Marcos Llope

Publications

Llope M, Blenckner T, Vasilakopoulos P, McGinty N, Lynam CP, Helaouët P, Möllmann C, Frelat R, Durant JM, Stige LC, Stenseth NC (2026) Stress and resilience in northern European marine ecosystems. Proceedings of the National Academy of Sciencies, http://www.pnas.org/doi/10.1073/pnas.2527939123 

Seck M, Kloff S, Baldé BS, M’Bengue B, Llope M (2026) Offshore gas in West Africa: assessment of impacts and redistribution of pressures. Frontiers in Ocean Sustainability, https://doi.org/10.3389/focsu.2026.1702958

Cabrera-Busto J, Ramírez-Romero E, Buttay L, Llope M (2026) Regionalisation and multidecadal change in the southern Canary Current Large Marine Ecosystem. Journal of Geophysical Research – Oceans, https://doi.org/10.1029/2025JC023075

Cabrera-Busto J, Buttay L, Ramírez-Romero E, Llope M, Planque B (2026) Causal modelling of fishing and upwelling impact on Sardinella aurita and Sardina pilchardus in the Canary Current Upwelling. ICES Journal of Marine Science, https://doi.org/10.1093/icesjms/fsaf242

Llope M, Diop B, Kloff S (2025) Fisheries, non-renewables and migration in West Africa, the need for an integrated socio-ecosystem-based management. ICES Journal of Marine Science, https://doi.org/10.1093/icesjms/fsaf193.

Feng J, Wan X, Wang R, Li S, Wei M, Dakos V, Llope M, Lu X, Stenseth NC (2025) Mediating role of food web structure in linking diversity to multidimensional stability: Evidence from global marine ecosystems. Science Advances, https://www.science.org/doi/10.1126/sciadv.adv3841

Smith P, Ballal VP, Cherubini F, Díaz-José J, Duchková H, Gupta H, Ito A, Khan S, Llope M, Hori M, Tirado MC, Tourinho L, Vale MM, Xu X, Chudasama H, Eriksen SH, Mason-D’Croz D, Phang SC, Srivastava Y, Harrison PA, McElwee PD, van Huysen T, Ricketts T, Herrero M, Singh PK (2025) Impacts of climate change interventions on biodiversity, water, the food system and human health and well-being. Global Change Biology, https://doi.org/10.1111/gcb.70444

García-de-Vinuesa A, Florido D, Vilas C, Torres MA, Delgado M, Muñoz I, Cabrera-Castro R, Ramos F, Llope M (2025) Framing of social systems for ecosystem-based management: The Guadalquivir estuary-Gulf of Cadiz coupled socio-ecosystem as case study. Environmental Development, https://doi.org/10.1016/j.envdev.2025.101206

Maureaud AA, et al. (2025) FISHGLOB: A collaborative infrastructure to bridge the gap between scientific monitoring and marine biodiversity conservation. Conservation Science and Practice, https://doi.org/10.1111/csp2.70035

Singh PK, Smith P, Cherubini F, Díaz-José J, Duchková H, Gupta H, Hori M, Ito A, Khan S, Llope M, Tirado MC, Vale MM, Xu X (2024). Chapter 5: Options for delivering sustainable biodiversity-related approaches to climate change, adaptation and mitigation, including relevant aspects of the energy system. In: Thematic Assessment of the Interlinkages among Biodiversity, Water, Food and Health of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. Harrison PA, McElwee PD, van Huysen TL (eds.). IPBES secretariat, Bonn, Germany.

Soto M, Fernández-Peralta L, Rey J, Czewinski I, García-Cancela R, Llope M, Cabrera-Busto J, Liébana M, Meissa B, Pennino MG (2023) Modelling drivers of trawl fisheries discards using Bayesian spatio-temporal models. Fisheries Research, https://doi.org/10.1016/j.fishres.2023.106830

Carvalho-Souza GF, Torres MA, Farias C, Acosta JJ, Tornero J, Sobrino I, Ramos F, Llope M (2021) International politics must be considered together with climate and fisheries regulation as a driver of marine ecosystems. Global Environmental Change, https://doi.org/10.1016/j.gloenvcha.2021.102288

van Denderen PD, Holah H, Robson LM, Hiddink JG, Menot L, Pedreschi D, Kazanidis G, Llope M, Turner PJ, Stirling D, Murillo FJ, Kenny A, Campbell N, Allcock AL, Braga-Henriques A, González-Irusta JM, Johnston G, Orejas C, Serrano A, Xavier JR, Hopkins P, Kenchington E, Nixon E, Valanko S (2021) A Policy-Based Framework for the Determination of Management Options to Protect Vulnerable Marine Ecosystems under the EU Deep-sea Access Regulations. ICES Journal of Marine Science, https://doi.org/10.1093/icesjms/fsab237

Bastardie F, Brown EJ, Andonegi E, Arthur R, Beukhof E, Depestele J, Döring R, Eigaard, OR, García-Barón I, Llope M, Mendes H, Piet G, Reid D (2021) A Review characterizing 25 ecosystem challenges to be addressed by an Ecosystem Approach to Fisheries Management in Europe. Frontiers in Marine Science, https://doi.org/10.3389/fmars.2020.629186

Llope M, Carvalho-Souza GF, Baldó F, González-Cabrera C, Jiménez MP, Licandro P, Vilas C (2020) Gulf of Cadiz zooplankton: community structure, zonation and temporal variation. Progress in Oceanography, https://doi.org/10.1016/j.pocean.2020.102379

Carvalho-Souza GF, González-Ortegón E, Baldó F, Vilas C, Drake P, Llope M (2018) Natural and anthropogenic drivers on the early life stages of European anchovy in one of its Essential Fish Habitats, the Guadalquivir estuary. Marine Ecology Progress Series, https://doi.org/10.3354/meps12562

Carvalho-Souza GF, Llope M, Tinôco MS, Medeiros DV, Maia-Nogueira R, Sampaio CLS (2018) Marine litter disrupts ecological processes in reef systems. Marine Pollution Bulletin, https://doi.org/10.1016/j.marpolbul.2018.05.049

Lynam CP, Llope M, Möllmann C, Helaouët P, Bayliss-Brown GA, Stenseth NC (2017) Interaction between top-down and bottom-up control in marine food webs. Proceedings of the National Academy of Sciences, https://doi.org/10.1073/pnas.1621037114

Llope M (2017) The ecosystem approach in the Gulf of Cadiz. A perspective from the southernmost European Atlantic regional sea. ICES Journal of Marine Science, https://doi.org/10.1093/icesjms/fsw165

Blenckner T, Llope M, Möllmann C, Voss R, Quaas MF, Casini M, Lindegren M, Folke C, Stenseth NC (2015) Climate and fishing steer ecosystem regeneration to uncertain economic futures. Proceedings of the Royal Society B: Biological Sciences, https://doi.org/10.1098/rspb.2014.2809

Jianfeng F, Durant JM, Stige LC, Hessen DO, Hjermann DØ, Zhu L, Llope M, Stenseth NC (2015) Contrasting correlation patterns between environmental factors and chlorophyll levels in the global ocean. Global Biogeochemical Cycle, https://doi.org/10.1002/2015GB005216

Jianfeng F, Stige LC, Durant JM, Hessen DO, Zhu L, Hjermann DØ, Llope M, Stenseth NC (2014) Large-scale season-dependent effects of temperature and zooplankton on phytoplankton in the North Atlantic. Marine Ecology Progress Series, https://doi.org/10.3354/meps10724

Otto SA, Kornilovs G, Llope M, Möllmann C (2014) Interactions among density, climate, and food web effects determine long-term life cycle dynamics of a key copepod. Marine Ecology Progress Series, https://doi.org/10.3354/meps10613

Nicolas D, Rochette S, Llope M, Licandro P (2014) Spatio-temporal variability of the North Sea cod recruitment in relation to temperature and zooplankton. PLoS ONE, https://doi.org/10.1371/journal.pone.0088447

Raab K, Llope M, Nagelkerke LAJ, Rijnsdorp AD, Teal LR, Licandro P, Ruardij P, Dickey-Collas M (2013) Influence of temperature and food availability on juvenile European anchovy Engraulis encrasicolus at its northern boundary. Marine Ecology Progress Series, https://doi.org/10.3354/meps10408

Casini M, Blenckner T, Möllmann C, Gårdmark A, Lindegren M, Llope M, Kornilovs G, Plikshs M, Stenseth NC (2012) Predator transitory spillover induces trophic cascades in ecological sinks. Proceedings of the National Academy of Sciences, https://doi.org/10.1073/pnas.1113286109

Llope M, Licandro P, Chan K-S, Stenseth NC (2012) Spatial variability of the plankton trophic interaction in the North Sea: a new feature after the early 1970s. Global Change Biology, https://doi.org/10.1111/j.1365-2486.2011.02492.x

Cardinale M, Bartolino V, Llope M, Maiorano L, Sköld M, Jacob Hågberg J (2011) Historical spatial baselines in conservation and management of marine resources. Fish and Fisheries, https://doi.org/10.1111/j.1467-2979.2010.00393.x

Llope M, Daskalov GM, Rouyer TA, Mihneva V, Grishin AN, Chan K-S, Stenseth NC (2011) Overfishing of top predators eroded the resilience of the Black Sea system regardless of the climate and anthropogenic conditions. Global Change Biology, https://doi.org/10.1111/j.1365-2486.2010.02331.x

Olsen EM, Ottersen G, Llope M, Chan K-S, Beaugrand G, Stenseth NC (2011) Spawning stock and recruitment in North Sea cod shaped by food and climate. Proceedings of the Royal Society B: Biological Sciences, https://doi.org/10.1098/rspb.2010.1465

Iriarte A, Aravena G, Villate F, Uriarte I, Ibañez B, Llope M, Stenseth NC (2010) Dissolved oxygen in contrasting estuaries of the Bay of Biscay: effects of temperature, river discharge and chlorophyll a. Marine Ecology Progress Series, https://doi.org/10.3354/meps08812

Llope M, Chan K-S, Ciannelli L, Reid PC, Stige L, Stenseth NC (2009) Effects of environmental conditions on the seasonal distribution of phytoplankton biomass in the North Sea. Limnology and Oceanography, https://doi.org/10.4319/lo.2009.54.2.0512

Payne MR, Hatfield EMC, Dickey-Collas M, Falkenhaug T, Gallego A, Gröger J, Licandro P, Llope M, Munk P, Röckmann C, Schmidt JO, Nash RDM (2009) Recruitment in a changing environment: the 2000s North Sea herring recruitment failure. ICES Journal of Marine Science, https://doi.org/10.1093/icesjms/fsn211

Valdés L, López-Urrutia A, Cabal J, Álvarez-Ossorio M, Bode A, Miranda A, Cabanas M, Huskin I, Anadón R, Alvarez-Marqués F, Llope M, Rodríguez N (2007) A decade of sampling in the Bay of Biscay: What are the zooplankton time series telling us?. Progress in Oceanography, https://doi.org/10.1016/j.pocean.2007.04.016

Llope M, Anadón R, Sostres JA, Viesca L (2007) Nutrients dynamics in the southern Bay of Biscay (1993-2003): Winter supply, stoichiometry, long-term trends and their effects on the phytoplankton community. Journal of Geophysical Research, https://doi.org/10.1029/2006JC003573

Stenseth NC, Llope M, Anadón R, Ciannelli L, Chan K-S, Hjermann DØ, Bagøien E, Ottersen G (2006) Seasonal plankton dynamics along a cross-shelf gradient. Proceedings of the Royal Society B: Biological Sciences, https://doi.org/10.1098/rspb.2006.3658

Llope M, Anadón R, Viesca L, Quevedo M, González-Quirós R, Stenseth NC (2006) Hydrography of the southern Bay of Biscay shelf-break region: integrating the multi-scale physical variability over the period 1993-2003. Journal of Geophysical Research, https://doi.org/10.1029/2005JC002963

Isla JA, Llope M, Anadón R (2004) Size-fractionated mesozooplankton biomass, metabolism and grazing along a 50ºN to 30ºS transect of the Atlantic Ocean. Journal of Plankton Research, https://doi.org/10.1093/plankt/fbh121

• Cabrera-Busto, Juncal; Ramírez-Romero, Eduardo; Buttay, Lucie & Peri, Marcos Llope (2026). Regionalization and Multidecadal Change in the Southern Canary Current Large Marine Ecosystem. Journal of Geophysical Research (JGR): Oceans. ISSN 2169-9275. 131(5). doi: 10.1029/2025JC023075. Full text in Research Archive Show summary

  The current delineation of the southern Canary Current Large Marine Ecosystem fisheries management zones responds only to political divisions rather than ecological features or a combination of both. In this paper, we implemented three time series analysis techniques on spatially explicit daily and monthly resolved data sets from satellite and reanalysis. Methods are based on climatologies, wavelet analysis and clusters and seasonal regression models, thus covering different aspects of the upwelling dynamics. Due to the overall convergence of the outputs of the analysis, by describing the spatiotemporal dynamics of oceanographic processes and biogeochemical features, we delineate distinct robust subregions and examine their recent trends. Our results identified four subregions: (a) a tropical subregion (12–18–21°N), (b) a Western Sahara (18–21°N to 25°N), (c) a South Morocco (26–28°N) and (d) a North Morocco subregion (29–35°N). The tropical subregion continues to exhibit a highly seasonal upwelling pattern with decreasing duration of winter upwelling events, in contrast with the other subregions that show slight variations in their seasonality. Although wind-driven upwelling has intensified in Western Sahara, leading to a cooling trend in sea surface temperature, chlorophyll-a does not show a concomitant increase. The North Morocco subregion is experiencing an increase in both the intensity and duration of the upwelling, making it more closely resemble the permanent upwelling zone of Western Sahara. This revisited subdivision provides a robust framework for ecosystem-based management by incorporating current ecological variability. Monitoring of hydrographic and biological trends is essential to adapt boundaries to future conditions.

• Cabrera-Busto, Juncal; Buttay, Lucie; Ramírez-Romero, Eduardo; Peri, Marcos Llope & Planque, Benjamin (2026). Unraveling fishing and upwelling impacts on Sardinella aurita and Sardina pilchardus in the Canary Current Upwelling. ICES Journal of Marine Science. ISSN 1054-3139. 83(4). doi: 10.1093/icesjms/fsaf242. Full text in Research Archive Show summary

  In the Canary Current Large Marine Ecosystem (CCLME), significant variations in upwelling dynamics have been observed in the last decades, but their impact on small pelagic fish biomass remains unclear. Identifying the factors driving these stock variations is challenging, particularly for heavily overexploited species. In this study, we attempt to disentangle the influence of fishing and environmental factors on the biomass of Sardinella aurita and Sardina pilchardus using structural causal models. Fisheries data alone cannot explain the year-to-year variations in biomass, indicating that other factors are at play. Furthermore, unrealistic results using fishing data point to high levels of illegal, unreported, and unregulated catches, leading to an underestimation of S. aurita biomass. In the tropical subregion (southernmost CCLME), fluctuations in the biomass of S. aurita are primarily driven by upwelling dynamics rather than temperature or salinity. A similar pattern is observed in the Western Sahara subregion, where primary productivity, along with the previous year’s biomass of S. aurita, is the main driver of biomass fluctuations. Additionally, a negative relationship between S. aurita biomass in the two subregions suggests that the Western Sahara may serve as a refuge during periods of declining suitable habitat in the tropical subregion. In the Western Sahara subregion, fluctuations in S. pilchardus biomass are influenced by primary productivity, while in the South Morocco subregion the driving factor is likely temperature. The results highlight the significant influence of environmental variables on pelagic fish biomass, underscoring the potential impacts of climate change on the distribution and abundance of these species. Such shifts could profoundly affect local economies and food security, while also heightening geopolitical tensions among nations competing for shared marine resources. Comprehensive evaluation and effective management strategies that account for environmental changes are urgently required.

• Peri, Marcos Llope; Blenckner, Thorsten; Vasilakopoulos, Paraskevas; McGinty, Niall; Lynam, Christopher P. & Hélaouët, Pierre [Show all 11 contributors for this article] (2026). Stress and resilience in northern European marine ecosystems. Proceedings of the National Academy of Sciences of the United States of America (PNAS). ISSN 0027-8424. 123(8), p. 1–7. doi: 10.1073/pnas.2527939123. Full text in Research Archive Show summary

  Marine ecosystems are facing multiple pressures from human activities, such as fishing and nutrient inputs from farming and agriculture, compounded by the global effects of climate change. As a result, the ecosystem services that societies depend on are at risk from the cumulative impact of these pressures. To better understand how different ecosystems respond, it is essential to assess their resilience. While predicting ecosystem resilience remains challenging, significant progress has been made in developing methods to measure it. In this study, we assessed resilience in four northern European marine ecosystems: the Icelandic Waters, the Barents Sea, the Baltic Sea, and the North Sea, spanning 3 to 5 decades. “Folded stability landscapes” were constructed for the Baltic and North Sea. This region provides a unique setting, with a south-north gradient from temperate to subarctic environments and a southeast-northwest gradient from open to semienclosed topographies. The Icelandic Waters and Barents Sea evolved relatively continuously, while the Baltic Sea and North Sea underwent more drastic changes. By comparing results across these gradients, we explore the role of isolation, level of pressures, and food web complexity in shaping resilience patterns and discuss the implications for managing resilient marine ecosystems in the future.

• Feng, Jianfeng; Wan, Xuhao; Wang, Ruyue; Li, Shengpeng; Wei, Maohong & Dakos, Vasilis [Show all 9 contributors for this article] (2025). Mediating role of food web structure in linking diversity to multidimensional stability: Evidence from global marine ecosystems. Science Advances. 11(41). doi: 10.1126/sciadv.adv3841. Full text in Research Archive Show summary

  The diversity-stability relationship persists as a central ecological debate, as existing work primarily examines direct diversity and structural drivers, while empirical evidence clarifying how direct and structure-mediated pathways jointly shape ecosystem multistability remains limited. Here, we analyzed 217 global marine food webs, quantifying structural properties and multidimensional stability to evaluate how diversity and structure jointly influence stability. Our analyses reveal that diversity is consistently linked to stability through dual pathways: positively associated with resistance and resilience via indirect structural mediation, yet negatively correlated with local stability unless interaction strength is accounted for. Critically, omitting structural mediation yields a net negative diversity-stability correlation, whereas integrating food web structure uncovers context-dependent positive relationships, underscoring structural metrics as pivotal explanatory variables. Our findings reconcile the diversity-stability debate by showing that the food web structure mediates context-dependent stability outcomes—integrating direct and indirect pathways resolves contradictions and advances actionable metrics for conservation strategies resilient to environmental changes.

• Smith, Pete; Singh, Pramod K.; Ballal, Vedant Pushpahas; Cherubini, Francesco; Díaz-José, Julio & Duchková, Helena [Show all 25 contributors for this article] (2025). Impacts of Climate Change Interventions on Biodiversity, Water, the Food System and Human Health and Well-Being. Global Change Biology. ISSN 1354-1013. 31(9), p. 1–28. doi: 10.1111/gcb.70444. Full text in Research Archive
• Denderen, P. Daniël Van; Holah, Helen; Robson, Laura M.; Hiddink, Jan Geert; Menot, Lénaick & Pedreschi, Debbi [Show all 24 contributors for this article] (2022). A policy-based framework for the determination of management options to protect vulnerable marine ecosystems under the EU deep-sea access regulations. ICES Journal of Marine Science. ISSN 1054-3139. 79(1), p. 34–49. doi: 10.1093/icesjms/fsab237. Full text in Research Archive
• Maureaud, Aurore; Frelat, Romain; Pecuchet, Laurène Anne Marie; Shackell, Nancy; Merigot, Bastien & Pinsky, Malin L. [Show all 75 contributors for this article] (2020). Are we ready to track climate‐driven shifts in marine species across international boundaries? ‐ A global survey of scientific bottom trawl data. Global Change Biology. ISSN 1354-1013. 27(2), p. 220–236. doi: 10.1111/gcb.15404. Full text in Research Archive Show summary

  Marine biota are redistributing at a rapid pace in response to climate change and shifting seascapes. While changes in fish populations and community structure threaten the sustainability of fisheries, our capacity to adapt by tracking and projecting marine species remains a challenge due to data discontinuities in biological observations, lack of data availability, and mismatch between data and real species distributions. To assess the extent of this challenge, we review the global status and accessibility of ongoing scientific bottom trawl surveys. In total, we gathered metadata for 283,925 samples from 95 surveys conducted regularly from 2001 to 2019. We identified that 59% of the metadata collected are not publicly available, highlighting that the availability of data is the most important challenge to assess species redistributions under global climate change. Given that the primary purpose of surveys is to provide independent data to inform stock assessment of commercially important populations, we further highlight that single surveys do not cover the full range of the main commercial demersal fish species. An average of 18 surveys is needed to cover at least 50% of species ranges, demonstrating the importance of combining multiple surveys to evaluate species range shifts. We assess the potential for combining surveys to track transboundary species redistributions and show that differences in sampling schemes and inconsistency in sampling can be overcome with spatio‐temporal modeling to follow species density redistributions. In light of our global assessment, we establish a framework for improving the management and conservation of transboundary and migrating marine demersal species. We provide directions to improve data availability and encourage countries to share survey data, to assess species vulnerabilities, and to support management adaptation in a time of climate‐driven ocean changes.

• Carvalho-Souza, Gustavo F. de; González-Ortegón, Enrique; Baldó, Francisco; Vilas, César; Drake, Pilar & Llope, Marcos (2018). Natural and anthropogenic effects on the early life stages of European anchovy in one of its essential fish habitats, the Guadalquivir estuary. Marine Ecology Progress Series. ISSN 0171-8630. p. 1–13. doi: 10.3354/meps12562. Full text in Research Archive
• Lynam, Christopher Philip; Llope, Marcos; Möllmann, Christian; Helaouët, Pierre; Bayliss-Brown, Georgia Anne & Stenseth, Nils Christian (2017). Interaction between top-down and bottom-up control in marine food webs. Proceedings of the National Academy of Sciences of the United States of America (PNAS). ISSN 0027-8424. 114(8), p. 1952–1957. doi: 10.1073/pnas.1621037114. Full text in Research Archive
• Llope, Marcos (2016). The ecosystem approach in the Gulf of Cadiz. A perspective from the southernmost European Atlantic regional sea. ICES Journal of Marine Science. ISSN 1054-3139. 74(1), p. 382–390. doi: 10.1093/icesjms/fsw165. Full text in Research Archive
• Blenckner, Thorsten; Llope, Marcos; Möllmann, Christian; Voss, Rudi; Quaas, Martin & Casini, Michele [Show all 9 contributors for this article] (2015). Climate and fishing steer ecosystem regeneration to uncertain economic futures. Proceedings of the Royal Society of London. Biological Sciences. ISSN 0962-8452. 282(1803), p. 1–9. doi: 10.1098/rspb.2014.2809. Full text in Research Archive
• Llope, Marcos; Licandro, Priscilla; Chan, Kung-Sik & Stenseth, Nils Christian (2012). Spatial variability of the plankton trophic interaction in the North Sea: a new feature after the early 1970s. Global Change Biology. ISSN 1354-1013. 18(1), p. 106–117. doi: 10.1111/j.1365-2486.2011.02492.x. Full text in Research Archive Show summary

  Traditionally, marine ecosystem structure was thought to be bottom-up controlled. In recent years, a number of studies have highlighted the importance of top-down regulation. Evidence is accumulating that the type of trophic forcing varies temporally and spatially, and an integrated view – considering the interplay of both types of control – is emerging. Correlations between time series spanning several decades of the abundances of adjacent trophic levels are conventionally used to assess the type of control: bottom-up if positive or top-down if this is negative. This approach implies averaging periods which might show time-varying dynamics and therefore can hide part of this temporal variability. Using spatially referenced plankton information extracted from the Continuous Plankton Recorder, this study addresses the potential dynamic character of the trophic structure at the planktonic level in the North Sea by assessing its variation over both temporal and spatial scales. Our results show that until the early-1970s a bottom-up control characterized the base of the food web across the whole North Sea, with diatoms having a positive and homogeneous effect on zooplankton filter-feeders. Afterwards, different regional trophic dynamics were observed, in particular a negative relationship between total phytoplankton and zooplankton was detected off the west coast of Norway and the Skagerrak as opposed to a positive one in the southern reaches. Our results suggest that after the early 1970s diatoms remained the main food source for zooplankton filter-feeders east of Orkney–Shetland and off Scotland, while in the east, from the Norwegian Trench to the German Bight, filter-feeders were mainly sustained by dinoflagellates.

• Casini, Michele; Blenckner, Thorsten; Möllmann, Christian; Gårdmark, Anna; Lindegren, Martin & Llope, Marcos [Show all 9 contributors for this article] (2012). Predator transitory spillover induces trophic cascades in ecological sinks. Proceedings of the National Academy of Sciences of the United States of America (PNAS). ISSN 0027-8424. 109(21), p. 8185–8189. doi: 10.1073/pnas.1113286109. Full text in Research Archive
• Cardinale, Massimiliano; Bartolino, Valerio; Llope, Marcos; Maiorano, Luigi; Sköld, Mattias & Hagberg, Jacob (2011). Historical spatial baselines in conservation and management of marine resources. Fish and Fisheries. ISSN 1467-2960. 12(3), p. 289–298. doi: 10.1111/j.1467-2979.2010.00393.x. Full text in Research Archive
• Olsen, Esben Moland; Ottersen, Geir; Llope, Marcos; Chan, Kung-Sik; Beaugrand, Grégory & Stenseth, Nils Christian (2011). Spawning stock and recruitment in North Sea cod shaped by food and climate. Proceedings of the Royal Society of London. Biological Sciences. ISSN 0962-8452. 278(1705), p. 504–510. doi: 10.1098/rspb.2010.1465. Full text in Research Archive Show summary

  In order to provide better fisheries management and conservation decisions, there is a need to discern the underlying relationship between the spawning stock and recruitment of marine fishes, a relationship which is influenced by the environmental conditions. Here, we demonstrate how the environmental conditions (temperature and the food availability for fish larvae) influence the stock–recruitment relationship and indeed what kind of stock–recruitment relationship we might see under different environmental conditions. Using unique zooplankton data from the Continuous Plankton Recorder, we find that food availability (i.e. zooplankton) in essence determines which model applies for the once large North Sea cod (Gadus morhua) stock. Further, we show that recruitment is strengthened during cold years and weakened during warm years. Our combined model explained 45 per cent of the total variance in cod recruitment, while the traditional Ricker and Beverton–Holt models only explained about 10 per cent. Specifically, our approach predicts that a full recovery of the North Sea cod stock might not be expected until the environment becomes more favourable.

• Llope, Marcos; Daskalov, Georgi; Rouyer, Tristan Alexander; Mihneva, Vesselina; Chan, Kung-Sik & Grishin, Alexander [Show all 7 contributors for this article] (2011). Overfishing of top predators eroded the resilience of the Black Sea system regardless of the climate and anthropogenic conditions. Global Change Biology. ISSN 1354-1013. 17(3), p. 1251–1265. doi: 10.1111/j.1365-2486.2010.02331.x. Full text in Research Archive
• Iriarte, Arantza; Aravena, Guillermo Hernan; Villate, Fernando; Uriarte, Ibon; Ibanez, Berta & Llope, Marcos [Show all 7 contributors for this article] (2010). Dissolved oxygen in contrasting estuaries of the Bay of Biscay: effects of temperature, river discharge and chlorophyll a. Marine Ecology Progress Series. ISSN 0171-8630. 418, p. 57–71. doi: 10.3354/meps08812. Full text in Research Archive Show summary

  The effects of environmental factors on variations of dissolved oxygen (DO) saturation was assessed in 2 estuaries (Bilbao and Urdaibai) in the Bay of Biscay with contrasting hydrography and anthropogenic impact, for the period 1997 to 2006, using generalised additive models (GAMs) and generalised additive mixed models (GAMMs). Spatial and temporal variations in DO saturation and the current severity and extent of hypoxia were higher in the culturally eutrophicated and stratified Bilbao estuary. Seasonal patterns of DO saturation and the hierarchy of factors controlling them differed both between estuaries and between salinity zones. In the inner Bilbao estuary, the main factor explaining DO-saturation variations was temperature, with a dominant effect on oxygen consumption processes. In contrast, in the outer Bilbao estuary, the main factor explaining DO-saturation variations was chlorophyll a, suggesting that the effect of oxygen production through photosynthesis was stronger than that of oxygen consumption. In the inner Urdaibai estuary, both a negative relationship with temperature and a positive relationship with chlorophyll a were detected. The outer Urdaibai estuary showed strong water-column mixing and very high tidal flushing, and ventilation processes were more important than oxygen production or consumption processes. River discharge affected DO saturation only at the inner waters, but when the effects of several variables on DO-saturation variability were tested in an additive model, river discharge was excluded from the models. The effects of temperature-dependent processes on DO-saturation dynamics decreased from the inner to the outer zones of both estuaries.

• Llope, Marcos; Chan, KS; Ciannelli, L; Reid, PC; Stige, Leif Christian & Stenseth, Nils Christian (2009). Effects of environmental conditions on the seasonal distribution of phytoplankton biomass in the North Sea. Limnology and Oceanography. ISSN 0024-3590. 54(2), p. 512–524. Full text in Research Archive
• Payne, MR; Hatfield, EMC; Dickey-Collas, M; Falkenhaug, T; Gallego, A & Groger, J [Show all 12 contributors for this article] (2009). Recruitment in a changing environment: the 2000s North Sea herring recruitment failure. ICES Journal of Marine Science. ISSN 1054-3139. 66(2), p. 272–277. doi: 10.1093/icesjms/fsn211. Full text in Research Archive

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• Maureaud, Aurore A.; Kitchel, Zoë; Fredston, Alexa; Guralnick, Robert; Palacios-Abrantes, Juliano & Palomares, Maria L. D. [Show all 63 contributors for this article] (2025). FISHGLOB: A collaborative infrastructure to bridge the gap between scientific monitoring and marine biodiversity conservation. Conservation Science and Practice (CSP). doi: 10.1111/csp2.70035. Full text in Research Archive
• Llope, Marcos; Blenckner, Thorsten; Vasilakopoulos, Paris; McGinty, Niall; Lynam, Christopher Philip & Helaouët, Pierre [Show all 10 contributors for this article] (2018). Resilience of northeast Atlantic marine ecosystems. Full text in Research Archive
• Llope, Marcos; Blenckner, Thorsten; Vasilakopoulos, Paris; McGinty, Niall; Lynam, Christopher Philip & Helaouët, Pierre [Show all 10 contributors for this article] (2018). Continuous and abrupt changes in the resilience of northeast Atlantic marine ecosystems. Full text in Research Archive
• Llope, Marcos; Blenckner, Thorstein; McGinty, Niall; Stige, Leif Christian; Marteinsdóttir, Gudrun & Stenseth, Nils Christian (2017). A look at the Arctic ecosystems time series can help us project their future. Full text in Research Archive
• Llope, Marcos (2016). Cod response to past and current warm phases in the Seas of Iceland, a time series analysis. Full text in Research Archive
• Llope, Marcos (2012). Regime shifts and resilience in a eutrophic marine ecosystem: the Kattegat as a case study for ecosystem management. Full text in Research Archive
• Llope, Marcos (2012). Resilience and thresholds in the Baltic Sea ecosystem. Full text in Research Archive
• Llope, Marcos (2012). The Baltic Sea regime shift, can it flip back? Full text in Research Archive
• Llope, Marcos; Licandro, P; Chan, Kung-Sik & Stenseth, Nils Christian (2009). Spatio-temporal shift in the plankton trophic interactions in the North Sea over the last 50 years. Full text in Research Archive
• Iriarte, A; Aravena, G; Uriarte, I; Villate, F; Llope, Marcos & Stenseth, Nils Christian (2009). Modelling the effect of temperature and chlorophyll - a on dissolved oxygen in two contrasting estuaries of the Bay of Biscay. Full text in Research Archive
• Llope, Marcos; Daskalov, GM; Rouyer, Tristan Alexander; Chan, Kung-Sik & Stenseth, Nils Christian (2009). Predicting regime shifts under different scenarios in the Black Sea. Full text in Research Archive
• Llope, Marcos (2009). Assessing the stability of trophic relationships in marine food webs, the Black and North Seas as case studies. Full text in Research Archive
• Dingsør, Gjert Endre; Huse, G & Llope, Marcos (2009). Spatial distribution and population dynamics of herring and cod in the North Sea. Full text in Research Archive
• Cardinale, M; Bartolino, V; Llope, Marcos; Maiorano, L; Skold, M & Hågberg, J (2009). Historical distribution (1901-2007) of Plaice in the Kattegat-Skagerrak and implications for stock management and conservation. Full text in Research Archive
• Llope, Marcos; Licandro, P; Chan, Kung-Sik & Stenseth, Nils Christian (2009). Spatio-temporal shift in the plankton trophic interactions in the North Sea over the last 50 years. Full text in Research Archive
• Llope, Marcos (2008). Modelling the effect of the North Sea regime shift on the distribution of plankton functional groups and biomass. Full text in Research Archive
• Llope, Marcos (2008). Is a changing North Sea environment making sustainable exploitation of herring more difficult? Full text in Research Archive
• Llope, Marcos (2008). Sea surface warming in the southern Bay of Biscay modulated by oceanic advection. Full text in Research Archive
• Llope, Marcos (2008). Modelling the effect of the environmental condition on the seasonal distribution of chlorophyll in the North Sea. Full text in Research Archive
• Llope, Marcos (2008). Investigating a possible link between zooplankton changes and recent poor recruitment of herring in the North Sea. Full text in Research Archive
• Llope, Marcos (2008). The effect of the North Sea regime shift on the distribution of plankton functional groups and biomass. Full text in Research Archive

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