameters for each population were estimated first followed by the target factors. The estimation of each parameter set (parameter type * population) combination was separated, and values were estimated jointly within each parameter set. To account for the specificity of each population model dynamics (global age-based for sole, spatial and size-based for hake, spatial, sex and size-based for Norway lobster), an objective function is defined for each population to calibrate their accessibility. More details on objective functions and procedures are provided in Supplementary Information’s section 2.5, as well as estimated values in Tables S17–S19.Hake accessibilityThe calibration for hake accessibility is based on a procedure developed for a former version of the database25. One parameter was estimated per quarter, all values being equal across length classes. The model outputs were fitted to hake catch observations in weight in the Bay of Biscay in 2010–2012 per length class.Sole accessibilityOne parameter was estimated per age class. The model outputs were fitted to WGBIE fishing mortality per age class for sole27 in 2010–2012.Norway lobster accessibilityOne parameter was calibrated per sex and length class. The model outputs were fitted to catch in numbers per length class and sex in 2010 per quarter provided by WGBIE.About target factorsTarget factors drive how the effort is distributed between populations, métiers and season*year combinations. They were split in 3 components: a fixed component derived from the SACROIS effort dataset analysis (Tables S25–S27), another fixed component driving inter-annual variations of fishing effort (Tables S22–S24), derived from catch observations, and finally an estimated component (Tables S28–S30), allowing to tune the model’s dynamics to observed catch. This section focuses on the estimation of the latter.Hake target factors20 parameters were defined, for each combination of the 5 groups of métiers (longliners, gillnetters, whitefish trawler (coastal), whitefish trawler (not coastal), Norway lobster trawler, see definition Table S8) and 4 quarters. We fitted the model’s outputs to the same data and with the same objective function as for hake’s accessibilities estimation.Sole target factors1 estimated component per group of métiers (gillnetters, Norway lobster trawlers and whitefish trawlers) and quarter. We fitted the model’s outputs to sole catch in weight on 2010–2012 for each métier and quarter.Norway lobster target factors1 estimated component per group of métiers (Norway lobster trawlers and whitefish trawlers). We fitted the model’s outputs to monthly Norway lobster landings data per length and sex class for 2010.Base simulationThe base simulation ran from January 2010 to December 2020 inclusive, with a monthly time step, using the database and parameters values described in this document. Several outputs of interest may be explored after a run: catch (discards and landings), as done in several figures in this paper, but also biomass (total biomass or mature biomass), fishing mortality values, or effort, all at a fine spatio-temporal scale. More

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