Ecology

Keenan, T. F. & Williams, C. A. The terrestrial carbon sink. Annu. Rev. Environ. Resour. 43, 219–243 (2018).Article 

Google Scholar 
Terrer, C. et al. A trade-off between plant and soil carbon storage under elevated CO2. Nature 591, 599–603 (2021).Article 

Google Scholar 
Walker, A. P. et al. Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO2. N. Phytol. 229, 2413–2445 (2021).Article 

Google Scholar 
Fossum, C. et al. Belowground allocation and dynamics of recently fixed plant carbon in a California annual grassland. Soil Biol. Biochem. 165, 108519 (2022).Article 

Google Scholar 
Rasse, D. P., Rumpel, C. & Dignac, M.-F. Is soil carbon mostly root carbon? Mechanisms for a specific stabilisation. Plant Soil 269, 341–356 (2005).Article 

Google Scholar 
Sokol, N. W., Kuebbing, Sara, E., Karlsen-Ayala, E. & Bradford, M. A. Evidence for the primacy of living root inputs, not root or shoot litter, in forming soil organic carbon. N. Phytol. 221, 233–246 (2019).Article 

Google Scholar 
Calvo, O. C., Franzaring, J., Schmid, I. & Fangmeier, A. Root exudation of carbohydrates and cations from barley in response to drought and elevated CO2. Plant Soil 438, 127–142 (2019).Article 

Google Scholar 
Fransson, P. M. A. & Johansson, E. M. Elevated CO2 and nitrogen influence exudation of soluble organic compounds by ectomycorrhizal root systems. FEMS Microbiol. Ecol. 71, 186–196 (2009).Article 

Google Scholar 
Johansson, E. M., Fransson, P. M. A., Finlay, R. D. & van Hees, P. A. W. Quantitative analysis of soluble exudates produced by ectomycorrhizal roots as a response to ambient and elevated CO2. Soil Biol. Biochem. 41, 1111–1116 (2009).Article 

Google Scholar 
Phillips, R. P., Finzi, A. C. & Bernhardt, E. S. Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation. Ecol. Lett. 14, 187–194 (2011).Article 

Google Scholar 
Jilling, A., Keiluweit, M., Gutknecht, J. L. M. & Grandy, A. S. Priming mechanisms providing plants and microbes access to mineral-associated organic matter. Soil Biol. Biochem. 158, 108265 (2021).Article 

Google Scholar 
Cotrufo, M. F., Wallenstein, M. D., Boot, C. M., Denef, K. & Paul, E. The Microbial Efficiency-Matrix Stabilization (MEMS) framework integrates plant litter decomposition with soil organic matter stabilization: do labile plant inputs form stable soil organic matter? Glob. Change Biol. 19, 988–995 (2013).Article 

Google Scholar 
Sokol, N. W., Sanderman, J. & Bradford, M. A. Pathways of mineral-associated soil organic matter formation: integrating the role of plant carbon source, chemistry, and point of entry. Glob. Change Biol. 25, 12–24 (2019).Article 

Google Scholar 
Bradford, M. A., Keiser, A. D., Davies, C. A., Mersmann, C. A. & Strickland, M. S. Empirical evidence that soil carbon formation from plant inputs is positively related to microbial growth. Biogeochemistry 113, 271–281 (2013).Article 

Google Scholar 
Keiluweit, M. et al. Mineral protection of soil carbon counteracted by root exudates. Nat. Clim. Change 5, 588–595 (2015).Article 

Google Scholar 
Kuzyakov, Y., Friedel, J. K. & Stahr, K. Review of mechanisms and quantification of priming effects. Soil Biol. Biochem. 32, 1485–1498 (2000).Article 

Google Scholar 
Jones, D. L., Dennis, P. G., Owen, A. G. & van Hees, P. A. W. Organic acid behavior in soils—misconceptions and knowledge gaps. Plant Soil 248, 31–41 (2003).Article 

Google Scholar 
Cleveland, C. C. & Liptzin, D. C:N:P stoichiometry in soil: is there a “Redfield ratio” for the microbial biomass? Biogeochemistry 85, 235–252 (2007).Article 

Google Scholar 
Meier, I. C., Finzi, A. C. & Phillips, R. P. Root exudates increase N availability by stimulating microbial turnover of fast-cycling N pools. Soil Biol. Biochem. 106, 119–128 (2017).Article 

Google Scholar 
Canarini, A., Kaiser, C., Merchant, A., Richter, A. & Wanek, W. Root exudation of primary metabolites: mechanisms and their roles in plant responses to environmental stimuli. Front. Plant Sci. 10, 157 (2019).Article 

Google Scholar 
Koo, B.-J., Adriano, D. C., Bolan, N. S. & Barton, C. D. in Encyclopedia of Soils in the Environment (ed. Hillel, D.) 421–428 (Elsevier, 2005); https://doi.org/10.1016/B0-12-348530-4/00461-6Oldfield, E. E., Crowther, T. W. & Bradford, M. A. Substrate identity and amount overwhelm temperature effects on soil carbon formation. Soil Biol. Biochem. 124, 218–226 (2018).Article 

Google Scholar 
Mason-Jones, K., Schmücker, N. & Kuzyakov, Y. Contrasting effects of organic and mineral nitrogen challenge the N-mining hypothesis for soil organic matter priming. Soil Biol. Biochem. 124, 38–46 (2018).Article 

Google Scholar 
Sokol, N. W. & Bradford, M. A. Microbial formation of stable soil carbon is more efficient from belowground than aboveground input. Nat. Geosci. 12, 46–53 (2019).Article 

Google Scholar 
Drake, J. E. et al. Stoichiometry constrains microbial response to root exudation—insights from a model and a field experiment in a temperate forest. Biogeosciences 10, 821–838 (2013).Article 

Google Scholar 
Falchini, L., Naumova, N., Kuikman, P. J., Bloem, J. & Nannipieri, P. CO2 evolution and denaturing gradient gel electrophoresis profiles of bacterial communities in soil following addition of low molecular weight substrates to simulate root exudation. Soil Biol. Biochem. 35, 775–782 (2003).Article 

Google Scholar 
Rasmussen, C., Southard, R. J. & Horwath, W. R. Soil mineralogy affects conifer forest soil carbon source utilization and microbial priming. Soil Sci. Soc. Am. J. 71, 1141–1150 (2007).Article 

Google Scholar 
Frey, S. D., Lee, J., Melillo, J. M. & Six, J. The temperature response of soil microbial efficiency and its feedback to climate. Nat. Clim. Change 3, 395–398 (2013).Article 

Google Scholar 
Angst, G., Mueller, K. E., Nierop, K. G. J. & Simpson, M. J. Plant- or microbial-derived? A review on the molecular composition of stabilized soil organic matter. Soil Biol. Biochem. 156, 108189 (2021).Article 

Google Scholar 
Craig, M. E. et al. Fast-decaying plant litter enhances soil carbon in temperate forests but not through microbial physiological traits. Nat. Commun. 13, 1229 (2022).Article 

Google Scholar 
Blagodatsky, S., Blagodatskaya, E., Yuyukina, T. & Kuzyakov, Y. Model of apparent and real priming effects: linking microbial activity with soil organic matter decomposition. Soil Biol. Biochem. 42, 1275–1283 (2010).Article 

Google Scholar 
Hill, P. W., Farrar, J. F. & Jones, D. L. Decoupling of microbial glucose uptake and mineralization in soil. Soil Biol. Biochem. 40, 616–624 (2008).Article 

Google Scholar 
Asmar, F., Eiland, F. & Nielsen, N. E. Interrelationship between extracellular enzyme activity, ATP content, total counts of bacteria and CO2 evolution. Biol. Fertil. Soils 14, 288–292 (1992).Article 

Google Scholar 
Fontaine, S., Mariotti, A. & Abbadie, L. The priming effect of organic matter: a question of microbial competition? Soil Biol. Biochem. 35, 837–843 (2003).Article 

Google Scholar 
McFarlane, K. J. et al. Comparison of soil organic matter dynamics at five temperate deciduous forests with physical fractionation and radiocarbon measurements. Biogeochemistry 112, 457–476 (2013).Article 

Google Scholar 
Post, W. M., Emanuel, W. R., Zinke, P. J. & Stangenberger, A. G. Soil carbon pools and world life zones. Nature 298, 156–159 (1982).Article 

Google Scholar 
Smith, W. H. Character and significance of forest tree root exudates. Ecology 57, 324–331 (1976).Article 

Google Scholar 
Dong, J. et al. Impacts of elevated CO2 on plant resistance to nutrient deficiency and toxic ions via root exudates: a review. Sci. Total Environ. 754, 142434 (2021).Article 

Google Scholar 
White, M. A., Running, S. W. & Thornton, P. E. The impact of growing-season length variability on carbon assimilation and evapotranspiration over 88 years in the eastern US deciduous forest. Int. J. Biometeorol. 42, 139–145 (1999).Article 

Google Scholar 
Giasson, M.-A. et al. Soil respiration in a northeastern US temperate forest: a 22-year synthesis. Ecosphere 4, 140 (2013).Article 

Google Scholar 
Mrak, T. et al. Elevated ozone prevents acquisition of available nitrogen due to smaller root surface area in poplar. Plant Soil 450, 585–599 (2020).Article 

Google Scholar 
Cotrufo, M. F., Ranalli, M. G., Haddix, M. L., Six, J. & Lugato, E. Soil carbon storage informed by particulate and mineral-associated organic matter. Nat. Geosci. 12, 989–994 (2019).Article 

Google Scholar 
Brookes, P. C., Landman, A., Pruden, G. & Jenkinson, D. S. Chloroform fumigation and the release of soil nitrogen: a rapid direct extraction method to measure microbial biomass nitrogen in soil. Soil Biol. Biochem. 17, 837–842 (1985).Article 

Google Scholar 
Haney, R. L., Franzluebbers, A. J., Hons, F. M. & Zuberer, D. A. Soil C extracted with water or K2SO4: pH effect on determination of microbial biomass. Can. J. Soil Sci. 79, 529–533 (1999).Article 

Google Scholar 
Ahmed, M. J. & Hossan, J. Spectrophotometric determination of aluminium by morin. Talanta 42, 1135–1142 (1995).Article 

Google Scholar 
Viollier, E., Inglett, P. W., Hunter, K., Roychoudhury, A. N. & Van Cappellen, P. The ferrozine method revisited: Fe(II)/Fe(III) determination in natural waters. Appl. Geochem. 15, 785–790 (2000).Article 

Google Scholar  More

Curtis, P. G., Slay, C. M., Harris, N. L., Tyukavina, A. & Hansen, M. C. Classifying drivers of global forest loss. Science 361, 1108–1111 (2018).CAS 
PubMed 

Google Scholar 
Laurance, W. F., Sayer, J. & Cassman, K. G. Agricultural expansion and its impacts on tropical nature. Trends Ecol. Evol. 29, 107–116 (2014).PubMed 

Google Scholar 
Pendrill, F. et al. Agricultural and forestry trade drives large share of tropical deforestation emissions. Glob. Environ. Change 56, 1–10 (2019).
Google Scholar 
Haupt, F., Bakhtary, H., Schulte, I., Galt, H. & Streck, C. Progress on Corporate Commitments and their Implementation (Tropical Forest Alliance, 2018); https://www.tropicalforestalliance.org/assets/Uploads/Progress-on-Corporate-Commitments-and-their-Implementation.pdfAustin, K. G. et al. Mapping and monitoring zero-deforestation commitments. Bioscience 71, 1079–1090 (2021).PubMed 
PubMed Central 

Google Scholar 
Leijten, F. C., Sim, S., King, H. & Verburg, P. H. Which forests could be protected by corporate zero deforestation commitments? A spatial assessment. Environ. Res. Lett. 15, 064021 (2020).
Google Scholar 
Garrett, R. D. et al. Criteria for effective zero-deforestation commitments. Glob. Environ. Change https://doi.org/10.1016/j.gloenvcha.2018.11.003 (2019).Lehmann, C. E. R. & Parr, C. L. Tropical grassy biomes: linking ecology, human use and conservation. Phil. Trans. R. Soc. B https://doi.org/10.1098/rstb.2016.0329 (2016).Miles, L. et al. A global overview of the conservation status of tropical dry forests. J. Biogeogr. 33, 491–505 (2006).
Google Scholar 
Gibbs, H. K. et al. Brazil’s soy moratorium. Science https://doi.org/10.1126/science.aaa0181 (2015).Jopke, P. & Schoneveld, G. C. Corporate Commitments to Zero Deforestation: An Evaluation of Externality Problems and Implementation Gaps (CIFOR, 2018); https://doi.org/10.17528/cifor/006827Parr, C. L., Lehmann, C. E. R., Bond, W. J., Hoffmann, W. A. & Andersen, A. N. Tropical grassy biomes: misunderstood, neglected, and under threat. Trends Ecol. Evol. https://doi.org/10.1016/j.tree.2014.02.004 (2014).Ratnam, J. et al. When is a ‘forest’ a savanna, and why does it matter? Glob. Ecol. Biogeogr. https://doi.org/10.1111/j.1466-8238.2010.00634.x (2011).Sanchez-Azofeifa, G. A. et al. Research priorities for neotropical dry forests. Biotropica 37, 477–485 (2005).
Google Scholar 
Vijay, V., Pimm, S. L., Jenkins, C. N. & Smith, S. J. The impacts of oil palm on recent deforestation and biodiversity loss. PLoS ONE 11, e0159668 (2016).PubMed 
PubMed Central 

Google Scholar 
Principles & Criteria for the Production of Sustainable Palm Oil (RSPO, 2018).Rosoman, G. et al. (eds) The HCS Approach Toolkit (HCS Approach Steering Group, 2017).Brown, E. & Senior, M. J. M. (eds) Common Guidance for the Identification of High Conservation Values (HCV Resource Network, 2017).Furumo, P. R. & Aide, T. M. Characterizing commercial oil palm expansion in Latin America: land use change and trade. Environ. Res. Lett. 12, 024008 (2017).
Google Scholar 
Dinerstein, E. et al. An ecoregion-based approach to protecting half the terrestrial realm. BioScience https://doi.org/10.1093/biosci/bix014 (2017).Descals, A. et al. High-resolution global map of smallholder and industrial closed-canopy oil palm plantations. Earth Syst. Sci. Data 13, 1211–1231 (2021).
Google Scholar 
Woittiez, L. S., van Wijk, M. T., Slingerland, M., van Noordwijk, M. & Giller, K. E. Yield gaps in oil palm: a quantitative review of contributing factors. Eur. J. Agron. 83, 57–77 (2017).
Google Scholar 
Kuepper, B., Drost, S. & Piotrowski, M. Latin American Palm Oil Linked to Social Risks, Local Deforestation (Chain Reaction Research, 2021); https://chainreactionresearch.com/wp-content/uploads/2021/12/Latin-American-Palm-Oil-Linked-to-Social-Issues-Local-Deforestation-1.pdfHoyle, D. et al. RSPO New Planting Procedures: Summary Report of ESIA, HCV Assessments and Management Plan (Terea, Proforest and Olam Palm Gabon, 2017).Universal Mill List (World Resources Institute, Rainforest Alliance, Proforest & Daemeter, 2018); https://data.globalforestwatch.org/documents/gfw::universal-mill-list/aboutPirker, J., Mosnier, A., Kraxner, F., Havlík, P. & Obersteiner, M. What are the limits to oil palm expansion? Glob. Environ. Change 40, 73–81 (2016).
Google Scholar 
Fischer, G. et al. Global Agro-Ecological Zones 4 (GAEZ v4) – Model Documentation (FAO, 2021); https://doi.org/10.4060/cb4744enGlobal Agro-Ecological Zoning Version 4 (GAEZ v4) (FAO & IIASA, 2021); http://www.fao.org/gaez/Tao, H. H. et al. Long-term crop residue application maintains oil palm yield and temporal stability of production. Agron. Sustain. Dev. https://doi.org/10.1007/s13593-017-0439-5 (2017).Wei, L., John Martin, J. J., Zhang, H., Zhang, R. & Cao, H. Problems and prospects of improving abiotic stress tolerance and pathogen resistance of oil palm. Plants 10, 2622 (2021).CAS 
PubMed 
PubMed Central 

Google Scholar 
Corley, R. H. & Tinker, P. B. The Oil Palm (Wiley-Blackwell, 2016).Barona, E., Ramankutty, N., Hyman, G. & Coomes, O. T. The role of pasture and soybean in deforestation of the Brazilian Amazon. Environ. Res. Lett. 5, 024002 (2010).
Google Scholar 
ten Kate, A., Kuepper, B. & Piotrowski, M. NDPE Policies Cover 83% of Palm Oil Refineries; Implementation at 78% (Chain Reaction Research, 2020); https://chainreactionresearch.com/wp-content/uploads/2020/04/NDPE-Policies-Cover-83-of-Palm-Oil-Refining-Market.pdfThe Trase Yearbook: The State Of Forest Risk Supply Chains (Trase, 2020); https://insights.trase.earth/yearbook/summaryAustin, K. G. et al. Shifting patterns of oil palm driven deforestation in Indonesia and implications for zero-deforestation commitments. Land Use Policy 69, 41–48 (2017).
Google Scholar 
Furumo, P. R., Rueda, X., Rodríguez, J. S. & Parés Ramos, I. K. Field evidence for positive certification outcomes on oil palm smallholder management practices in Colombia. J. Clean. Prod. 245, 118891 (2020).
Google Scholar 
Carlson, K. M. et al. Effect of oil palm sustainability certification on deforestation and fire in Indonesia. Proc. Natl Acad. Sci. USA 115, 121–126 (2018).CAS 
PubMed 

Google Scholar 
Heilmayr, R., Carlson, K. M. & Benedict, J. J. Deforestation spillovers from oil palm sustainability certification. Environ. Res. Lett. 15, 075002 (2020).CAS 

Google Scholar 
Impact (RSPO, 2022); https://www.rspo.org/impactBastos Lima, M. G., Persson, U. M. & Meyfroidt, P. Leakage and boosting effects in environmental governance: a framework for analysis. Environ. Res. Lett. 14, 105006 (2019).
Google Scholar 
Corley, R. H. V. How much palm oil do we need? Environ. Sci. Policy https://doi.org/10.1016/j.envsci.2008.10.011 (2009).FAOSTAT: Food and Agriculture Data (FAO, 2020); https://www.fao.org/faostat/en/Olson, D. M. et al. Terrestrial ecoregions of the world: a new map of life on Earth: a new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. BioScience 51, 933–938 (2001).
Google Scholar 
Murphy, B. P., Andersen, A. N. & Parr, C. L. The underestimated biodiversity of tropical grassy biomes. Phil. Trans. R. Soc. B 371, 20150319 (2016).PubMed 
PubMed Central 

Google Scholar 
Smith, J. R., Hendershot, J. N., Nova, N. & Daily, G. C. The biogeography of ecoregions: descriptive power across regions and taxa. J. Biogeogr. https://doi.org/10.1111/jbi.13871 (2020).Klink, C. A. & Machado, R. B. Conservation of the Brazilian Cerrado. Conserv. Biol. 19, 707–713 (2005).
Google Scholar 
Strassburg, B. B. N. et al. Moment of truth for the Cerrado hotspot. Nat. Ecol. Evol. https://doi.org/10.1038/s41559-017-0099 (2017).le Polain de Waroux, Y. et al. The restructuring of South American soy and beef production and trade under changing environmental regulations. World Dev. 121, 188–202 (2019).
Google Scholar 
Nepstad, L. S. et al. Pathways for recent Cerrado soybean expansion: extending the soy moratorium and implementing integrated crop livestock systems with soybeans. Environ. Res. Lett. 14, 044029 (2019).
Google Scholar 
Searchinger, T. D. et al. High carbon and biodiversity costs from converting Africa’s wet savannahs to cropland. Nat. Clim. Change https://doi.org/10.1038/nclimate2584 (2015).Cardoso Da Silva, J. M. & Bates, J. M. Biogeographic patterns and conservation in the South American Cerrado: a tropical savanna hotspot. BioScience 52, 225–233 (2002).
Google Scholar 
Poggio, L. et al. SoilGrids 2.0: producing soil information for the globe with quantified spatial uncertainty. SOIL 7, 217–240 (2021).Hill, T. C., Williams, M., Bloom, A. A., Mitchard, E. T. A. & Ryan, C. M. Are inventory based and remotely sensed above-ground biomass estimates consistent? PLoS ONE https://doi.org/10.1371/journal.pone.0074170 (2013).Ryan, C. M. et al. Ecosystem services from southern African woodlands and their future under global change. Phil. Trans. R. Soc. B https://doi.org/10.1098/rstb.2015.0312 (2016).Grace, J., Jose, J. S., Meir, P., Miranda, H. S. & Montes, R. A. Productivity and carbon fluxes of tropical savannas. J. Biogeogr. 33, 387–400 (2006).
Google Scholar 
Scharlemann, J. P., Tanner, E. V., Hiederer, R. & Kapos, V. Global soil carbon: understanding and managing the largest terrestrial carbon pool. Carbon Manag. 5, 81–91 (2014).CAS 

Google Scholar 
Quezada, J. C., Etter, A., Ghazoul, J., Buttler, A. & Guillaume, T. Carbon neutral expansion of oil palm plantations in the Neotropics. Sci. Adv. 5, eaaw4418 (2019).CAS 
PubMed 
PubMed Central 

Google Scholar 
Aleman, J. C., Blarquez, O. & Staver, C. A. Land-use change outweighs projected effects of changing rainfall on tree cover in sub-Saharan Africa. Glob. Change Biol. https://doi.org/10.1111/gcb.13299 (2016).Espírito-Santo, M. M. et al. Understanding patterns of land-cover change in the Brazilian Cerrado from 2000 to 2015. Phil. Trans. R. Soc. B https://doi.org/10.1098/rstb.2015.0435 (2016).Overbeck, G. E. et al. Conservation in Brazil needs to include non-forest ecosystems. Divers. Distrib. https://doi.org/10.1111/ddi.12380 (2015).Hoekstra, J. M., Boucher, T. M., Ricketts, T. H. & Roberts, C. Confronting a biome crisis: global disparities of habitat loss and protection. Ecol. Lett. https://doi.org/10.1111/j.1461-0248.2004.00686.x (2005).RTRS Standard for Responsible Soy Production Version 3.1 (RTRS, 2017); https://responsiblesoy.org/wp-content/uploads/2019/08/RTRS%20Standard%20Responsible%20Soy%20production%20V3.1%20ING-LOW.pdfBatlle-Bayer, L., Batjes, N. H. & Bindraban, P. S. Changes in organic carbon stocks upon land use conversion in the Brazilian Cerrado: a review. Agric. Ecosyst. Environ. 137, 47–58 (2010).CAS 

Google Scholar 
Rockström, J., Falkenmark, M., Lannerstad, M. & Karlberg, L. The planetary water drama: dual task of feeding humanity and curbing climate change. Geophys. Res. Lett. 39, LXXXXX (2012).
Google Scholar 
Ocampo-Peñuela, N., Garcia-Ulloa, J., Ghazoul, J. & Etter, A. Quantifying impacts of oil palm expansion on Colombia’s threatened biodiversity. Biol. Conserv. https://doi.org/10.1016/j.biocon.2018.05.024 (2018).Gilroy, J. J. et al. Minimizing the biodiversity impact of Neotropical oil palm development. Glob. Change Biol. 21, 1531–1540 (2015).
Google Scholar 
Bonn Challenge 2020 Report (IUCN, 2020); https://www.bonnchallenge.org/resources/bonn-challenge-2020-reportGilroy, J. J. et al. Cheap carbon and biodiversity co-benefits from forest regeneration in a hotspot of endemism. Nat. Clim. Change 4, 503–507 (2014).
Google Scholar 
Evans, M. C. et al. Carbon farming via assisted natural regeneration as a cost-effective mechanism for restoring biodiversity in agricultural landscapes. Environ. Sci. Policy 50, 114–129 (2015).CAS 

Google Scholar 
Hunter, M. C., Smith, R. G., Schipanski, M. E., Atwood, L. W. & Mortensen, D. A. Agriculture in 2050: recalibrating targets for sustainable intensification. BioScience https://doi.org/10.1093/biosci/bix010 (2017).Beyer, R. & Rademacher, T. Species richness and carbon footprints of vegetable oils: can high yields outweigh palm oil’s environmental impact? Sustainability 13, 1813 (2021).
Google Scholar 
Lee, J. S. H., Ghazoul, J., Obidzinski, K. & Koh, L. P. Oil palm smallholder yields and incomes constrained by harvesting practices and type of smallholder management in Indonesia. Agron. Sustain. Dev. 34, 501–513 (2014).
Google Scholar 
Murphy, D. J. The future of oil palm as a major global crop: opportunities and challenges. J. Oil Palm Res. 26, 1–24 (2014).
Google Scholar 
Giam, X., Koh, L. P. & Wilcove, D. S. Tropical crops: cautious optimism. Science https://doi.org/10.1126/science.346.6212.928-a (2014).Villoria, N. B., Golub, A., Byerlee, D. & Stevenson, J. Will yield improvements on the forest frontier reduce greenhouse gas emissions? A global analysis of oil palm. Am. J. Agric. Econ. 95, 1301–1308 (2013).
Google Scholar 
Koh, L. P. & Lee, T. M. Sensible consumerism for environmental sustainability. Biol. Conserv. https://doi.org/10.1016/j.biocon.2011.10.029 (2012).Fick, S. E. & Hijmans, R. J. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37, 4302–4315 (2017).
Google Scholar 
Harris, N., Goldman, E. & Gibbes, S. Spatial Database of Planted Trees (SDPT) Version 1.0 (World Resources Institute, 2019); https://data.globalforestwatch.org/datasets/tree-plantationsSutanudjaja, E. H. et al. PCR-GLOBWB 2: a 5 arcmin global hydrological and water resources model. Geosci. Model Dev. 11, 2429–2453 (2018).
Google Scholar 
Global Land Cover (Copernicus, 2019); https://lcviewer.vito.be/Tsendbazar, N.-E. et al. Copernicus Global Land Operations ‘Vegetation and Energy’ ‘CGLOPS−1’ Validation Report. Moderate Dynamic Land Cover 100m Version 2 (WUR, 2019); https://land.copernicus.eu/global/sites/cgls.vito.be/files/products/CGLOPS1_VR_LC100m-V2.0_I1.00.pdfSantoro, M. et al. GlobBiomass – global datasets of forest biomass. PANGAEA https://doi.org/10.1594/PANGAEA.894711 (2018).Santoro, M. et al. A detailed portrait of the forest aboveground biomass pool for the year 2010 obtained from multiple remote sensing observations. Geophys. Res. Abstr. https://doi.org/10.1002/joc.5086 (2018).Hansen, M. C. et al. High-resolution global maps of 21st-century forest cover change. Science https://doi.org/10.1126/science.1244693 (2013).Gumbricht, T. et al. Tropical and Subtropical Wetlands Distribution Version 7 (CIFOR, 2017); https://doi.org/10.17528/CIFOR/DATA.00058The IUCN Red List of Threatened Species Version 2018−1 (IUCN, 2018); https://www.iucnredlist.orgBird Species Distribution Maps of the World Version 6.0 (BirdLife International & Handbook of the Birds of the World, 2016); http://datazone.birdlife.org/species/requestdisR Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2018).Silalertruksa, T. et al. Environmental sustainability of oil palm cultivation in different regions of Thailand: greenhouse gases and water use impact. J. Clean. Prod. 167, 1009–1019 (2017).CAS 

Google Scholar 
Fourcade, Y., Engler, J. O., Rödder, D. & Secondi, J. Mapping species distributions with Maxent using a geographically biased sample of presence data: a performance assessment of methods for correcting sampling bias. PLoS ONE 9, e97122 (2014).PubMed 
PubMed Central 

Google Scholar 
Liu, Z. et al. Shifts in the extent and location of rice cropping areas match the climate change pattern in China during 1980–2010. Reg. Environ. Change https://doi.org/10.1007/s10113-014-0677-x (2015).Singh, K., McClean, C. J., Büker, P., Hartley, S. E. & Hill, J. K. Mapping regional risks from climate change for rainfed rice cultivation in India. Agric. Syst. 156, 76–84 (2017).PubMed 
PubMed Central 

Google Scholar 
Estes, L. D. et al. Comparing mechanistic and empirical model projections of crop suitability and productivity: implications for ecological forecasting. Glob. Ecol. Biogeogr. https://doi.org/10.1111/geb.12034 (2013).Thuiller, W., Georges, D., Engler, R. & Breiner, F. biomod2: Ensemble Platform for Species Distribution Modelling (2016).Hernandez, P. A., Graham, C. H., Master, L. L. & Albert, D. L. The effect of sample size and species characteristics on performance of different species distribution modeling methods. Ecography 29, 773–785 (2006).
Google Scholar 
Merow, C., Smith, M. J., Silander, J. A., Merow, C. & Silander, J. A. A practical guide to Maxent for modeling species’ distributions: what it does, and why inputs and settings matter. Ecography 36, 1058–1069 (2013).
Google Scholar 
Phillips, S. J., Anderson, R. P. & Schapire, R. E. Maximum entropy modeling of species geographic distributions. Ecol. Modell. https://doi.org/10.1016/j.ecolmodel.2005.03.026 (2006).VanDerWal, J., Shoo, L. P., Graham, C. & Williams, S. E. Selecting pseudo-absence data for presence-only distribution modeling: how far should you stray from what you know? Ecol. Modell. 220, 589–594 (2009).
Google Scholar 
Hirzel, A. H., Le Lay, G., Helfer, V., Randin, C. F. & Guisan, A. Evaluating the ability of habitat suitability models to predict species presences. Ecol. Modell. 199, 142–152 (2006).
Google Scholar 
Engler, R., Guisan, A. & Rechsteiner, L. An improved approach for predicting the distribution of rare and endangered species from occurrence and pseudo-absence data. J. Appl. Ecol. https://doi.org/10.1111/j.0021-8901.2004.00881.x (2004).Allouche, O., Tsoar, A. & Kadmon, R. Assessing the accuracy of species distribution models: prevalence, kappa and the true skill statistic (TSS). J. Appl. Ecol. https://doi.org/10.1111/j.1365-2664.2006.01214.x (2006).Global Spatially-Disaggregated Crop Production Statistics Data for 2010 Version 1.1. (International Food Policy Research Institute, 2019); https://doi.org/10.7910/DVN/PRFF8V/M2EMBNHofste, R. W. et al. Aqueduct 3.0: Updated Decision-Relevant Global Water Risk Indicators (World Resources Institute, 2019); https://www.wri.org/research/aqueduct-30-updated-decision-relevant-global-water-risk-indicatorsCarr, M. K. V. The water relations and irrigation requirements of oil palm (Elaeis guineensis): a review. Exp. Agric. 47, 629–652 (2011).
Google Scholar 
Yusop, Z., Hui, C. M., Garusu, G. J. & Katimon, A. Estimation of evapotranspiration in oil palm catchments by short-time period water-budget method. Malays. J. Civ. Eng. 20, 160–174 (2008).
Google Scholar 
Hargreaves, G. H. & Allen, R. G. History and evaluation of Hargreaves evapotranspiration equation. J. Irrig. Drain. Eng. 129, 53–63 (2003).
Google Scholar 
Trabucco, A. & Zomer, R. J. Global aridity index and potential evapotranspiration (ET0) climate database v2. Figshare https://doi.org/10.6084/m9.figshare.7504448.v3 (2019).Protected Planet: The World Database on Protected Areas (WDPA) (UNEP-WCMC & IUCN, 2020); www.protectedplanet.net/en/thematic-areas/wdpaDudley, N. (ed.) Guidelines for Applying Protected Area Management Categories (IUCN, 2008).Juffe-Bignoli, D. et al. World Database on Protected Areas User Manual 1.5 (UNEP-WCMC, 2017); https://www.protectedplanet.net/en/resources/wdpa-manualChave, J. J. et al. Tree allometry and improved estimation of carbon stocks and balance in tropical forests. Oecologia 145, 87–99 (2005).CAS 
PubMed 

Google Scholar 
Jetz, W., Wilcove, D. S. & Dobson, A. P. Projected impacts of climate and land-use change on the global diversity of birds. PLoS Biol. https://doi.org/10.1371/journal.pbio.0050157 (2007).Beyer, R. M. & Manica, A. Historical and projected future range sizes of the world’s mammals, birds, and amphibians. Nat. Commun. 11, 5633 (2020).CAS 
PubMed 
PubMed Central 

Google Scholar 
Beyer, R. M. & Manica, A. Global and country-level data of the biodiversity footprints of 175 crops and pasture. Data Brief 36, 106982 (2021).CAS 
PubMed 
PubMed Central 

Google Scholar 
Cobertura de la Tierra 100K Periodo 2018 (IDEAM, Instituto de Hidrología, Meteorología y Estudios Ambientales, 2021); http://www.siac.gov.co/catalogo-de-mapasSouza, C. M. et al. Reconstructing three decades of land use and land cover changes in Brazilian biomes with Landsat archive and Earth Engine. Remote Sens. 12, 2735 (2020).
Google Scholar 
MapBiomas Project – Collection 6 of the Annual Series of Land Use and Land Cover Maps of Brazil (MapBiomas, 2021); https://plataforma.brasil.mapbiomas.org/Veldman, J. W. & Putz, F. E. Grass-dominated vegetation, not species-diverse natural savanna, replaces degraded tropical forests on the southern edge of the Amazon Basin. Biol. Conserv. https://doi.org/10.1016/j.biocon.2011.01.011 (2011).Portillo-Quintero, C. A. & Sánchez-Azofeifa, G. A. Extent and conservation of tropical dry forests in the Americas. Biol. Conserv. https://doi.org/10.1016/j.biocon.2009.09.020 (2010).Veldman, J. W. et al. Toward an old-growth concept for grasslands, savannas, and woodlands. Front. Ecol. Environ. https://doi.org/10.1890/140270 (2015).Zaloumis, N. P. & Bond, W. J. Reforestation or conservation? The attributes of old growth grasslands in South Africa. Phil. Trans. R. Soc. B https://doi.org/10.1098/rstb.2015.0310 (2016).Garnett, S. T. et al. A spatial overview of the global importance of Indigenous lands for conservation. Nat. Sustain. https://doi.org/10.1038/s41893-018-0100-6 (2018).Djoudi, H., Vergles, E., Blackie, R. R., Koame, C. K. & Gautier, D. Dry forests, livelihoods and poverty alleviation: understanding current trends. Int. For. Rev. https://doi.org/10.1505/146554815815834868 (2015).Ground-Truthing to Improve Due Diligence on Human Rights in Deforestation-Risk Supply Chains (Forest Peoples Programme, 2020); https://www.forestpeoples.org/en/ground-truthing-to-improve-due-diligenceDrost, S., Rijk, G. & Piotrowski, M. Oil Palm Growers Exposed to USD 0.4-5.9B in Social Compensation Risk (Chain Reaction Research, 2019); https://chainreactionresearch.com/wp-content/uploads/2019/12/Social-compensation-risks-for-palm-growers-4.pdf More

Extent and location of critical natural assetsCritical natural assets providing the 12 local NCP (Fig. 1a) occupy only 30% (41 million km2) of total land area (excluding Antarctica) and 24% (34 million km2) of marine Exclusive Economic Zones (EEZs), reflecting the steep slope of the aggregate NCP accumulation curve (Fig. 1b). Despite this modest proportion of global land area, the shares of countries’ land areas that are designated as critical can vary substantially. The 20 largest countries require only 24% of their land area, on average, to maintain 90% of current levels of NCP, while smaller countries (10,000 to 1.5 million km2) require on average 40% of their land area (Supplementary Data 1). This high variability in the NCP–area relationship is primarily driven by the proportion of countries’ land areas made up by natural assets (that is, excluding barren, ice and snow, and developed lands), but even when this is accounted for, there are outliers (Extended Data Fig. 2). Outliers may be due to spatial patterns in human population density (for example, countries with dense population centres and vast expanses with few people, such as Canada and Russia, require far less area to achieve NCP targets) or large ecosystem heterogeneity (if greater ecosystem diversity yields higher levels of diverse NCP in a smaller proportion of area, which may explain patterns in Chile and Australia).The highest-value critical natural assets (the locations delivering the highest magnitudes of NCP in the smallest area, denoted by the darkest blue or green shades in Fig. 1c) often coincide with diverse, relatively intact natural areas near or upstream from large numbers of people. Many of these high-value areas coincide with areas of greatest spatial congruence among multiple NCP (Extended Data Fig. 3). Spatially correlated pairs of local NCP (Supplementary Table 4) include those related to water (flood risk reduction with nitrogen retention and nitrogen with sediment retention); forest products (timber and fuelwood); and those occurring closer to human-modified habitats (pollination with nature access and with nitrogen retention). Coastal risk reduction, forage production for grazing, and riverine fish harvest are the most spatially distinct from other local NCP. In the marine realm, there is substantial overlap of fisheries with coastal risk reduction and reef tourism (though not between the latter two, which each have much smaller critical areas than exist for fisheries).Number of people benefitting from critical natural assetsWe estimate that ~87% of the world’s current population, 6.4 billion people, benefit directly from at least one of the 12 local NCP provided by critical natural assets, while only 16% live on the lands providing these benefits (and they may also benefit; Fig. 2a). To quantify the number of beneficiaries of critical natural assets, we spatially delineate their benefitting areas (which varies on the basis of NCP: for example, areas downstream, within the floodplain, in low-lying areas near the coast, or accessible by a short travel). While our optimization selects for the provision of 90% of the current value of each NCP, it is not guaranteed that 90% of the world’s population would benefit (since it does not include considerations for redundancy in adjacent pixels and therefore many of the areas selected benefit the same populations), so it is notable that an estimated 87% do. This estimate of ‘local’ beneficiaries probably underestimates the total number of people benefitting because it includes only NCP for which beneficiaries can be spatially delineated to avoid double-counting, yet it is striking that the vast majority, 6.1 billion people, live within 1 h travel (by road, rail, boat or foot, taking the fastest path17) of critical natural assets, and more than half of the world’s population lives downstream of these areas (Fig. 2b). Material NCP are often delivered locally, but many also enter global supply chains, making it difficult to delineate beneficiaries spatially for these NCP. However, past studies have calculated that globally more than 54 million people benefit directly from the timber industry18, 157 million from riverine fisheries19, 565 million from marine fisheries20 and 1.3 billion from livestock grazing21, and across the tropics alone 2.7 billion are estimated to be dependent on nature for one or more basic needs22.Fig. 2: People benefitting from and living on critical natural assets (CNA).a,b, ‘Local’ beneficiaries were calculated through the intersection of areas benefitting from different NCP, to avoid double-counting people in areas of overlap; only those NCP for which beneficiaries could be spatially delineated were included (that is, not material NCP that enter global supply chains: fisheries, timber, livestock or crop pollination). Bars show percentages of total population globally and for large and small countries (a) or the percentage of relevant population globally (b). Numbers inset in bars show millions of people making up that percentage. Numbers to the right of bars in b show total relevant population (in millions of people, equivalent to total global population from Landscan 2017 for population within 1 h travel or downstream, but limited to the total population living within 10 km of floodplains or along coastlines 80%) of their populations benefitting from critical natural assets, but small countries have much larger proportions of their populations living within the footprint of critical natural assets than do large countries (Fig. 2a and Supplementary Data 2). When people live in these areas, and especially when current levels of use of natural assets are not sustainable, regulations or incentives may be needed to maintain the benefits these assets provide. While protected areas are an important conservation strategy, they represent only 15% of the critical natural assets for local NCP (Supplementary Table 5); additional areas should not necessarily be protected using designations that restrict human access and use, or they could cease to provide some of the diverse values that make them so critical23. Other area-based conservation measures, such as those based on Indigenous and local communities’ governance systems, Payments for Ecosystem Services programmes, and sustainable use of land- and seascapes, can all contribute to maintaining critical flows of NCP in natural and semi-natural ecosystems24.Overlaps between local and global prioritiesUnlike the 12 local NCP prioritized here at the national scale, certain benefits of natural assets accrue continentally or even globally. We therefore optimize two additional NCP at a global scale: vulnerable terrestrial ecosystem carbon storage (that is, the amount of total ecosystem carbon lost in a typical disturbance event25, hereafter ‘ecosystem carbon’) and vegetation-regulated atmospheric moisture recycling (the supply of atmospheric moisture and precipitation sustained by plant life26, hereafter ‘moisture recycling’). Over 80% of the natural asset locations identified as critical for the 12 local NCP are also critical for the two global NCP (Fig. 3). The spatial overlap between critical natural assets for local and global NCP accounts for 24% of land area, with an additional 14% of land area critical for global NCP that is not considered critical for local NCP (Extended Data Fig. 4). Together, critical natural assets for securing both local and global NCP require 44% of total global land area. When each NCP is optimized individually (carbon and moisture NCP at the global scale; the other 12 at the country scale), the overlap between carbon or moisture NCP and the other NCP exceeds 50% for all terrestrial (and freshwater) NCP except coastal risk reduction (which overlaps only 36% with ecosystem carbon, 5% with moisture recycling; Supplementary Table 4).Fig. 3: Spatial overlaps between critical natural assets for local and global NCP.Red and teal denote where critical natural assets for global NCP (providing 90% of ecosystem carbon and moisture recycling globally) or for local NCP (providing 90% of the 12 NCP listed in Fig. 1), respectively, but not both, occur; gold shows areas where the two overlap (24% of the total area). Together, local and global critical natural assets account for 44% of total global land area (excluding Antarctica). Grey areas show natural assets not defined as ‘critical’ by this analysis, though still providing some values to certain populations. White areas were excluded from the optimization.Full size imageSynergies can also be found between NCP and biodiversity and cultural diversity. Critical natural assets for local NCP at national levels overlap with part or all of the area of habitat (AOH, mapped on the basis of species range maps, habitat preferences and elevation27) for 60% of 28,177 terrestrial vertebrates (Supplementary Data 3). Birds (73%) and mammals (66%) are better represented than reptiles and amphibians (44%). However, these critical natural assets represent only 34% of the area for endemic vertebrate species (with 100% of their AOH located within a given country; Supplementary Data 3) and 16% of the area for all vertebrates if using a more conservative representation target framework based on the IUCN Red List criteria (though, notably, achieving Red List representation targets is impossible for 24% of species without restoration or other expansion of existing AOH; Supplementary Data 4). Cultural diversity (proxied by linguistic diversity) has far higher overlaps with critical natural assets than does biodiversity; these areas intersect 96% of global Indigenous and non-migrant languages28 (Supplementary Data 5). The degree to which languages are represented in association with critical natural assets is consistent across most countries, even at the high end of language diversity (countries containing >100 Indigenous and non-migrant languages, such as Indonesia, Nigeria and India). This high correspondence provides further support for the importance of safeguarding rights to access critical natural assets, especially for Indigenous cultures that benefit from and help maintain them. Despite the larger land area required for maintaining the global NCP compared with local NCP, global NCP priority areas overlap with slightly fewer languages (92%) and with only 2% more species (60% of species AOH), although a substantially greater overlap is seen with global NCP if Red List criteria are considered (36% compared with 16% for local NCP; Supplementary Data 4). These results provide different insights than previous efforts at smaller scales, particularly a similar exercise in Europe that found less overlap with priority areas for biodiversity and NCP29. However, the 40% of all vertebrate species whose habitats did not overlap with critical natural assets could drive very different patterns if biodiversity were included in the optimization.Although these 14 NCP are not comprehensive of the myriad ways that nature benefits and is valued by people23, they capture, spatially and thematically, many elements explicitly mentioned in the First Draft of the CBD’s post-2020 Global Biodiversity Framework13: food security, water security, protection from hazards and extreme events, livelihoods and access to green and blue spaces. Our emphasis here is to highlight the contributions of natural and semi-natural ecosystems to human wellbeing, specifically contributions that are often overlooked in mainstream conservation and development policies around the world. For example, considerations for global food security often include only crop production rather than nature’s contributions to it via pollination or vegetation-mediated precipitation, or livestock production without partitioning out the contribution of grasslands from more intensified feed production.Gaps and next stepsOur synthesis of these 14 NCP represents a substantial advance beyond other global prioritizations that include NCP limited to ecosystem carbon stocks, fresh water and marine fisheries30,31,32, though still falls short of including all important contributions of nature such as its relational values33. Despite the omission of many NCP that were not able to be mapped, further analyses indicate that results are fairly robust to inclusion of additional NCP. Dropping one of the 12 local NCP at a time results in More

Aristegui, J., Gasol, J. M., Duarte, C. M. & Herndl, G. J. Microbial oceanography of the dark ocean’s pelagic realm. Limnol. Oceanogr. 54, 1501–1529 (2009).Article 

Google Scholar 
Jannasch, H. W., Eimhjellen, K., Wirsen, C. O. & Farmanfarmaian, A. Microbial degradation of organic matter in the deep sea. Science 171, 672–675 (1971).Article 

Google Scholar 
Tamburini, C., Boutrif, M., Garel, M., Colwell, R. R. & Deming, J. W. Prokaryotic responses to hydrostatic pressure in the ocean – a review. Environ. Microbiol. 15, 1262–1274 (2013).Article 

Google Scholar 
Yayanos, A. A. Microbiology to 10,500 meters in the deep-sea. Annu. Rev. Microb. 49, 777–805 (1995).Article 

Google Scholar 
Jebbar, M., Franzetti, B., Girard, E. & Oger, P. Microbial diversity and adaptation to high hydrostatic pressure in deep-sea hydrothermal vents prokaryotes. Extremophiles 19, 721–740 (2015).Article 

Google Scholar 
Yayanos, A. A. Evolutional and ecological implications of the properties of deep-sea barophilic bacteria. Proc. Natl Acad. Sci. USA 83, 9542–9546 (1986).Article 

Google Scholar 
Nagata, T. et al. Emerging concepts on microbial processes in the bathypelagic ocean – ecology, biogeochemistry, and genomics. Deep-Sea Res. II 57, 1519–1536 (2010).Article 

Google Scholar 
Picard, A. & Daniel, I. Pressure as an environmental parameter for microbial life – a review. Biophys. Chem. 183, 30–41 (2013).Article 

Google Scholar 
Herndl, G. J. & Reinthaler, T. Microbial control of the dark end of the biological pump. Nat. Geosci. 6, 718–724 (2013).Article 

Google Scholar 
Marietou, A. & Bartlett, D. H. Effects of high hydrostatic pressure on coastal bacterial community abundance and diversity. Appl. Environ. Microbiol. 80, 5992–6003 (2014).Article 

Google Scholar 
Lauro, F. M. & Bartlett, D. H. Prokaryotic lifestyles in deep sea habitats. Extremophiles 12, 15–25 (2008).Article 

Google Scholar 
Peoples, L. M. et al. Distinctive gene and protein characteristics of extremely piezophilic Colwellia. BMC Genom. 21, 692 (2020).Article 

Google Scholar 
Reinthaler, T. et al. Prokaryotic respiration and production in the meso- and bathypelagic realm of the eastern and western North Atlantic basin. Limnol. Oceanogr. 51, 1262–1273 (2006).Article 

Google Scholar 
Steinberg, D. K. et al. Bacterial vs. zooplankton control of sinking particle flux in the ocean’s twilight zone. Limnol. Oceanogr. 53, 1327–1338 (2008).Article 

Google Scholar 
Burd, A. B. et al. Assessing the apparent imbalance between geochemical and biochemical indicators of meso- and bathypelagic biological activity: what the @$#! is wrong with present calculations of carbon budgets? Deep-Sea Res. II 57, 1557–1571 (2010).Article 

Google Scholar 
Boyd, P. W., Claustre, H., Levy, M., Siegel, D. A. & Weber, T. Multi-faceted particle pumps drive carbon sequestration in the ocean. Nature 568, 327–335 (2019).Article 

Google Scholar 
Kirchman, D., Knees, E. & Hodson, R. Leucine incorporation and its potential as a measure of protein-synthesis by bacteria in natural aquatic systems. Appl. Environ. Microbiol. 49, 599–607 (1985).Article 

Google Scholar 
Nielsen, J. L., Christensen, D., Kloppenborg, M. & Nielsen, P. H. Quantification of cell-specific substrate uptake by probe-defined bacteria under in situ conditions by microautoradiography and fluorescence in situ hybridization. Environ. Microbiol. 5, 202–211 (2003).Article 

Google Scholar 
Sintes, E. & Herndl, G. J. Quantifying substrate uptake by individual cells of marine bacterioplankton by catalyzed reporter deposition fluorescence in situ hybridization combined with micro autoradiography. Appl. Environ. Microbiol. 72, 7022–7028 (2006).Article 

Google Scholar 
Garel, M. et al. Pressure-retaining sampler and high-pressure systems to study deep-sea microbes under in situ conditions. Front. Microbiol 10, 453 (2019).Article 

Google Scholar 
Peoples, L. M. et al. A full-ocean-depth rated modular lander and pressure-retaining sampler capable of collecting hadal-endemic microbes under in situ conditions. Deep-Sea Res. I 143, 50–57 (2019).Article 

Google Scholar 
Gross, M. & Jaenicke, R. Proteins under pressure – the influence of high hydrostatic pressure on structure, function and assembly of proteins and protein complexes. Eur. J. Biochem. 221, 617–630 (1994).Article 

Google Scholar 
Kirchman, D. L. Growth rates of microbes in the oceans. Annu. Rev. Mar. Sci. 8, 285–309 (2016).Article 

Google Scholar 
Ashburner, M. et al. Gene ontology: tool for the unification of biology. Nat. Genet. 25, 25–29 (2000).Article 

Google Scholar 
Xie, Z., Jian, H., Jin, Z. & Xiao, X. Enhancing the adaptability of the deep-sea bacterium Shewanella piezotolerans WP3 to high pressure and low temperature by experimental evolution under H2O2 stress. Appl. Environ. Microbiol. 84, e02342–02317 (2018).Article 

Google Scholar 
Tamburini, C. et al. Effects of hydrostatic pressure on microbial alteration of sinking fecal pellets. Deep-Sea Res. II 56, 1533–1546 (2009).Article 

Google Scholar 
Ivars-Martinez, E. et al. Comparative genomics of two ecotypes of the marine planktonic copiotroph Alteromonas macleodii suggests alternative lifestyles associated with different kinds of particulate organic matter. ISME J. 2, 1194–1212 (2008).Article 

Google Scholar 
Zhao, Z., Baltar, F. & Herndl, G. J. Linking extracellular enzymes to phylogeny indicates a predominantly particle-associated lifestyle of deep-sea prokaryotes. Sci. Adv. 6, eaaz4354 (2020).Article 

Google Scholar 
Bochdansky, A. B., van Aken, H. M. & Herndl, G. J. Role of macroscopic particles in deep-sea oxygen consumption. Proc. Natl Acad. Sci. USA 107, 8287–8291 (2010).Article 

Google Scholar 
Chikuma, S., Kasahara, R., Kato, C. & Tamegai, H. Bacterial adaptation to high pressure: a respiratory system in the deep-sea bacterium Shewanella violacea DSS12. FEMS Microbiol. Lett. 267, 108–112 (2007).Article 

Google Scholar 
Qin, Q. L. et al. Oxidation of trimethylamine to trimethylamine N-oxide facilitates high hydrostatic pressure tolerance in a generalist bacterial lineage. Sci. Adv. 7, eabf9941 (2021).Article 

Google Scholar 
Mestre, M. et al. Sinking particles promote vertical connectivity in the ocean microbiome. Proc. Natl Acad. Sci. USA 115, E6799–E6807 (2018).Article 

Google Scholar 
Thiele, S., Fuchs, B. M., Amann, R. & Iversen, M. H. Colonization in the photic zone and subsequent changes during sinking determine bacterial community composition in marine snow. Appl. Environ. Microbiol. 81, 1463–1471 (2015).Article 

Google Scholar 
Tada, Y. et al. Differing growth responses of major phylogenetic groups of marine bacteria to natural phytoplankton blooms in the western North Pacific Ocean. Appl. Environ. Microbiol. 77, 4055–4065 (2011).Article 

Google Scholar 
Cottrell, M. T. & Kirchman, D. L. Natural assemblages of marine proteobacteria and members of the Cytophaga-Flavobacter cluster consuming low- and high-molecular-weight dissolved organic matter. Appl. Environ. Microbiol. 66, 1692–1697 (2000).Article 

Google Scholar 
Poff, K. E., Leu, A. O., Eppley, J. M., Karl, D. M. & DeLong, E. F. Microbial dynamics of elevated carbon flux in the open ocean’s abyss. Proc. Natl Acad. Sci. USA 118, e2018269118 (2021).Article 

Google Scholar 
Ducklow, H. in Microbial Ecology of the Oceans (ed. Kirchman, D. L.) Ch. 4, 85–120 (Wiley-Liss, 2000).Herndl, G. J. et al. Contribution of archaea to total prokaryotic production in the deep Atlantic Ocean. Appl. Environ. Microbiol. 71, 2303–2309 (2005).Article 

Google Scholar 
Baltar, F., Aristegui, J., Gasol, J. M. & Herndl, G. J. Prokaryotic carbon utilization in the dark ocean: growth efficiency, leucine-to-carbon conversion factors, and their relation. Aquat. Microb. Ecol. 60, 227–232 (2010).Article 

Google Scholar 
Edgcomb, V. P. et al. Comparison of Niskin vs. in situ approaches for analysis of gene expression in deep Mediterranean Sea water samples. Deep-Sea Res. II 129, 213–222 (2016).Article 

Google Scholar 
Cario, A., Oliver, G. C. & Rogers, K. L. Exploring the deep marine biosphere: challenges, innovations, and opportunities. Front. Earth Sci. 7, 225 (2019).Article 

Google Scholar 
Giering, S. L. C. et al. Reconciliation of the carbon budget in the ocean’s twilight zone. Nature 507, 480–483 (2014).Article 

Google Scholar 
Simon, M. & Azam, F. Protein content and protein synthesis rates of planktonic marine bacteria. Mar. Ecol. Prog. Ser. 51, 201–213 (1989).Article 

Google Scholar 
Gasol, J. M. et al. Mesopelagic prokaryotic bulk and single-cell heterotrophic activity and community composition in the NW Africa-Canary Islands coastal-transition zone. Prog. Oceanogr. 83, 189–196 (2009).Article 

Google Scholar 
DeLong, E. F. et al. Community genomics among stratified microbial assemblages in the ocean’s interior. Science 311, 496–503 (2006).Article 

Google Scholar 
Teira, E., Reinthaler, T., Pernthaler, A., Pernthaler, J. & Herndl, G. J. Combining catalyzed reporter deposition-fluorescence in situ hybridization and microautoradiography to detect substrate utilization by bacteria and archaea in the deep ocean. Appl. Environ. Microbiol. 70, 4411–4414 (2004).Article 

Google Scholar 
Woebken, D., Fuchs, B. M., Kuypers, M. M. M. & Amann, R. Potential interactions of particle-associated anammox bacteria with bacterial and archaeal partners in the Namibian upwelling system. Appl. Environ. Microbiol. 73, 4648–4657 (2007).Article 

Google Scholar 
Wand, M. P. Data-based choice of histogram bin width. Am. Stat. 51, 59–64 (1997).
Google Scholar 
Acinas, S. G. et al. Deep ocean metagenomes provide insight into the metabolic architecture of bathypelagic microbial communities. Commun. Biol. 4, 604 (2021).Article 

Google Scholar 
Sunagawa, S. et al. Structure and function of the global ocean microbiome. Science 348, 1261359 (2015).Article 

Google Scholar 
Delmont, T. O. et al. Nitrogen-fixing populations of Planctomycetes and Proteobacteria are abundant in surface ocean metagenomes. Nat. Microbiol. 3, 804–813 (2018).Article 

Google Scholar 
Li, D., Liu, C. M., Luo, R., Sadakane, K. & Lam, T. W. MEGAHIT: an ultra-fast single-node solution for large and complex metagenomics assembly via succinct de Bruijn graph. Bioinformatics 31, 1674–1676 (2015).Article 

Google Scholar 
Wu, Y. W., Tang, Y. H., Tringe, S. G., Simmons, B. A. & Singer, S. W. MaxBin: an automated binning method to recover individual genomes from metagenomes using an expectation-maximization algorithm. Microbiome 2, 26 (2014).Article 

Google Scholar 
Kang, D. D. et al. MetaBAT 2: an adaptive binning algorithm for robust and efficient genome reconstruction from metagenome assemblies. Peerj 7, e7359 (2019).Article 

Google Scholar 
Olm, M. R., Brown, C. T., Brooks, B. & Banfield, J. F. dRep: a tool for fast and accurate genomic comparisons that enables improved genome recovery from metagenomes through de-replication. ISME J. 11, 2864–2868 (2017).Article 

Google Scholar 
Chaumeil, P. A., Mussig, A. J., Hugenholtz, P. & Parks, D. H. GTDB-Tk: a toolkit to classify genomes with the Genome Taxonomy Database. Bioinformatics 36, 1925–1927 (2020).
Google Scholar 
Hyatt, D. et al. Prodigal: prokaryotic gene recognition and translation initiation site identification. BMC Bioinf. 11, 119 (2010).Article 

Google Scholar 
Li, W. & Godzik, A. Cd-hit: a fast program for clustering and comparing large sets of protein or nucleotide sequences. Bioinformatics 22, 1658–1659 (2006).Article 

Google Scholar 
Eng, J. K., McCormack, A. L. & Yates, J. R. An approach to correlate tandem mass spectral data of peptides with amino acid sequences in a protein database. J. Am. Soc. Mass. Spectrom. 5, 976–989 (1994).Article 

Google Scholar 
Elias, J. E. & Gygi, S. P. Target-decoy search strategy for increased confidence in large-scale protein identifications by mass spectrometry. Nat. Methods 4, 207–214 (2007).Article 

Google Scholar 
Riffle, M. et al. MetaGOmics: a web-based tool for peptide-centric functional and taxonomic analysis of metaproteomics data. Proteomes 6, 2 (2017).Article 

Google Scholar 
Reinthaler, T., van Aken, H. M. & Herndl, G. J. Major contribution of autotrophy to microbial carbon cycling in the deep North Atlantic’s interior. Deep-Sea Res. II 57, 1572–1580 (2010).Article 

Google Scholar 
Yokokawa, T., Yang, Y. H., Motegi, C. & Nagata, T. Large-scale geographical variation in prokaryotic abundance and production in meso- and bathypelagic zones of the central Pacific and Southern Ocean. Limnol. Oceanogr. 58, 61–73 (2013).Article 

Google Scholar 
Frank, A. H., Garcia, J. A., Herndl, G. J. & Reinthaler, T. Connectivity between surface and deep waters determines prokaryotic diversity in the North Atlantic Deep Water. Environ. Microbiol. 18, 2052–2063 (2016).Article 

Google Scholar 
Herndl, G. J., Bayer, B., Baltar, F. & Reinthaler, T. Prokaryotic life in the deep ocean’s water column. Annu. Rev. Mar. Sci. (in the press).Uchimiya, M., Ogawa, H. & Nagata, T. Effects of temperature elevation and glucose addition on prokaryotic production and respiration in the mesopelagic layer of the western North Pacific. J. Oceanogr. 72, 419–426 (2016).Article 

Google Scholar 
Antia, A. N. et al. Basin-wide particulate carbon flux in the Atlantic Ocean: regional export patterns and potential for atmospheric CO2 sequestration. Glob. Biogeochem. Cycles 15, 845–862 (2001).Article 

Google Scholar 
Behrenfeld, M. J. & Falkowski, P. G. Photosynthetic rates derived from satellite-based chlorophyll concentration. Limnol. Oceanogr. 42, 1–20 (1997).Article 

Google Scholar  More

Austin, K. G. et al. Land Use Policy 69, 41–48 (2017).Article 

Google Scholar 
Busch, J. et al. Environ. Res. Lett. 17, 014035 (2022).Article 
CAS 

Google Scholar 
Fleiss, S. et al. Nat. Ecol. Evol. https://doi.org/10.1038/s41559-022-01941-6 (2022).Qaim, M. et al. Annu. Rev. Resour. Econ. 12, 321–344 (2020).Article 

Google Scholar 
Haupt, F. et al. Progress on Corporate Commitments and their Implementation (Tropical Forest Alliance, 2018).Brooks, T. et al. Nat. Ecol. Evol. 1, 0099 (2017).Article 

Google Scholar 
Buisson, E. et al. Biol. Rev. 94, 590–609 (2019).Article 
PubMed 

Google Scholar 
López-Ricaurte, L. et al. Biol. Conserv. 213, 225–233 (2017).Article 

Google Scholar 
Furumo, P. R. & Aide, T. M. Environ. Res. Lett. 12, 024008 (2017).Article 

Google Scholar 
RTRS Standard for Responsible Soy Production Version 3.1 (RTRS, 2017). More

General description of sequencesAfter the quality filtering step, removal of chimeric fragments, and read merging, a total of 3,378,323 reads with 3007 different features was obtained, with an average of 27,244 sequences per individual sample. After quality filtering, none of the samples was excluded from the analysis of microbial communities.Amoxicillin and thiamphenicol treatments influence microbial diversity and the abundance of specific taxaUsing 16S rRNA NGS, the gut microbial community composition of the chicks in each group was characterized at different time points. At phylum level, microbiota composition varied with age rather than with treatment (Supplementary Fig. S1). Proteobacteria were the most abundant phyla at 1 day of age (d.o.a.), Firmicutes became dominant at later stages, while Bacteroidota were highly abundant in caecum samples collected at 46 d.o.a. Similar dynamics were observed also at family level, since Enterobacteriaceae and Clostridiaceae were significantly more abundant at 1 d.o.a. in all groups, Lactobacillaceae, Lachnospiraceae, and Ruminococcaceae seemed to bloom at 8 d.o.a., and Rikenellaceae were the dominant family in the caecum samples collected at 46 d.o.a. (Fig. 1; Supplementary Fig. S2).Figure 1Heatmap representing the microbial community composition at family level. The heatmap was generated in R (version 4.2.1) (https://www.r-project.org/) using package pheatmap (version 1.0.12).Full size imageEarly-age administrationIn both α-diversity indices (Fig. 2A,B), there was a trend towards increasing diversity from early to late time points in all groups; however, the only significant differences were between the group treated with amoxicillin (AMX1) and the other groups on day 21 post treatment (p.t.), and within AMX1 group between day 21 p.t. and the other time points. PERMANOVA showed that the microbial community was significantly different between the group treated with thiamphenicol (THP1) and the other two groups (i.e. AMX1 and control) on day 1 p.t. (p  More

Total catch control regulation does not lead to the recovery of fisheries and the maintenance of community functionTo contain the decline of wild capture fisheries by overfishing, a series of management regulations have been in place in China to mitigate the fishing impacts as much as possible and maintain sustainable stocks. The “zero-growth” policy is one of the most outstanding representatives. The results showed certain achievements after the implementation of the policy. Simulating the status without the “zero-growth” policy, B/Bmsy fell below 0.5 by 2010 and close to zero by 2019, indicating the impossibility for recovery. However, the policy is not enough for fishery recovery and community health, failing to stop the degradation of fishery resources. Under the implementation of the “zero-growth” policy, B/Bmsy was in a healthy state in 1998, fell below 1 for the first time in 2003, and dropped to 0.52 in 2019, accompanying by F/Fmsy as 1.60. If fishing pressure were maintained at the level of 2019 (F = 1.56 Fmsy), the resource would decline to the depletion state by 2030 (B/Bmsy close to zero, F/Fmsy = 3.64, catch = 35 T). Therefore, a great degree of negative production growth as well as the strict implementation is extremely important. A rapid reduction in the catch control under 0.5 Fmsy scenario would expect to achieve a quick recovery with B/Bmsy over 1 in 2025. Nevertheless, a significant reduction in production would lead to the decline of fishery economics, livelihood difficulties for fishermen and a series of derivative social problems28. An alternative of 1.0 Fmsy would be feasible, under which B/Bmsy could rise to 1 by 2030 with a production of 11.64 MT, close to MSY.The “zero growth” policy faces some inherent challenges, at least from the point of view of ensuring the sustainable use of individual species stocks. Attention should be also paid at the catch quota control of individual species. Because the variation of the intrinsic growth rate of different species, the B is dynamic, and the F changes with the change of B. In a constant production, r-strategic species could remain a higher B/Bmsy than 1 even at a large proportion in catch, but K-strategic species did not show the same fortune. The control of total catch volume rather than individual species could not prevent the community structure from becoming fragile, with the exhaustion of high-trophic species and the decrease of mean trophic level.Individual species have different responses to overfishing that highly associated with their biological characteristicsHigh trophic level species can be sensitive to overfishing, and difficult to rebuild stocks after collapseHairtails Trichiurus spp. are the largest contribution group to China marine capture fisheries, at 0.90 MT about 8.3% of the total production in 20202. They are carnivorous and aggressive with a mean trophic level of 4.4, mainly feeding on fishes in the adult stage, and Mysidacea and Euphausiacea in the juvenile stage29,30. The spawning seasons of Trichiurus spp. are mainly from April to June, and from September to November in Chinese waters31.China coastal areas are excellent foraging and spawning grounds for Trichiurus spp, sustaining a large stock size. If the “zero-growth” policy was not implemented since 1999, the resources of Trichiurus spp. would be exhausted by 2027, having no possibility to recovery at 1.0 Fmsy. Although the total fisheries production has been controlled, and the fishing moratorium period partly covered the spawning seasons of Trichiurus spp., their resource continuous declined into a “destroying” state in 2007, due to the time-lag effect of fishing on high trophic level predators characterized by long population doubling time-consuming32. Under intensive fishing pressure, Trichiurus spp. have showed astonishing fisheries-induced adaption33 by reducing the age and size of maturity, which effectively alleviates the decline rate of B value, resulting the maintenance of Trichiurus spp. capture production. Under the rebuilding scenario of fishing pressure as 1.0 Fmsy, Trichiurus spp. B/Bmsy rose to 0.87 by 2030, lower than the recovery rate of national total capture fisheries, suggesting the recovery rate of high trophic level species could be slow34. Furthermore, in this study fisheries rebuilding only considers the responses of species to fishing pressure, irrespective of a series of factors sensitive to high trophic level species such as pollution and climate change, which indicated a longer period is needed for resource recovery.Middle trophic level species seems non sensitive to total catch control policyAs a representative of middle trophic level species, L. polyactis performed different from Trichiurus spp. Under high fishing pressure. It forms spawning and over-wintering aggregations between nearshore and offshore waters, as well as vertical migration, rising at dusk and falling at dawn35. The spawning season is from mid-February to early May, prior to the national fishing moratorium, indicating young juveniles are in effective protection rather than spawning stock. In the 1950s, L. polyactis was one of the few important species in domestic marine capture fisheries in Chinese waters, producing more than 100,000 T annually5. The catch volumes then showed a downward trend and fell significantly to less than 50,000 T in the 1960–1980s. After 3 decades low catch volumes, the annual capture production rebounded significantly to more than 200,000 T and maintained at such high levels for 2 decades5, showing high resilience to overfishing.Despite many concerns on the risk of resource exhaustion of L. polyactis stocks5,36, official statistics showed that the annual catch remains high. The L. polyactis production broke through 150,000 tons in 1995, and was above 300,000 tons after 2005. There is likely to have a large offshore stock of L. polyactis, which gradually joined the catch under increasing fishing efforts offshore. Furthermore, the L. polyactis stocks can be resilience to high pressure for several reasons: (1) its miscellaneous diet makes them be able to receive sufficient food sources; (2) size and age at sexual maturity reduced37,38; and (3) the over consumption of top predators relieves the prey pressure on middle trophic level species, such as L. polyactis, snappers, and flatfishes. A good job is the difficulties of artificial propagation and seedling breeding of small yellow croaker were broken for the first time in 201539 and the whole artificial cultivation was successfully realized in 2020 (https://www.chinanews.com.cn/cj/2020/07-02/9227715.shtml), which would effectively alleviate the market demand and wild stock sustain of small yellow croaker.Pelagic small fish stocks may not recovery quickly as early cognitionSmall pelagic fishes enjoy assembling in large schools of tens of thousands of individuals, and are more vulnerable to predators. Species S. sagax mainly filter plankton with a low trophic level about 2.8. It spawns in May–June, with high fecundity (an absolute fecundity of 30,000–100,000 pelagic eggs) and fast growth, and has short generation time of 1.4 years40. S. sagax shows strong phototropy, and can be caught using light purse seine, gill net, and fixed net fishing at night41,42.In 1989, the biomass of S. sagax was about twice of Bmsy. With the decreasing capture production of traditional economic fishes, S. sagax became a target species using specific fishing methods43, resulted in catch increase accompanied with B/Bmsy decline into a state of extremely unhealthy in 2019. Recovery of small pelagic species stocks would be delayed by the total catch control policy, mainly because the removal of large numbers of predator species left more opportunities for their feeding objects44. Resource rebuilding of S. sagax was not as quick as expected, as small pelagic species had to endure increasing predation pressure from the recovery of high-trophic species under the total catch control. At 1.0 Fmsy scenario, B/Bmsy would be only 0.88 by 2030, in need of a longer period to healthy state.Well-planned restocking can enhance resource recoverySwimming crab P. trituberculatus has high reproductive capacity, with a female can release two to three batches of eggs during a breeding season, and a batch contains about 1–6 million eggs45. Under the complementary of existing management measures and restocking programmes, the production of P. trituberculatus was kept in a certain amount close to a healthy state, and there is not an urgent need for its stock rebuilding. Since the 1990s, restocking of hatchery-produced larvae of P. trituberculatus has been promoted in coastal waters of China. Large-scaled restocking programmes were documented: 33 million larvae were released into the Yellow Sea by Shandong Province in June 2013 (http://hyj.shandong.gov.cn/xwzx/sjdt/201311/t20131120_507389.html); 50.3 million larvae with carapace width over 6 mm were released in the northern Yellow Sea by Liaoning Province in June 2020 (http://nync.ln.gov.cn/fwzx/zxdt/202007/t20200707_3902016.html); 16.1 million larvae were released into the East China Sea by Daishan County of Zhejiang Province in June 2021 (http://www.daishan.gov.cn/art/2021/6/8/art_1383064_59012675.html). What should be of concern is when, where, and how many seedlings are released46,47,48, to maximumly utilize the environmental resources without encroaching on the benefits of other species.Short-living species can be resilience to overfishingThe main cephalopod species in Chinese fisheries are Sepiella maindroni, mainly distributes in the East China Sea35 and Sepia esculenta, mainly distributes in the Bohai Sea, the Yellow Sea and the East China Sea49. As a 1-year lifespan species with fast growth rate, S. maindroni forms spawning migration from deep water to shallow nearshore bays in spring, partly within the fishing moratorium period. Due to the positive phototaxis, the cuttlefishes can be captured by light seining. Sepiella esculenta was the most important cephalopod economically in the northern coastal seas and one of the four major fisheries in the Bohai Sea and the Yellow Sea until the 1970s50. The abundance of this species has been greatly reduced with continuous fishing pressures and dwindling spawning grounds51.Total catch control and fishing moratorium showed significant output on the short lifespan cuttlefishes. Without the implementation of the “zero-growth” policy, the cuttlefishes resources would have been exhausted by 2015 and impossible to rebuild. According to the current state of resources, by 2030 the cuttlefish stocks can be recovered under the 1.0 Fmsy scenario. Moreover, the extent of cuttlefishes stock recovery relies on food supply.Ways to sustain fisheriesThe conflict between rising demand for fishery products and declining resources under multiple pressures including overfishing, climate change, and marine pollution has put heavy pressures at a global scale52. Chinese government has undertaken serious reforms to effectively replan the fishery industry.The effective recovery and rational utilization of resources depend on the support by sufficient reliable data. China started fishery statistics right after the foundation of the People’s Republic of China, completed by MOA (1949–2017) and MARA (since 2018). However, the statistical dataset has been questioned internationally53. According to the explanation by FAO54, before 2000s, especially from 1979 to the late 1990s, as the central government raced to meet the increasing demand for seafood and to grow the domestic production, the local governments had frequently overreported their local catch. In addition, fishermen may falsely claim to increase their production for surplus compensation, after the government introduced fishing subsidies. On the contrary, the production might have been underreported since the early 2000s55,56, which could be attributed to the existence of a large number of “black ships” (fishing vessels without relevant legal permits). Moreover, the lack of professionals in the early period and inaccurate knowledge of species identification by fishermen also lead to data uncertainty. Reasonable fisheries data should be consistent with the species functional traits and life history characteristics. However, in the actual fishing activities, the intentional and high-intensity selective fishing of species may greatly deviate the catch data from the data predicted by models. The Chinese government has been trying to improve the statistical system, including data coefficient adjustment, training of fishermen and professional, and supervision of statistical authorities5. In this study, selected objects are inshore species: the species are familiar to fishermen; the fishing vessel supervision is in place; the data collection is relatively rational and complete; all these are conducive to the reliability of the results.The zero-growth policy, which has been implemented since 1999, is an important measure in the history of marine fishery development and management in China. That is, the total catch of marine fisheries in the current year cannot be higher than that of the previous year. However, the “12th Five-year Plan” for national fishery development (2011–2015) issued by the Ministry of Agriculture canceled the mandatory targets of controlling the production but to encourage more catches of marine fisheries (http://www.moa.gov.cn/gk/ghjh_1/201110/t20111017_2357716.htm). In 2013, the State Council published the first state-level marine fishery development document as “Several Advices on Promoting Marine Sustainable and Healthy Development”, incorporating marine fishery development into the strategy of building a maritime power (http://www.gov.cn/zwgk/2013-06/25/content_2433577.htm). This policy shift was clearly reflected in the significant increase in the national annual catch from 12 to 14 MT. Until the “13th Five-year Plan” for national fishery development (2016–2020) issued in 2016, the zero-even negative-growth policy was revalidated, and the volume of annual output control was clearly proposed as 8–10 MT57, which was determined by multiplying the fishing coefficient by the total stock size derived from the assessment of surveys on the zoning of fisheries and the supplementary survey of marine biological resources in the exclusive economic zone and the continental shelf7. To achieve the target of keeping fishing capacity at a high level of sustainability, significant reductions in fishing pressures over a period of time are required, as well as rational updates of control policies.Many policies were introduced together or around the same time as the “zero-growth” policy, such as summer fishing moratorium, fishing license system, and fishing fuel subsides. However, the achievements are far from satisfactory. The fishing fuel subsidy policy together with the license system induced the direct fishing vessel construction boom which resulted in fewer but bigger and more powerful fishing vessels. Fishing moratorium is the most promising policy, by leaving enough time and space for fish to successfully reproduce. However, the truth is that, right after the fishing closure season, almost all fishing vessels immediately rush into the sea and fishermen try their best to fish as much as possible within the gears and engine power permission of their fishing licenses, attempting to earn a year’s income in a short period of 2–4 months. As a result of such high fishing effort, the achievements of seasonal fishing bans were largely offset and resource densities fell to low levels after autumn. The number of legally binding standards for mesh size is not enough, only 6 at present of at least 40 fishing target species and over 10 fishing gears, leaving many fishing gears and fish species outside the regulation of existing standards6,58. Ideally, standards of mesh size should be updated corresponding to the changes of species traits, however, it is a challenge because the main fishing mode is multiple species fishery by bottom trawling. Moreover, species in China seas are diverse, and the spawning period of different species may not fall into the fishing closure season5. The lack of specificity to sufficiently cover all the species may result an unbalance of community composition. Another system “Double Control” aims to limit both the numbers of fishing vessels and the total power. Unfortunately, the inspections of fishing vessels and their power are not very strict, due to the need of developing local economy and guaranteeing the fishermen’s income, e.g., under a nominal power mask the low-power engines have been replaced by high-power engines, some fishing vessels do not have the fishing licenses28. The limitation of the license number and engine power also stimulate the technological improvement for more catch7.The structure adjustment of fisheries composition is the main management measure at present. The high degree of self-sufficiency in fishery products in China has been achieved through overfishing of domestic fishery resources, resulting in the rapid depletion of fisheries in China’s coastal waters59. Aquaculture, accounting for more than 70% of China’s total fisheries production2, is identified as a successful way. Accompanying by aquaculture development, a series of problems also arise, particularly, the demand of low-value/trash fish and fish meal that significantly drives further expansion of capture fisheries60. Cooperation with other countries to promote regional aquaculture may be an alternative way to meeting global growing demand for seafood and combating overfishing61,62. Seeking resources from the high seas and EEZs of other countries is also a choice, of course, on the premise of taking full account of ecology, maritime, and food security of other countries63,64,65.In addition, this study pointed out a new focus for fisheries management, in which differences in species biological traits, including species vulnerability, population multiplication, and resilience to environmental pressures, should be given full consideration. On this basis, more detailed and targeted management schemes are supposed to propose to achieve the dual purpose of recoverable fisheries resource and balanced species composition, so as to become a truly sustainable fishery. In short, the effective implementation of various management measures is an indispensable guarantee. More

Way, M. J., Hopkins, B. & Smith, P. M. Photoperiodism and diapause in insects. Nature 164, 615 (1949).Article 
PubMed 

Google Scholar 
Beck, S. Photoperiod induction of diapause in an insect. Biol. Bull. 122, 1–12 (1962).Article 

Google Scholar 
Denlinger, D. L. & Armbruster, P. A. Mosquito diapause. Annu. Rev. Entomol. 59, 73–93 (2014).Article 
PubMed 

Google Scholar 
Readio, J., Chen, M. H. & Meola, R. Juvenile hormone biosynthesis in diapausing and nondiapausing Culex pipiens (Diptera: Culicidae). J. Med. Entomol. 36, 355–360 (1999).Article 
PubMed 

Google Scholar 
Eldridge, B. F. & Bailey, C. L. Experimental hibernation studies in Culex pipiens (Diptera: Culicidae): reactivation of ovarian development and blood-feeding in prehibernating females. J. Med Entomol. 15, 462–467 (1979).Article 
PubMed 

Google Scholar 
Spielman, A. & Wong, J. Environmental control of ovarian diapause in Culex pipiens. Ann. Entomol. Soc. Am. 66, 905–907 (1973).Article 

Google Scholar 
Sanburg, L. L. & Larsen, J. R. Effect of photoperiod and temperature on ovarian development in Culex pipiens pipiens. J. Insect Physiol. 19, 1173–1190 (1973).Article 
PubMed 

Google Scholar 
Eldridge, B. F. The effect of temperature and photoperiod on blood-feeding and ovarian development in mosquitoes of the Culex pipiens complex. Am. J. Trop. Med. Hyg. 17, 133–140 (1968).Article 
PubMed 

Google Scholar 
Bowen, M. F. Patterns of sugar feeding in diapausing and nondiapausing Culex pipiens (Diptera: Culicidae) females. J. Med. Entomol. 29, 843–849 (1992).Article 
PubMed 

Google Scholar 
Robich, R. M. & Denlinger, D. L. Diapause in the mosquito Culex pipiens evokes a metabolic switch from blood feeding to sugar gluttony. Proc. Natl Acad. Sci. USA 102, 15912–15917 (2005).Article 
PubMed 
PubMed Central 

Google Scholar 
Eldridge, B. F. Environmental control of ovarian development in mosquitoes of the Culex pipiens complex. Am. Assoc. Adv. Sci. 151, 826–828 (1966).
Google Scholar 
Vinogradova, A. B. Culex pipiens Pipiens Mosquitoes: Taxonomy, Distribution, Ecology, Physiology, Genetics, Applied Importance And Control (Pensoft, 2000).Benoit, J. B. & Denlinger, D. L. Suppression of water loss during adult diapause in the northern house mosquito, Culex pipiens. J. Exp. Biol. 210, 217–226 (2007).Article 
PubMed 

Google Scholar 
Li, A. & Denlinger, D. L. Pupal cuticle protein is abundant during early adult diapause in the mosquito Culex pipiens. J. Med. Entomol. 46, 1382–1386 (2009).Article 
PubMed 

Google Scholar 
Yang, L., Denlinger, D. L. & Piermarini, P. M. The diapause program impacts renal excretion and molecular expression of aquaporins in the northern house mosquito, Culex pipiens. J. Insect Physiol. 98, 141–148 (2017).Article 
PubMed 

Google Scholar 
King, B., Li, S., Liu, C., Kim, S. J. & Sim, C. Suppression of glycogen synthase expression reduces glycogen and lipid storage during mosquito overwintering diapause. J. Insect Physiol. 120, 103971 (2020).Article 
PubMed 

Google Scholar 
Sim, C. & Denlinger, D. L. Transcription profiling and regulation of fat metabolism genes in diapausing adults of the mosquito Culex pipiens. Physiol. Genomics 39, 202–209 (2009).Article 
PubMed 
PubMed Central 

Google Scholar 
Sim, C. & Denlinger, D. L. Insulin signaling and FOXO regulate the overwintering diapause of the mosquito Culex pipiens. Proc. Natl Acad. Sci. USA 105, 6777–6781 (2008).Article 
PubMed 
PubMed Central 

Google Scholar 
Zhou, G. & Miesfeld, R. L. Energy metabolism during diapause in Culex pipiens mosquitoes. J. Insect Physiol. 55, 40–46 (2009).Article 
PubMed 

Google Scholar 
Chang, J. et al. Solid-state NMR reveals differential carbohydrate utilization in diapausing Culex pipiens. Sci. Rep. 6, 37350 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Madder, D. J., Surgeoner, G. A. & Helson, B. V. Induction of diapause in Culex pipiens and Culex restuans (Diptera: Culicidae) in Southern Ontario. Can. Entomol. 115, 877–883 (1983).Article 

Google Scholar 
Spielman, A. Effect of synthetic juvenile hormone on ovarian diapause of Culex pipiens mosquitoes. J. Med. Entomol. 11, 223–225 (1974).Article 
PubMed 

Google Scholar 
Sim, C. & Denlinger, D. L. Insulin signaling and the regulation of insect diapause. Front. Physiol. 4, 189 (2013).Article 
PubMed 
PubMed Central 

Google Scholar 
Robich, R. M., Rinehart, J. P., Kitchen, L. J. & Denlinger, D. L. Diapause-specific gene expression in the northern house mosquito, Culex pipiens L., identified by suppressive subtractive hybridization. J. Insect Physiol. 53, 235–245 (2007).Article 
PubMed 

Google Scholar 
Sim, C., Kang, D. S., Kim, S., Bai, X. & Denlinger, D. L. Identification of FOXO targets that generate diverse features of the diapause phenotype in the mosquito Culex pipiens. Proc. Natl Acad. Sci. USA 112, 3811–3816 (2015).Article 
PubMed 
PubMed Central 

Google Scholar 
Kang, D. S., Cotten, M. A., Denlinger, D. L. & Sim, C. Comparative transcriptomics reveals key gene expression differences between diapausing and non-diapausing adults of Culex pipiens. PLoS ONE 11, e0154892 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Spielman, A. Structure and seasonality of nearctic Culex pipiens populations. Ann. N. Y. Acad. Sci. 951, 220–234 (2001).Article 
PubMed 

Google Scholar 
Wilton, D. P. & Smith, G. C. Ovarian diapause in three geographic strains of Culex pipiens (Diptera: Culicidae). J. Med. Entomol. 22, 524–528 (1985).Article 
PubMed 

Google Scholar 
Eldridge, B. F. Diapause and related phenomena in Culex mosquitoes: their relation to arbovirus disease ecology. In: Current Topics in Vector Research (ed. Harris, K. F.) 1–28 (Springer, 1987).Meuti, M. E., Short, C. A. & Denlinger, D. L. Mom matters: diapause characteristics of Culex pipiens-Culex quinquefasciatus (Diptera: Culicidae) hybrid mosquitoes. J. Med. Entomol. 52, 131–137 (2015).Article 
PubMed 
PubMed Central 

Google Scholar 
Zhang, C. et al. Understanding the regulation of overwintering diapause molecular mechanisms in Culex pipiens pallens through comparative proteomics. Sci. Rep. 9, 6845 (2019).PubMed 
PubMed Central 

Google Scholar 
Dunphy, B. M. et al. Long-term surveillance defines spatial and temporal patterns implicating Culex tarsalis as the primary vector of West Nile virus. Sci. Rep. 9, 6637 (2019).Article 
PubMed 
PubMed Central 

Google Scholar 
Dunphy, B. M., Rowley, W. A. & Bartholomay, L. C. A Taxonomic checklist of the mosquitoes of Iowa. J. Am. Mosq. Control Assoc. 30, 119–121 (2014).Article 
PubMed 

Google Scholar 
Sucaet, Y., Van Hemert, J., Tucker, B. & Bartholomay, L. C. A web-based relational database for monitoring and analyzing mosquito population dynamics. J. Med. Entomol. 45, 775–784 (2008).Article 
PubMed 

Google Scholar 
Ryan, S. F., Valella, P., Thivierge, G., Aardema, M. L. & Scriber, J. M. The role of latitudinal, genetic and temperature variation in the induction of diapause of Papilio glaucus (Lepidoptera: Papilionidae). Insect Sci. 25, 328–336 (2018).Article 
PubMed 

Google Scholar 
Huang, L. et al. Diapause incidence and critical day length of Asian corn borer (Ostrinia furnacalis) populations exhibit a latitudinal cline in both pure and hybrid strains. J. Pest Sci. 93, 559–568 (2020).Article 

Google Scholar 
Bradshaw, W. E. Geography of photoperiodic response in diapausing mosquito. Nature 262, 384–386 (1976).Article 
PubMed 

Google Scholar 
Bradshaw, W. E. & Lounibos, L. P. Evolution of dormancy and its photoperiodic control in pitcher-plant mosquitoes. Nature 31, 546–567 (1977).
Google Scholar 
Kothera, L., Zimmerman, E. M., Richards, C. M. & Savage, H. M. Microsatellite characterization of subspecies and their hybrids in Culex pipiens complex (Diptera: Culicidae) mosquitoes along a North-South transect in the central United States. J. Med. Entomol. 46, 236–248 (2009).Article 
PubMed 

Google Scholar 
Darsie, R. F. R. & Ward, R. A. R. Identification and Geographical Distribution of the Mosquitoes of North America, North of Mexico (University Press of Florida, 2005).Huang, S., Molaei, G. & Andreadis, T. G. Reexamination of Culex pipiens hybridization zone in the eastern United States by ribosomal DNA-based single nucleotide polymorphism markers. Am. J. Trop. Med. Hyg. 85, 434–441 (2011).Article 
PubMed 
PubMed Central 

Google Scholar 
Reisen, W. K. Overwintering studies on Culex tarsalis (Diptera: Culicidae) in Kern County, California: life stages sensitive to diapause induction cues. Ann. Entomol. Soc. Am. 79, 674–676 (1986).Article 

Google Scholar 
Haba, Y. & McBride, L. Origin and status of Culex pipiens mosquito ecotypes. Curr. Biol. 32, R237–R246 (2022).Article 
PubMed 

Google Scholar 
Holzapfel, C. M. & Bradshaw, W. E. Geography of larval dormancy in the tree-hole mosquito, Aedes triseriatus (Say). Can. J. Zool. 59, 1014–1021 (1981).Article 

Google Scholar 
Rinehart, J. P., Robich, R. M. & Denlinger, D. L. Enhanced cold and desiccation tolerance in diapausing adults of Culex pipiens, and a role for Hsp70 in response to cold shock but not as a component of the diapause program. J. Med. Entomol. 43, 713–722 (2006).Article 
PubMed 

Google Scholar 
Faraji, A. & Gaugler, R. Experimental host preference of diapause and non-diapause induced Culex pipiens pipiens (Diptera: Culicidae). Parasit. Vectors 8, 389 (2015).Article 
PubMed 
PubMed Central 

Google Scholar 
Washino, R. K. The physiological ecology of gonotrophic dissociation and related phenomena in mosquitoes. J. Med. Entomol. 13, 381–388 (1977).Article 
PubMed 

Google Scholar 
Christophers, S. The development of the egg follicle in Anophelines. Paludism 1, 73–88 (1911).
Google Scholar 
Nelms, B. M., Macedo, P. A., Kothera, L., Savage, H. M. & Reisen, W. K. Overwintering biology of Culex (Diptera: Culicidae) mosquitoes in the Sacramento Valley of California. J. Med. Entomol. 50, 773–790 (2013).Article 
PubMed 

Google Scholar 
Diniz, D. F. A., De Albuquerque, C. M. R., Oliva, L. O., De Melo-Santos, M. A. V. & Ayres, C. F. J. Diapause and quiescence: dormancy mechanisms that contribute to the geographical expansion of mosquitoes and their evolutionary success. Parasites Vectors 10, 1–13 (2017).Article 

Google Scholar 
Kingsolver, J. G. & Nagle, A. Evolutionary divergence in thermal sensitivity and diapause of field and laboratory populations of Manduca sexta. Physiol. Biochem. Zool. 80, 473–479 (2007).Article 
PubMed 

Google Scholar 
Brent, C. S. & Spurgeon, D. W. Diapause response of laboratory reared and native lygus hesperus knight (Hemiptera: Miridae). Environ. Entomol. 40, 455–461 (2011).Article 

Google Scholar 
Rinehart, J. P., Yocum, G. D., Leopold, R. A. & Robich, R. M. Cold storage of Culex pipiens in the absence of diapause. J. Med. Entomol. 47, 1071–1076 (2014).Article 

Google Scholar 
Arora, A. K., Sim, C., Severson, D. W. & Kang, D. S. Random forest analysis of impact of abiotic factors on Culex pipiens and Culex quinquefasciatus occurrence. Front. Ecol. Evol. 9, 773360 (2022).Article 

Google Scholar 
Focks, D. A., Linda, S. B., Craig Jnr, G. B., Hawley, W. A. & Pumpuni, C. B. Aedes albopictus (Diptera: Culicidae): a statistical model of the role of temperature, photoperiod, and geography in the induction of egg diapause. J. Med. Entomol. 31, 278–286 (1994).Article 
PubMed 

Google Scholar 
Urbanski, J. et al. Rapid adaptive evolution of photoperiodic response during invasion and range expansion across a climatic gradient. Am. Nat. 179, 490–500 (2012).Article 
PubMed 

Google Scholar 
Kothera, L., Godsey, M. S., Doyle, M. S. & Savage, H. M. Characterization of Culex pipiens complex (Diptera: Culicidae) populations in Colorado, USA using microsatellites. PLoS ONE 7, e0047602 (2012).Article 

Google Scholar 
Kothera, L., Nelms, B. M., Reisen, W. K. & Savage, H. M. Population genetic and admixture analyses of Culex pipiens complex (Diptera: Culicidae) populations in California, United States. Am. J. Trop. Med. Hyg. 89, 1154–1167 (2013).Article 
PubMed 
PubMed Central 

Google Scholar 
Kothera, L. et al. Bloodmeal, Host selection, and genetic admixture analyses of Culex pipiens Complex (Diptera: Culicidae) mosquitoes in Chicago, IL. J. Med. Entomol. 57, 78–87 (2020).Article 
PubMed 

Google Scholar 
Huang, S., Molaei, G. & Andreadis, T. G. Genetic insights into the population structure of Culex pipiens (Diptera: Culicidae) in the Northeastern United States by using microsatellite analysis. Am. J. Trop. Med Hyg. 79, 518–527 (2008).Article 
PubMed 

Google Scholar 
Barr, A. R. The Distribution of Culex p. pipiens and Cp quinquefasciatus in North America. Am. J. Trop. Med. Hyg. 6, 153–165 (1957).Article 
PubMed 

Google Scholar 
Iltis, W. G. Biosystematics of the Culex pipiens Complex in Northern California. Thesis, University of California, Davis. (1966).Urbanelli, S., Silvestrini, F., Reisen, W. K., De Vito, E. & Bullini, L. Californian hybrid zone between Culex pipiens pipiens and Cx. p. quinquefasciatus revisited (Diptera: Culicidae). J. Med. Entomol. 34, 116–127 (1997).Article 
PubMed 

Google Scholar 
Nelms, B. M. et al. Phenotypic variation among Culex pipiens complex (Diptera: Culicidae) populations from the Sacramento Valley, California: Horizontal and vertical transmission of West Nile virus, diapause potential, autogeny, and host selection. Am. J. Trop. Med. Hyg. 89, 1168–1178 (2013).Article 
PubMed 
PubMed Central 

Google Scholar 
Dodson, B. L., Kramer, L. D. & Rasgon, J. L. Effects of larval rearing temperature on immature development and West Nile virus vector competence of Culex tarsalis. Parasit. Vectors 5, 199 (2012).Article 
PubMed 
PubMed Central 

Google Scholar 
Ciota, A. T., Matacchiero, A. C., Marm Kilpatrick, A. & Kramer, L. D. The effect of temperature on life history traits of Culex mosquitoes. J. Med Entomol. 51, 55–62 (2014).Article 
PubMed 

Google Scholar 
Carrington, L. B., Seifert, S. N., Willits, N. H., Lambrechts, L. & Scott, T. W. Large diurnal temperature fluctuations negatively influence Aedes aegypti (Diptera: Culicidae) life-history traits. J. Med. Entomol. 50, 43–51 (2013).Article 
PubMed 

Google Scholar 
Lambrechts, L. et al. Impact of daily temperature fluctuations on dengue virus transmission by Aedes aegypti. Proc. Natl Acad. Sci. USA 108, 7460–7465 (2011).Article 
PubMed 
PubMed Central 

Google Scholar 
Karki, S., Brown, W. M., Uelmen, J., O’Hara Ruiz, M. & Smith, R. L. The drivers of West Nile virus human illness in the Chicago, Illinois, USA area: fine scale dynamic effects of weather, mosquito infection, social, and biological conditions. PLoS ONE 15, e0227160 (2020).Article 
PubMed 
PubMed Central 

Google Scholar 
Andreadis, T. G., Anderson, J. F., Vossbrinck, C. R. & Main, A. J. Epidemiology of West Nile virus in Connecticut: a five-year analysis of mosquito data 1999–2003. Vector-Borne Zoonotic Dis. 4, 360–378 (2004).Article 
PubMed 

Google Scholar 
Anderson, J. F. & Main, A. J. Importance of vertical and horizontal transmission of West Nile virus by Culex pipiens in the northeastern United States. J. Infect. Dis. 194, 1577–1579 (2006).Article 
PubMed 

Google Scholar 
Nasci, R. S. et al. West Nile virus in overwintering Culex mosquitoes, New York City, 2000. Emerg. Infect. Dis. 7, 742–744 (2001).Article 
PubMed 
PubMed Central 

Google Scholar 
Kampen, H., Tews, B. A. & Werner, D. First evidence of West Nile virus overwintering in mosquitoes in Germany. Viruses 13, 1–7 (2021).Article 

Google Scholar 
Farajollahi, A. et al. Detection of West Nile viral RNA from an overwintering pool of Culex pipens pipiens (Diptera: Culicidae) in New Jersey, 2003. J. Med. Entomol. 42, 490–494 (2005).Article 
PubMed 

Google Scholar 
Baqar, S., Hayes, C. G., Murphy, J. R. & Watts, D. M. Vertical transmission of West Nile virus by Culex and Aedes species mosquitoes. Am. J. Trop. Med. Hyg. 48, 757–762 (1993).Article 
PubMed 

Google Scholar 
Miller, B. R. et al. First field evidence for natural vertical transmission of West Nile virus in Culex univittatus complex mosquitoes from Rift Valley Province, Kenya. Am. J. Trop. Med. Hyg. 62, 240–246 (2000).Article 
PubMed 

Google Scholar 
Peffers, C. S., Pomeroy, L. W. & Meuti, M. E. Critical photoperiod and its potential to predict mosquito distributions and control medically important pests. J. Med. Entomol. 58, 1610–1618 (2021).Article 
PubMed 

Google Scholar 
Bradshaw, W. E. & Holzapfel, C. M. Genetic shift in photoperiodic response correlated with global warming. Proc. Natl Acad. Sci. USA 98, 14509–14511 (2001).Article 
PubMed 
PubMed Central 

Google Scholar 
Reiter, P. Climate change and mosquito-borne disease. Environ. Health Perspect. 109, 141–161 (2001).PubMed 
PubMed Central 

Google Scholar 
Colón-González, F. J. et al. Projecting the risk of mosquito-borne diseases in a warmer and more populated world: a multi-model, multi-scenario intercomparison modelling study. Lancet Planet. Heal. 5, e404–e414 (2021).Article 

Google Scholar 
Barreaux, A. M. G., Stone, C. M., Barreaux, P. & Koella, J. C. The relationship between size and longevity of the malaria vector Anopheles gambiae (s.s.) depends on the larval environment. Parasites Vectors 11, 485 (2018).Article 
PubMed 
PubMed Central 

Google Scholar 
Van Handel, E. & Day, J. F. Correlation between wing length and protein content of mosquitoes. J. Am. Mosq. Control Assoc. 5, 180–182 (1989).PubMed 

Google Scholar 
Ferreira-De-Freitas, L., Thrun, N. B., Tucker, B. J., Melidosian, L. & Bartholomay, L. C. An evaluation of characters for the separation of two Culex species (Diptera: Culicidae) based on material from the Upper Midwest. J. Insect Sci. 20, 21 (2020).Harrington, L. C. & Poulson, R. L. Considerations for accurate identification of adult Culex restuans (Diptera: Culicidae) in field studies. J. Med. Entomol. 45, 1–8 (2008).Article 
PubMed 

Google Scholar  More