Philippa Kaur

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

Effects of plant genetic diversity on multiple trophic groupsWe found that plant genetic diversity (i.e. diversification of cropping or plant cultivation systems; see Methods and Supplementary Table 15) decreased the overall performance of plant antagonists (effect size = −0.539, t = −2.070, P = 0.039) and several of its components (i.e., herbivores (effect size = −0.606, t = −4.127, P  More

Kimmel, K., Kull, A., Salm, J. & Mander, Ü. The status, conservation and sustainable use of Estonian wetlands. Wetl. Ecol. Manag. 18, 375–395. https://doi.org/10.1007/s11273-008-9129-z (2008).Article 

Google Scholar 
Engle, V. Estimating the provision of ecosystem services by Gulf of Mexico coastal wetlands. Wetlands 31, 179–193. https://doi.org/10.1007/s13157-010-0132-9 (2011).Article 

Google Scholar 
Ward, R., Teasdale, P., Burnside, N., Joyce, C. & Sepp, K. Recent rates of sedimentation on irregularly flooded Boreal Baltic coastal wetlands: Responses to recent changes in sea level. Geomorphology 217, 61–72. https://doi.org/10.1016/j.geomorph.2014.03.045 (2014).Article 

Google Scholar 
Villoslada Peciña, M. et al. Country-scale mapping of ecosystem services provided by semi-natural grasslands. Sci. Total Environ. 661, 212–225. https://doi.org/10.1016/j.scitotenv.2019.01.174 (2019).Article 
CAS 
PubMed 

Google Scholar 
Lima, M., Ward, R. & Joyce, C. Environmental drivers of sediment carbon storage in temperate seagrass meadows. Hydrobiologia 847, 1773–1792. https://doi.org/10.1007/s10750-019-04153-5 (2019).Article 
CAS 

Google Scholar 
Ward, R. Sedimentary response of Arctic coastal wetlands to sea level rise. Geomorphology 370, 107400. https://doi.org/10.1016/j.geomorph.2020.107400 (2020).Article 

Google Scholar 
Akumu, C., Pathirana, S., Baban, S. & Bucher, D. Examining the potential impacts of sea level rise on coastal wetlands in north-eastern NSW, Australia. J. Coast. Conserv. 15, 15–22. https://doi.org/10.1007/s11852-010-0114-3 (2010).Article 

Google Scholar 
Ward, R. Carbon sequestration and storage in Norwegian Arctic coastal wetlands: Impacts of climate change. Sci. Total Environ. 748, 141343. https://doi.org/10.1016/j.scitotenv.2020.141343 (2020).Article 
CAS 
PubMed 

Google Scholar 
Hossain, M., Hein, L., Rip, F. & Dearing, J. Integrating ecosystem services and climate change responses in coastal wetlands development plans for Bangladesh. Mitig. Adapt. Strateg. Glob. Chang. 20, 241–261. https://doi.org/10.1007/s11027-013-9489-4 (2015).Article 

Google Scholar 
Ward, R., Friess, D., Day, R. & Mackenzie, R. Impacts of climate change on global mangrove ecosystems: A regional comparison. Ecosyst. Health Sustain. 4, 1–25 (2016).
Google Scholar 
Graham, L. P. et al. Climate change. In The Baltic Sea Area Draft HELCOM Thematic Assessment. (Helsinki Commission, Baltic Marine Environmental Protection Commission, 2007).BACC. Assessment of Climate Change for the Baltic Sea Basin. (Springer Science & Business Media, 2008).
Google Scholar 
Rivis, R. et al. Trends in the development of Estonian coastal land cover and landscapes caused by natural changes and human impact. J. Coast. Conserv. 20, 199–209. https://doi.org/10.1007/s11852-016-0430-3 (2016).Article 

Google Scholar 
Cubasch, U. et al. Projections of future climate change. in IPCC Climate Change 2001: The Scientific Basis Contribution of Working Group I to the Third Assessment Report of the Intergovernmental Panel on Climate Change. (Cambridge University Press, 2001).Mafi-Gholami, D., Zenner, E., Jaafari, A. & Ward, R. Modeling multi-decadal mangrove leaf area index in response to drought along the semi-arid southern coasts of Iran. Sci. Total Environ. 656, 1326–1336. https://doi.org/10.1016/j.scitotenv.2018.11.462 (2019).Article 
CAS 
PubMed 

Google Scholar 
IPCC. Global Warming of 1.5 ºC. Ipcc.ch. https://www.ipcc.ch/sr15/ (2008).Omstedt, A., Pettersen, C., Rodhe, J. & Winsor, P. Baltic Sea climate: 200 yr of data on air temperature, sea level variation, ice cover, and atmospheric circulation. Clim. Res. 25, 205–216 (2004).Article 

Google Scholar 
Räisänen, J. Future climate change in the Baltic Sea Region and environmental impacts. Oxf. Res. Encycl. Clim. Sci. https://doi.org/10.1093/acrefore/9780190228620.013.634 (2017).Article 

Google Scholar 
Dippner, J. W. et al. Climate-related marine ecosystem change. in Team, B. A. Assessment of Climate Change for the Baltic Sea Basin. (SSBM, 2008).Ward, R., Burnside, N., Joyce, C., Sepp, K. & Teasdale, P. Improved modelling of the impacts of sea level rise on coastal wetland plant communities. Hydrobiologia 774, 203–216. https://doi.org/10.1007/s10750-015-2374-2 (2016).Article 

Google Scholar 
Vuorinen, I. Proportion of copepod biomass declines with decreasing salinity in the Baltic Sea. ICES Mar. Sci. 55, 767–774. https://doi.org/10.1006/jmsc.1998.0398 (1998).Article 

Google Scholar 
Berg, M., Joyce, C. & Burnside, N. Differential responses of abandoned wet grassland plant communities to reinstated cutting management. Hydrobiologia 692, 83–97. https://doi.org/10.1007/s10750-011-0826-x (2011).Article 

Google Scholar 
Short, F., Kosten, S., Morgan, P., Malone, S. & Moore, G. Impacts of climate change on submerged and emergent wetland plants. Aquat. Bot. 135, 3–17. https://doi.org/10.1016/j.aquabot.2016.06.006 (2016).Article 

Google Scholar 
Ward, R., Burnside, N., Joyce, C. & Sepp, K. Importance of microtopography in determining plant community distribution in Baltic coastal wetlands. J. Coast. Res. 321, 1062–1070. https://doi.org/10.2112/JCOASTRES-D-15-00065.1 (2016).Article 

Google Scholar 
Burnside, N., Joyce, C., Puurmann, E. & Scott, D. Use of vegetation classification and plant indicators to assess grazing abandonment in Estonian coastal wetlands. J. Veg. Sci. 18, 645–654. https://doi.org/10.1111/j.1654-1103.2007.tb02578.x (2007).Article 

Google Scholar 
Ward, R., Burnside, N., Joyce, C. & Sepp, K. The use of medium point density LiDAR elevation data to determine plant community types in Baltic coastal wetlands. Ecol. Indic. 33, 96–104. https://doi.org/10.1016/j.ecolind.2012.08.016 (2013).Article 

Google Scholar 
Goud, E., Watt, C. & Moore, T. Plant community composition along a peatland margin follows alternate successional pathways after hydrologic disturbance. Acta Oecol. 91, 65–72. https://doi.org/10.1016/j.actao.2018.06.006 (2018).Article 

Google Scholar 
Moreno, J., Terrones, A., Juan, A. & Alonso, M. Halophytic plant community patterns in Mediterranean saltmarshes: Shedding light on the connection between abiotic factors and the distribution of halophytes. Plant Soil 430, 185–204. https://doi.org/10.1007/s11104-018-3671-0 (2018).Article 
CAS 

Google Scholar 
Sharpe, P. & Baldwin, A. Tidal marsh plant community response to sea-level rise: A mesocosm study. Aquat. Bot. 101, 34–40. https://doi.org/10.1016/j.aquabot.2012.03.015 (2012).Article 

Google Scholar 
Lindig-Cisneros, R. & Zedler, J. Phalaris arundinacea seedling establishment: Effects of canopy complexity in fen, mesocosm, and restoration experiments. J. Bot. 80, 617–624. https://doi.org/10.1139/b02-042 (2002).Article 

Google Scholar 
Ahn, C. & Mitsch, W. Scaling considerations of mesocosm wetlands in simulating large created freshwater marshes. Ecol. Eng. 18, 327–342. https://doi.org/10.1016/S0925-8574(01)00092-1 (2002).Article 

Google Scholar 
Brotherton, S. & Joyce, C. Extreme climate events and wet grasslands: Plant traits for ecological resilience. Hydrobiologia 750, 229–243. https://doi.org/10.1007/s10750-014-2129-5 (2015).Article 

Google Scholar 
Stewart, R. I. et al. Mesocosm experiments as a tool for ecological climate-change research. Adv. Ecol. Res. AP. 48, 71–181 (2013).Article 

Google Scholar 
Kont, A., Ratas, U. & Puurmann, E. Sea-level rise impact on coastal areas of Estonia. Clim. Change 36, 175–184. https://doi.org/10.1023/A:1005352715752 (1997).Article 

Google Scholar 
Short, F. & Neckles, H. The effects of global climate change on seagrasses. Aquat. Bot. 63, 169–196. https://doi.org/10.1016/S0304-3770(98)00117-X (1999).Article 

Google Scholar 
Engels, J., Rink, F. & Jensen, K. Stress tolerance and biotic interactions determine plant zonation patterns in estuarine marshes during seedling emergence and early establishment. J. Ecol. 99, 277–287. https://doi.org/10.1111/j.1365-2745.2010.01745.x (2010).Article 

Google Scholar 
Rayner, D. et al. Intertidal wetland vegetation dynamics under rising sea levels. Sci. Total Environ. 766, 144237. https://doi.org/10.1016/j.scitotenv.2020.144237 (2021).Article 
CAS 
PubMed 

Google Scholar 
Toogood, S. & Joyce, C. Effects of raised water levels on wet grassland plant communities. Appl. Veg. Sci. 12, 283–294. https://doi.org/10.1111/j.1654-109X.2009.01028.x (2009).Article 

Google Scholar 
Humphreys, A., Gorsky, A., Bilkovic, D. & Chambers, R. Changes in plant communities of low-salinity tidal marshes in response to sea-level rise. Ecosphere https://doi.org/10.1002/ecs2.3630 (2021).Article 

Google Scholar 
Jarvis, J. C., McKenna, S. A. & Rasheed, M. A. Seagrass seed bank spatial structure and function following a large-scale decline. Mar. Ecol. Prog. Ser. 665, 75–87. https://doi.org/10.3354/meps13668 (2021).Article 

Google Scholar 
Elsey-Quirk, T. & Leck, M. Patterns of seed bank and vegetation diversity along a tidal freshwater river. Am. J. Bot. 102, 1996–2012. https://doi.org/10.3732/ajb.1500314 (2015).Article 
PubMed 

Google Scholar 
Jutila, H. Germination in Baltic coastal wetland meadows: Similarities and differences between vegetation and seed bank. Plant Ecol. 166, 275–293 (2003).Article 

Google Scholar 
Ellenberg, H. Zeigerwerte der Gefässpflanzen Mitteleuropas. 42–111. (Scr. Geobot., 1979).Joshi, R. et al. Salt adaptation mechanisms of halophytes: Improvement of salt tolerance in crop plants. in Elucidation of Abiotic Stress Signaling in Plants. (Springer, 2015).Tessier, M., Gloaguen, J. & Lefeuvre, J. Factors affecting the population dynamics of Suaeda maritima at initial stages of development. Plant Ecol. 147, 193–203 (2000).Article 

Google Scholar 
Hanslin, H. & Eggen, T. Salinity tolerance during germination of seashore halophytes and salt-tolerant grass cultivars. Seed Sci. Res. 15, 43–50. https://doi.org/10.1079/SSR2004196 (2005).Article 

Google Scholar 
Köster, T. et al. The management of the coastal grasslands of Estonia. WIT Trans. Ecol. Environ. https://doi.org/10.2495/CENV040051 (2004).Article 

Google Scholar 
Spencer, T. et al. Global coastal wetland change under sea-level rise and related stresses: The DIVA wetland change model. Glob. Planet. Change 139, 15–30. https://doi.org/10.1016/j.gloplacha.2015.12.018 (2016).Article 

Google Scholar 
Marani, M., D’Alpaos, A., Lanzoni, S., Carniello, L. & Rinaldo, A. Biologically-controlled multiple equilibria of tidal landforms and the fate of the Venice lagoon. Geophys. Res. Lett. https://doi.org/10.1029/2007GL030178 (2007).Article 

Google Scholar 
Petersen, K., Frank, H., Paytan, A. & Bar-Zeev, E. Impacts of seawater desalination on coastal environments. Sustain. Desalin. Handb. https://doi.org/10.1016/B978-0-12-809240-8.00011-3 (2018).Article 

Google Scholar 
Rannap, R. et al. Coastal meadow management for threatened waders has a strong supporting impact on meadow plants and amphibians. J. Nat. Conserv. 35, 77–91. https://doi.org/10.1016/j.jnc.2016.12.004 (2017).Article 

Google Scholar 
Krauss, K. et al. How mangrove forests adjust to rising sea level. New Phytol. 202, 19–34. https://doi.org/10.1111/nph.12605 (2014).Article 
PubMed 

Google Scholar 
Kirwan, M. et al. Limits on the adaptability of coastal marshes to rising sea level. Geophys. Res. Lett. https://doi.org/10.1029/2010GL045489 (2010).Article 

Google Scholar 
Burnside, N., Joyce, C., Berg, M. & Puurman, E. The relationship between microtopography and vegetation in Estonian coastal wetlands: Implications for climate change. Publ. Inst. Geogr. Univ. Tartu. 106, 19–23 (2008).
Google Scholar 
Hulisz, P., Piernik, A., Mantilla-Contreras, J. & Elvisto, T. Main driving factors for seacoast vegetation in the southern and eastern Baltic. Wetlands 36, 909–919. https://doi.org/10.1007/s13157-016-0803-2 (2016).Article 

Google Scholar 
Gough, L. & Grace, J. Effects of flooding, salinity and herbivory on coastal plant communities, Louisiana, United States. Oecologia 117, 527–535. https://doi.org/10.1007/s004420050689 (1998).Article 
PubMed 

Google Scholar 
Hannerz, F. & Destouni, G. Spatial characterization of the Baltic sea drainage basin and its unmonitored catchments. Ambio 35, 214–219. https://doi.org/10.1579/05-A-022R.1 (2006).Article 
PubMed 

Google Scholar 
Kont, A., Jaagus, J. & Aunap, R. Climate change scenarios and the effect of sea-level rise for Estonia. Glob. Planet. Change 36, 1–15. https://doi.org/10.1016/S0921-8181(02)00149-2 (2003).Article 

Google Scholar 
von Storch, H. & Omstedt, A. Introduction and summary. in Team, B. A. Assessment of Climate Change for the Baltic Sea Basin. (SSBM, 2008).Stigebrandt, A. Physical oceanography of the Baltic Sea. in A Systems Analysis of the Baltic Sea. 19–74 (Springer, 2001).Ingerpuu, N. & Sarv, M. Effect of grazing on plant diversity of coastal meadows in Estonia. Ann. Bot. Fenn. 52, 84–92. https://doi.org/10.5735/085.052.0210 (2015).Article 

Google Scholar 
Moinardeau, C., Mesléard, F., Ramone, H. & Dutoit, T. Short-term effects on diversity and biomass on grasslands from artificial dykes under grazing and mowing treatments. Environ. Conserv. 46, 132–139. https://doi.org/10.1017/S0376892918000346 (2019).Article 

Google Scholar 
Tardella, F. M., Bricca, A., Goia, I. G. & Catorci, A. How mowing restores montane Mediterranean grasslands following cessation of traditional livestock grazing. Agric. Ecosyst. Environ. 295, 1158. https://doi.org/10.1016/j.agee.2020.106880 (2020).Article 

Google Scholar 
Lindborg, R. & Eriksson, O. Historical landscape connectivity affects present plant species diversity. Ecology 85, 1840–1845. https://doi.org/10.1890/04-0367 (2004).Article 

Google Scholar 
Villoslada Peciña, M., Bergamo, T., Ward, R., Joyce, C. & Sepp, K. A novel UAV-based approach for biomass prediction and grassland structure assessment in coastal meadows. Ecol. Indic. 122, 107227. https://doi.org/10.1016/j.ecolind.2020.107227 (2021).Article 

Google Scholar 
Villoslada, M. et al. Fine scale plant community assessment in coastal meadows using UAV based multispectral data. Ecol. Indic. 111, 105979. https://doi.org/10.1016/j.ecolind.2019.105979 (2020).Article 

Google Scholar 
Araya, Y., Gowing, D. & Dise, N. A controlled water-table depth system to study the influence of fine-scale differences in water regime for plant growth. Aquat. Bot. 92, 70–74. https://doi.org/10.1016/j.aquabot.2009.10.004 (2010).Article 

Google Scholar 
Koch, E. et al. Non-linearity in ecosystem services: Temporal and spatial variability in coastal protection. Front. Ecol. Environ. 7, 29–37. https://doi.org/10.1890/080126 (2009).Article 

Google Scholar 
Church, J. & White, N. Sea-level rise from the late 19th to the early 21st century. Surv. Geophys. 32, 585–602. https://doi.org/10.1007/s10712-011-9119-1 (2011).Article 

Google Scholar 
Goodwillie, C., McCoy, M. & Peralta, A. Long-term nutrient enrichment, mowing, and ditch drainage interact in the dynamics of a wetland plant community. Ecosphere. https://doi.org/10.1002/ecs2.3252 (2020).Article 

Google Scholar 
Kindt, R. & Coe, R. Tree diversity analysis. A manual and software for common statistical methods for ecological and biodiversity studies. in World Agroforestry | Transforming Lives and Landscapes with Trees. http://www.worldagroforestry.org/output/tree-diversity-analysis (2005).Oksanen, J. et al. CRAN—Package Vegan. Cran.r-project.org. https://CRAN.R-project.org/package=vegan. (2022).Wickham, H. Create Elegant Data Visualisations Using the Grammar of Graphics. Ggplot2.tidyverse.org. https://ggplot2.tidyverse.org (2016).Avolio, M. et al. A comprehensive approach to analyzing community dynamics using rank abundance curves. Ecosphere. https://doi.org/10.1002/ecs2.2881 (2019).Article 

Google Scholar 
Curtis, J. & McIntosh, R. The interrelations of certain analytic and synthetic phytosociological characters. Ecology 31, 434–455. https://doi.org/10.2307/1931497 (1950).Article 

Google Scholar 
Porto, A. B., do Prado, M. A., Rodrigues, L. D. S. & Overbeck, G. E. Restoration of subtropical grasslands degraded by non-native pine plantations: Effects of litter removal and hay transfer. Restor. Ecol. https://doi.org/10.1111/rec.13773 (2022).Article 

Google Scholar 
Cáceres, M. D. & Legendre, P. Associations between species and groups of sites: Indices and statistical inference. Ecol. 90, 3566–3574. https://doi.org/10.1890/08-1823.1 (2009).Article 

Google Scholar 
Wickham, H., François, R., Henry, L. & Müller, K. dplyr: A Grammar of Data Manipulation. https://dplyr.tidyverse.org; https://github.com/tidyverse/dplyr (2022). More

Genetic variationThe best fitting substitution models for COI codons 1–3 were identified as TN + F + G4, F81 + F + I, and TN + F, respectively. The maximum likelihood (ML) and Bayesian inference (BI) trees showed similar topologies of the main nodes, although the BI tree displayed greater resolution of the ingroup branches (Fig. 1). Furthermore, the BI tree revealed three monophyletic clades, while two of those clades were merged in the ML tree. Two of the three molecular species delimitation methods (ASAP and TCS) recovered three groups in the BI tree as distinct taxa (Fig. 1), corresponding to the three previously described ESUs27,28. The third method (bPTP) recovered between 8 and 43 groups (mean = 28.03) suggesting that there is evidence of additional genetic differentiation within the three groups identified by ASAP and TCS. The outputs of the three methods are provided in the Supplementary information. The molecular diagnosis uncovered several fixed nucleotide differences COI characters for each taxon (Table 1: “W. carteri” I = 10; “W. carteri” II = 3; “W. carteri” III = 5). There were also 13 fixed nucleotide differences in W. carteri for the 16S gene. The remaining two taxa had no fixed nucleotide differences for the 16S gene.Figure 1Phylogenetic trees obtained by maximum likelihood (left) and Bayesian inference (right) analysis of “Westralunio carteri” mtDNA COI sequences, including support values for the major genetic clades [ultrafast bootstrap values (left) and Bayesian posterior probabilities (right)]. Colour coded bars show support for the three major clades by the species delimitation methods (ASAP = dark shade; TCS = lighter shade). Green = WcI = “W. carteri” I; blue = WcIII = “W. carteri” III; red = WcII = “W. carteri” II. Results of bPTP analysis not shown (see supplementary data). Haplotype names correspond to Benson et al.28. Outgroup taxa are Velesunio ambiguus (Philippi, 1847) (Hyriidae: Velesunioninae) and Cucumerunio novaehollandiae (Gray, 1834) (Hyriidae: Hyriinae: Hyridellini).Full size imageTable 1 Molecular diagnoses of “Westralunio carteri” Evolutionarily Significant Units (ESUs) from southwestern Australia (after Bolotov et al.122 with reanalysis of data from Klunzinger et al.27 and Benson et al.28).Full size tableVariation in shell morphologyBased on results from analyses of variances (ANOVAs), shells of “W. carteri” I were significantly larger (for size metrics total length (TL), maximum height (MH), beak height (BH) and beak length (BL)) and more elongated (i.e., had a lower maximum height index (MHI)) than shells of “W. carteri” II and “W. carteri” II + III combined (Table 2). However, there was no difference in size or shape metrics between “W. carteri” I and “W. carteri” III (Table 2). The lack of significant differences in beak height index (BHI) and beak length index (BLI) among any of the taxa (Table 2) indicates that wing and anterior shell development was not discernibly different between any of the ESUs.Table 2 Shell size metrics [mm], shape indices [%] and scores for the first two principal components (PC) obtained by Principal Component Analysis of shape indices and 18 Fourier coefficients generated by Fourier Shape Analysis for each “Westralunio carteri” species and subspecies-level Evolutionarily Significant Units (ESUs): n, number of specimens measured; minimum (min) to maximum (max) and mean (± standard error (SE)).Full size tableThis pattern was partly confirmed in the principal component analysis (PCA) of these three shell shape indices, where PC1, largely explained by variation in BLI (Fig. 2A), did not differ between the two species (i.e., “W. carteri” I vs. “W. carteri” II + III) or among the three taxa (Table 2). The PC2, largely explained by variation in MHI and BHI (Fig. 2A), differed significantly between “W. carteri” I and “W. carteri” II (Table 2). Accordingly, 70% (70% jack-knifed) of specimens were assigned to the correct species in the corresponding discriminant analysis (DA), whilst this was true for only 55% (54%) at the MOTU-level.Figure 2Scatterplots of the first two PC axes obtained by PCA on (A) calculated shape indices based on shell measurements, and (B) 18 Fourier coefficients for “Westralunio carteri” I, “W. carteri” II and “W. carteri” III. 95% Confidence Intervals are displayed at the species level, i.e., for “W. carteri” I (full line) and “W. carteri” II + III (dashed line). Extreme shell outlines in (B) are depicted to visualise trends in sagittal shell shape, along PC axes.Full size imageThe difference in shell elongation between “W. carteri” I and “W. carteri” II was confirmed by Fourier shape analysis. As visualised by synthetic outlines in Fig. 2B, shell elongation is expressed along the PC1 (explaining 15% of total variation in Fourier coefficients). The PC1 as well as PC2 scores differed significantly between the two species (i.e., “W. carteri” I vs. “W. carteri” II + III) as well as between “W. carteri” I and “W. carteri” II, respectively (Table 2). Combined with synthetic outlines, this indicated a tendency towards a more elongated, somewhat wedge-shaped shell in “W. carteri” I, whilst “W. carteri” II shells tended to be relatively high with a stout anterior margin (Fig. 2B). An analysis of similarities (ANOSIM) analysis on all Fourier coefficients revealed no significant difference between the two species (i.e., “W. carteri” I vs. “W. carteri” II + III; ANOSIM: R = − 0.018, p = 0.097), but did indicate a significant difference between the three ESUs (ANOSIM: R = 0.0625, p = 0.0051). Specifically, “W. carteri” I differed significantly from “W. carteri” II (Bonferroni-corrected p = 0.0009). Only 66% and 65% (62% and 62% jack-knifed) of specimens were assigned to the correct species and taxon in DAs on that dataset, respectively.Taxonomic accountsClass: Bivalvia Linnaeus, 175831.Subclass: Autobranchia Grobben, 189432.Infraclass: Heteroconchia Gray, 185433.Cohort: Palaeoheterodonta Newell, 196534.Order: Unionida Gray, 185433 in Bouchet & Rocroi, 201035.Superfamily: Unionoidea Rafinesque, 182036.Family: Hyriidae Parodiz & Bonetto 196337.Genus: Westralunio Iredale, 19349.Type species: Westralunio ambiguus carteri Iredale, 19349 (by original designation).Redescription: Westralunio carteri (Iredale, 1934)SynonymyUnio australis Lamarck38: Menke39, p. 38, specimen 219. (Non Unio australis Lamarck, 181938).Unio moretonicus Reeve40: Smith41, p. 3, pl. iv, Fig. 2. (misidentified reference to Unio moretonicus Reeve, 186540).Hyridella australis (Lam.38): Cotton & Gabriel42 (in part), p. 156. (misidentified reference to Unio australis Lamarck, 181938).Hyridella ambigua (Philippi26): Cotton & Gabriel42 (in part), p. 157. (misidentified reference to Unio ambiguus Philippi, 184726).Westralunio ambiguus carteri: Iredale, 19349, p. 62.Westralunio ambiguus (Philippi26): Iredale9, p. 62, pl. iii, Fig. 8, pl. iv, Fig. 8. (Non Unio ambiguus Phil. 184726), Iredale43, p. 190.Centralhyria angasi subjecta Iredale, 19349, p. 67 (in part), Iredale43, p. 190.Westralunio carteri Iredale9: McMichael & Hiscock10pl. viii, Figs. 1, 2, 3, 4, 5, 6 and 7, pl. xvii, Figs. 4, 5.Type materialLectotype: AMS C.61724 (Fig. 3A) Westralunio ambiguus carteri Iredale, 19349.Figure 3(A) Westralunio ambiguus carteri Iredale, 1934, Lectotype: Victoria Reservoir, Darling Range, 12 mi E of Perth, AMS C.061724. Detail of fusion in anterior muscle scars from either valve represented by dashed lines and black polygons. Bottom image showing detail of hinge teeth. Photos provided with permission by Dr Mandy Reid, AMS. (B) Valves and detail of sculptured umbo of a juvenile W. carteri from Yule Brook, Western Australia, UMZC 2013.2.9. Photo by Dr Michael W. Klunzinger. (C) Glochidia of W. carteri from Canning River, Western Australia. Photo by Dr Michael W. Klunzinger.Full size imageParalectotypes: AMS C.170635 Westralunio ambiguus carteri Iredale, 19349 (n = 4).Type locality: Victoria Reservoir, Darling Range, 12 miles east of Perth, Western Australia (Fig. 4A).Figure 4(A) Victoria Reservoir, Canning River, near Perth, Western Australia, type locality for W. carteri. Photo by Corey Whisson. (B) Goodga River, Western Australia, type locality for W. inbisi inbisi, at vertical slot fishway where holotype of W. inbisi inbisi was collected from. Photo provided with permission by Dr Stephen J. Beatty. (C) Margaret River, Western Australia, type locality for W. inbisi meridiemus, at Canebreak Pool. Photo by Dr Michael W. Klunzinger.Full size imageLectotype: BMNH 1840–10-21–29 Centralhyria angasi subjecta Iredale (selected by McMichael & Hiscock10).Type locality: Avon River, Western Australia.Material examined for redescription: For W. carteri (= “W. carteri” I), molecular data examined included 52 and 61 individual COI mtDNA and 16S rDNA sequences, respectively, for species delimitation. Additionally, Fourier shell shape outline analysis and traditional shell morphometric measurements were examined from 238 and 290 individuals, respectively. Complete details on all specimens examined are provided in Supplementary Table S1.ZooBank registration: urn:lsid:zoobank.org:act:6B740F4D-40C3-4D6A-8938-B0FD7FD1F6D7.Etymology: The species name carteri is most likely named after the surname of the collector who provided original type specimens to the Australian Museum, although Iredale9 did not specify this as the case. We have applied ICZN Articles 46.1 and 47.144, designating W. carteri as the nominotypical species.Revised diagnosis: Specimens of W. carteri are distinguished from other Australian Westralunio taxa by having shell series that are significantly larger and more elongated than Westralunio inbisi inbisi subsp. nov., but not different from Westralunio inbisi meridiemus subsp. nov. The species has 10 diagnostic nucleotides at COI (57 G, 117 T, 210 G, 249 T, 255 C, 345 G, 423 T, 447 T, 465 A, 499 T) and 13 at 16S (137 T, 155 C, 228 C, 229 T, 260 G, 290 A, 305 G, 307 T, 310 A, 311 C, 321 T, 330 A, 460 A), which differentiate it from its sister taxa, W. inbisi inbisi and W. inbisi meridiemus (each described below) using ASAP and TCS species delimitation models.RedescriptionThis species is of the ESU “W. carteri” I27,28.Shell morphology: Shells of relatively small to medium size, generally less than 70 mm in length, but to a maximum length of approximately 100 mm10,45, MHI 46–89%; anterior portion of shell with moderate development, BLI 22–49%; larger shells with abraded umbos scarcely winged; wing development variable, generally decreasing with size, BHI 76–104% (Table 2). Shell outline oblong-ovate to rounded; posterior end obliquely to squarely truncate, anterior end round; ventral edge slightly curved, nearly straight in larger specimens; hinge line curved, hinge strong. Umbos usually abraded in specimens  > 20 mm in length; unabraded umbos with distinctive v- or w-shaped plicated sculpturing (Fig. 3B and Zieritz et al.46). Shell substance typically thick; shells of medium width with pronounced posterior ridge; periostracum smooth, dark brown to reddish, with fine growth lines. Pallial line less developed in smaller specimens and prominent only in large specimens (e.g.,  > 60 mm TL). Lateral teeth longer and blade-like, slightly serrated to smooth and singular in left valve, fitting into deep groove in right valve; pseudocardinal tooth in right valve coarsely serrated, thick, and erect, fitting into deeply grooved socket in left valve. Anterior muscle scars well impressed and anchored deeply in larger specimens; anterior retractor pedis and protractor pedis scars both small and fused with adductor muscle scar; posterior muscle scars lightly impressed; dorsal muscle scars usually with two or three deep pits anchored to internal umbo region.Anatomy: Supra-anal opening absent, siphons of moderate size, not prominent but protrude beyond shell margin in actively filtering live specimens, pigmented dark brown with mottled lighter brown to orange splotches; inhalant siphon aperture about 1.5 times size of exhalant and bearing 2–4 rows of internal papillae (Fig. 5A); ctenidial diaphragm relatively long and perforated. Outer lamellae of outer ctenidia completely fused to mantle, inner lamellae of inner ctenidia fused to visceral mass then united to form diaphragm; palps relatively small, usually semilunar in shape; marsupium well developed as a distinctive swollen interlamellar space in the middle third of the inner ctenidium of females. Outer ctenidia in both sexes thin, with numerous, short intrafilamentary junctions and few, irregular interlamellar junctions; in females similar, but marsupium has numerous, tightly packed, well-developed interlamellar junctions. Thus, brooding in females is endobranchous.Figure 5Live specimens of actively filtering freshwater mussels in the burrowed position. (A) Westralunio carteri (Iredale, 1934), Canning River at Kelmscott, Western Australia, inhalant siphon with 2–4 rows of papillae oriented toward substrate. Photo by Dr Michael W. Klunzinger. (B) Westralunio inbisi meridiemus subsp. nov., Canebreak Pool, Margaret River, Western Australia; inhalant siphon edges lined with protruding papillae facing towards water surface, away from substrate. Photo by Dr Michael W. Klunzinger.Full size imageLife history: Sexes are separate in W. carteri, and hermaphroditism appears to be rare47,48,49. Males and females both produce gametes year-round but brooding of glochidia appears to be seasonal and ‘tachyticitc’ (i.e., as defined by Bauer & Wächtler19, fertilisation and embryonic development occurring in late winter/early spring and glochidia release in early summer)50. Glochidia are released within vitelline membranes, embedded in mucus which extrude from exhalant siphons of females (i.e., ‘amorphous mucus conglutinates’) during spring/summer. Glochidia attach to host fishes and live parasitically on fins, gills or body surfaces for 3–4 weeks while undergoing metamorphosis to the juvenile stage. Host fishes which have been shown to support glochidia metamorphosis to the juvenile stage in the laboratory include Afurcagobius suppositus (Sauvage, 188051), Gambusia holbrooki (Girard, 185952), Nannoperca vitttata (Castelnau, 187353), Pseudogobius olorum (Sauvage, 188051) and Tandanus bostocki Whitley, 194454 but not Carassisus auratus Linnaeus, 175831 or Geophagus brasiliensis (Quoy & Gaimard, 1824 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

Henry, L.-A. & Roberts, J. M. Biodiversity and ecological composition of macrobenthos on cold-water coral mounds and adjacent off-mound habitat in the bathyal Porcupine Seabight, NE Atlantic. Deep Sea Res. I(54), 654–672 (2007).
Google Scholar 
Buhl-Mortensen, L. et al. First observations of the structure and megafaunal community of a large Lophelia reef on the Ghanaian shelf (the Gulf of Guinea). Deep Sea Res. II(137), 148–156 (2017).
Google Scholar 
Price, D. M. et al. Using 3D photogrammetry from ROV video to quantify cold-water coral reef structural complexity and investigate its influence on biodiversity and community assemblage. Coral Reefs 38, 1007–1021 (2019).
Google Scholar 
Roberts, J. M., Wheeler, A. J. & Freiwald, A. Reefs of the deep: the biology and geology of cold-water coral ecosystems. Science 312, 543–547 (2006).CAS 
PubMed 

Google Scholar 
Henry, L. A., Nizinski, M. S. & Ross, S. W. Occurrence and biogeography of hydroids (Cnidaria: Hydrozoa) from deep-water coral habitats off the southeastern United States. Deep. Res. I(55), 788–800 (2008).
Google Scholar 
Henry, L.-A. & Roberts, J. M. Global Biodiversity in Cold-Water Coral Reef Ecosystems. In Marine Animal Forests (eds Rossi, S. et al.) 1–21 (Springer, 2016). https://doi.org/10.1007/978-3-319-17001-5_6-1.Chapter 

Google Scholar 
De Mol, B. et al. Large deep-water coral banks in the Porcupine Basin, southwest of Ireland. Mar. Geol. 188, 193–231 (2002).
Google Scholar 
Dorschel, B., Hebbeln, D., Rüggeberg, A., Dullo, W. C. & Freiwald, A. Growth and erosion of a cold-water coral covered carbonate mound in the Northeast Atlantic during the Late Pleistocene and Holocene. Earth Planet. Sci. Lett. 233, 33–44 (2005).CAS 

Google Scholar 
Hebbeln, D., Van Rooij, D. & Wienberg, C. Good neighbours shaped by vigorous currents: Cold-water coral mounds and contourites in the North Atlantic. Mar. Geol. 378, 171–185 (2016).
Google Scholar 
Wheeler, A. J. et al. Morphology and environment of cold-water coral carbonate mounds on the NW European margin. Int. J. Earth Sci. 96, 37–56 (2007).CAS 

Google Scholar 
Lo Iacono, C., Savini, A. & Basso, D. Cold-water carbonate bioconstructions. in Submarine Geomorphology, 425–455 (Springer, 2018).Hebbeln, D. Highly variable submarine landscapes in the Alborán sea created by cold-water corals. In Mediterranean Cold-Water Corals: Past, Present and Future (eds Orejas, C. & Jiménez, C.) 61–65 (Springer, 2019). https://doi.org/10.1007/978-3-319-91608-8_8.Chapter 

Google Scholar 
Addamo, A. M. et al. Merging scleractinian genera: The overwhelming genetic similarity between solitary Desmophyllum and colonial Lophelia. BMC Evol. Biol. 16, 1–17 (2016).
Google Scholar 
Wienberg, C. & Titschack, J. Framework-forming scleractinian cold-water corals through space and time: A late quaternary north atlantic perspective. in Marine Animal Forests 1–34 (Springer, 2017). https://doi.org/10.1007/978-3-319-17001-5_16-1Maier, C., Weinbauer, M. G. & Gattuso, J.-P. Fate of mediterranean scleractinian cold-water corals as a result of global climate change: A synthesis. In Mediterranean Cold-Water Corals: Past, Present and Future (eds Orejas, C. & Jiménez, C.) 517–529 (Springer, 2019). https://doi.org/10.1007/978-3-319-91608-8_44.Chapter 

Google Scholar 
Reynaud, S. & Ferrier-Pagès, C. Biology and ecophysiology of mediterranean cold-water corals. In Mediterranean Cold-Water Corals: Past, Present and Future (eds Orejas, C. & Jiménez, C.) 391–404 (Springer, 2019). https://doi.org/10.1007/978-3-319-91608-8_35.Chapter 

Google Scholar 
Hennige, S. J. et al. Using the Goldilocks principle to model coral ecosystem engineering. Proc. R. Soc. B Biol. Sci. 288, 20211260 (2021).CAS 

Google Scholar 
LoIacono, C. et al. The West Melilla cold water coral mounds, Eastern Alboran Sea: Morphological characterization and environmental context. Deep Sea Res. II(99), 316–326 (2014).
Google Scholar 
Mortensen, P. B., Hovland, T., Fosså, J. H. & Furevik, D. M. Distribution, abundance and size of Lophelia pertusa coral reefs in mid-Norway in relation to seabed characteristics. J. Mar. Biol. Assoc. 81, 581–597 (2001).
Google Scholar 
Mienis, F. et al. Hydrodynamic controls on cold-water coral growth and carbonate-mound development at the SW and SE Rockall Trough Margin, NE Atlantic. Ocean. Deep. Res. I(54), 1655–1674 (2007).
Google Scholar 
Davies, A. J. et al. Downwelling and deep-water bottom currents as food supply mechanisms to the cold-water coral Lophelia pertusa (Scleractinia) at the Mingulay Reef Complex. Limnol. Oceanogr. 54, 620–629 (2009).
Google Scholar 
Mohn, C. et al. Linking benthic hydrodynamics and cold-water coral occurrences: A high-resolution model study at three cold-water coral provinces in the NE Atlantic. Prog. Oceanogr. 122, 92–104 (2014).
Google Scholar 
Mienis, F. et al. Cold-water coral growth under extreme environmental conditions, the Cape Lookout area, NW Atlantic. Biogeosciences 11, 2543–2560 (2014).
Google Scholar 
Grasmueck, M. et al. Autonomous underwater vehicle (AUV) mapping reveals coral mound distribution, morphology, and oceanography in deep water of the Straits of Florida. Geophys. Res. Lett. 33, L23616 (2006).
Google Scholar 
Correa, T. B. S., Eberli, G. P., Grasmueck, M., Reed, J. K. & Correa, A. M. S. Genesis and morphology of cold-water coral ridges in a unidirectional current regime. Mar. Geol. 326–328, 14–27 (2012).
Google Scholar 
Lavaleye, M. et al. Cold-water corals on the tisler reef: Preliminary observations on the dynamic reef environment. Oceanography 22, 76–84 (2009).
Google Scholar 
Mortensen, P. B. et al. Seascape description of anunusual coral reef area off Vesteraålen, Northern Norway. in 4th International Symposium on deep-sea corals. (2008).Cathalot, C. et al. Cold-water coral reefs and adjacent sponge grounds: Hotspots of benthic respiration and organic carbon cycling in the deep sea. Front. Mar. Sci. 2, 37 (2015).
Google Scholar 
Buhl-Mortensen, P. & Sundahl, H. Environmental control of cold-water coral reef morphology. in 7th International Symposium on deep-sea corals. (2019).van der Kaaden, A.-S., van Oevelen, D., Rietkerk, M., Soetaert, K. & van de Koppel, J. Spatial self-organization as a new perspective on cold-water coral mound development. Front. Mar. Sci. 7, 631 (2020).
Google Scholar 
Buhl-Mortensen, L. et al. Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins. Mar. Ecol. 31, 21–50 (2010).
Google Scholar 
Jones, C. G., Lawton, J. H. & Shachak, M. Organisms as ecosystem engineers. Oikos 69, 373–386 (1994).
Google Scholar 
Mienis, F., Bouma, T., Witbaard, R., van Oevelen, D. & Duineveld, G. Experimental assessment of the effects of coldwater coral patches on water flow. Mar. Ecol. Prog. Ser. 609, 101–117 (2019).CAS 

Google Scholar 
van der Kaaden, A.-S. et al. Feedbacks between hydrodynamics and cold-water coral mound development. Deep Sea Res. I 178, 103641 (2021).
Google Scholar 
Mortensen, P. B., Hovland, M., Brattegard, T. & Farestveit, R. Deep water bioherms of the scleractinian coral Lophelia pertusa (L.) at 64° n on the norwegian shelf: Structure and associated megafauna. Sarsia 80, 145–158 (1995).
Google Scholar 
Corbera, G. et al. Ecological characterisation of a Mediterranean cold-water coral reef: Cabliers Coral Mound Province (Alboran Sea, western Mediterranean). Prog. Oceanogr. 175, 245–262 (2019).
Google Scholar 
Kano, A. et al. Age constraints on the origin and growth history of a deep-water coral mound in the northeast Atlantic drilled during Integrated Ocean Drilling Program Expedition 307. Geology 35, 1051–1054 (2007).CAS 

Google Scholar 
Douarin, M. et al. Growth of north-east Atlantic cold-water coral reefs and mounds during the Holocene: A high resolution U-series and 14C chronology. Earth Planet. Sci. Lett. 375, 176–187 (2013).CAS 

Google Scholar 
Orejas, C., Gori, A. & Gili, J. M. Growth rates of live Lophelia pertusa and Madrepora oculata from the Mediterranean Sea maintained in aquaria. Coral Reefs 27, 255–255 (2008).
Google Scholar 
Orejas, C. et al. Long-term growth rates of four Mediterranean cold-water coral species maintained in aquaria. Mar. Ecol. Prog. Ser. 429, 57–65 (2011).
Google Scholar 
Lartaud, F., Mouchi, V., Chapron, L., Meistertzheim, A.-L. & Le Bris, N. Growth Patterns of Mediterranean Calcifying Cold-Water Corals. in Mediterranean Cold-Water Corals: Past, Present and Future 405–422 (2019). https://doi.org/10.1007/978-3-319-91608-8_36.Büscher, J. V. et al. In situ growth and bioerosion rates of Lophelia pertusa in a Norwegian fjord and open shelf cold-water coral habitat. PeerJ 2019, 1–10 (2019).
Google Scholar 
Form, A. U. & Riebesell, U. Acclimation to ocean acidification during long-term CO2 exposure in the cold-water coral Lophelia pertusa. Glob. Chang. Biol. 18, 843–853 (2012).
Google Scholar 
Maier, C., Watremez, P., Taviani, M., Weinbauer, M. G. & Gattuso, J. P. Calcification rates and the effect of ocean acidification on Mediterranean cold-water corals. Proc. R. Soc. B Biol. Sci. 279, 1716–1723 (2012).CAS 

Google Scholar 
Lunden, J. J., McNicholl, C. G., Sears, C. R., Morrison, C. L. & Cordes, E. E. Acute survivorship of the deep-sea coral Lophelia pertusa from the Gulf of Mexico under acidification, warming, and deoxygenation. Front. Mar. Sci. 1, 78 (2014).
Google Scholar 
Gori, A., Reynaud, S., Orejas, C., Gili, J. M. & Ferrier-Pagès, C. Physiological performance of the cold-water coral Dendrophyllia cornigera reveals its preference for temperate environments. Coral Reefs 33, 665–674 (2014).
Google Scholar 
Huvenne, V. A. I. et al. Sediment dynamics and palaeo-environmental context at key stages in the Challenger cold-water coral mound formation: Clues from sediment deposits at the mound base. Deep. Res. I(56), 2263–2280 (2009).
Google Scholar 
Bartzke, G. et al. Investigating the prevailing hydrodynamics around a cold-water coral colony using a physical and a numerical approach. Front. Mar. Sci. 8, 3304 (2021).
Google Scholar 
Downs, C. A. et al. Cellular diagnostics and coral health: Declining coral health in the Florida Keys. Mar. Pollut. Bull. 51, 558–569 (2005).CAS 
PubMed 

Google Scholar 
Ayala, A., Muñoz, M. F. & Argüelles, S. Lipid peroxidation: Production, metabolism, and signaling mechanisms of malondialdehyde and 4-hydroxy-2-nonenal. Oxid. Med. Cell. Long. 2014, 1–10 (2014).CAS 

Google Scholar 
Oh, T. J., Kim, I. G., Park, S. Y., Kim, K. C. & Shim, H. W. NAD-dependent malate dehydrogenase protects against oxidative damage in Escherichia coli K-12 through the action of oxaloacetate. Environ. Toxicol. Pharmacol. 11, 9–14 (2002).CAS 
PubMed 

Google Scholar 
Dade, L., Hogg, A. & Boudreau, B. Physics of Flow Above the Sediment-Water Interface (Oxford University Press, 2001).
Google Scholar 
Gass, S. E. & Roberts, J. M. The occurrence of the cold-water coral Lophelia pertusa (Scleractinia) on oil and gas platforms in the North Sea: Colony growth, recruitment and environmental controls on distribution. Mar. Pollut. Bull. 52, 549–559 (2006).CAS 
PubMed 

Google Scholar 
Brooke, S. & Young, C. M. In situ measurement of survival and growth of Lophelia pertusa in the northern Gulf of Mexico. Mar. Ecol. Prog. Ser. 397, 153–161 (2009).
Google Scholar 
Lartaud, F. et al. A new approach for assessing cold-water coral growth in situ using fluorescent calcein staining. Aquat. Living Resour. 26, 187–196 (2013).
Google Scholar 
Sebens, K. P., Witting, J. & Helmuth, B. Effects of water flow and branch spacing on particle capture by the reef coral Madracis mirabilis (Duchassaing and Michelotti). J. Exp. Mar. Bio. Ecol. 211, 1–28 (1997).
Google Scholar 
Sebens, K. P., Grace, S. P., Helmuth, B., Maney, E. J. Jr. & Miles, J. S. Water flow and prey capture by three scleractinian corals, Madracis mirabilis, Montastrea cavernosa and Porites porites, in a field enclosure. Mar. Biol. 131, 347–360 (1998).
Google Scholar 
Purser, A., Larsson, A. I., Thomsen, L. & van Oevelen, D. The influence of flow velocity and food concentration on Lophelia pertusa (Scleractinia) zooplankton capture rates. J. Exp. Mar. Bio. Ecol. 395, 55–62 (2010).
Google Scholar 
Orejas, C. et al. The effect of flow speed and food size on the capture efficiency and feeding behaviour of the cold-water coral Lophelia pertusa. J. Exp. Mar. Bio. Ecol. 481, 34–40 (2016).
Google Scholar 
Duineveld, G. C. A. et al. Spatial and tidal variation in food supply to shallow cold-water coral reefs of the Mingulay Reef complex (Outer Hebrides, Scotland). Mar. Ecol. Prog. Ser. 444, 97–115 (2012).
Google Scholar 
De Clippele, L. H. et al. The effect of local hydrodynamics on the spatial extent and morphology of cold-water coral habitats at Tisler Reef, Norway. Coral Reefs 37, 253–266 (2018).PubMed 

Google Scholar 
Jokiel, P. L. Effects of water motion on reef corals. J. Exp. Mar. Biol. Ecol. 35, 87–97 (1978).
Google Scholar 
Shashar, N., Cohen, Y. & Loya, Y. Extreme diel fluctuations of oxygen in diffusive boundary layers surrounding stony corals. Biol. Bull. 185, 455–461 (1993).CAS 
PubMed 

Google Scholar 
Finelli, C. M., Helmuth, B. S. T., Pentcheff, N. D. & Wethey, D. S. Water flow influences oxygen transport and photosynthetic efficiency in corals. Coral Reefs 25, 47–57 (2006).
Google Scholar 
Atkinson, M. J. & Bilger, R. W. Effects of water velocity on phosphate uptake in coral reef-hat communities. Limnol. Oceanogr. 37, 273–279 (1992).CAS 

Google Scholar 
Mass, T., Genin, A., Shavit, U., Grinstein, M. & Tchernov, D. Flow enhances photosynthesis in marine benthic autotrophs by increasing the efflux of oxygen from the organism to the water. Proc. Natl. Acad. Sci. 107, 2527–2531 (2010).CAS 
PubMed 
PubMed Central 

Google Scholar 
Comeau, S., Edmunds, P. J., Lantz, C. A. & Carpenter, R. C. Water flow modulates the response of coral reef communities to ocean acidification. Sci. Rep. 4, 6681 (2014).CAS 
PubMed 
PubMed Central 

Google Scholar 
Larsson, A., Lundälv, T. & van Oevelen, D. Skeletal growth, respiration rate and fatty acid composition in the cold-water coral Lophelia pertusa under varying food conditions. Mar. Ecol. Prog. Ser. 483, 169–184 (2013).
Google Scholar 
Baussant, T., Nilsen, M., Ravagnan, E., Westerlund, S. & Ramanand, S. Physiological responses and lipid storage of the coral Lophelia pertusa at varying food density. J. Toxicol. Environ. Health. A 80, 266–284 (2017).CAS 
PubMed 

Google Scholar 
Bouma, T. J. et al. Spatial flow and sedimentation patterns within patches of epibenthic structures: Combining field, flume and modelling experiments. Cont. Shelf Res. 27, 1020–1045 (2007).
Google Scholar 
Brooke, S. D., Holmes, M. W. & Young, C. M. Sediment tolerance of two different morphotypes of the deep-sea coral Lophelia pertusa from the Gulf of Mexico. Mar. Ecol. Prog. Ser. 390, 137–144 (2009).
Google Scholar 
Bøe, R. et al. Giant sandwaves in the Hola glacial trough off Vesterålen, North Norway. Mar. Geol. 267, 36–54 (2009).
Google Scholar 
Huvenne, V. A. I. et al. The Magellan mound province in the Porcupine Basin. Int. J. Earth Sci. 96, 85–101 (2007).CAS 

Google Scholar 
De Haas, H. et al. Morphology and sedimentology of (clustered) cold-water coral mounds at the south Rockall Trough margins, NE Atlantic Ocean. Facies 55, 1–26 (2009).
Google Scholar 
Lim, A., Huvenne, V. A. I., Vertino, A., Spezzaferri, S. & Wheeler, A. J. New insights on coral mound development from groundtruthed high-resolution ROV-mounted multibeam imaging. Mar. Geol. 403, 225–237 (2018).
Google Scholar 
Olariaga, A., Gori, A., Orejas, C. & Gili, J. M. Development of an autonomous aquarium system for maintaining deep corals. Oceanography 22, 44–45 (2009).
Google Scholar 
Davies, A. J. et al. Short-term environmental variability in cold-water coral habitat at Viosca Knoll, Gulf of Mexico. Deep Sea Res. I(57), 199–212 (2010).
Google Scholar 
Mienis, F. et al. The influence of near-bed hydrodynamic conditions on cold-water corals in the Viosca Knoll area, Gulf of Mexico. Deep Sea Res. I(60), 32–45 (2012).
Google Scholar 
Flo, E., Garcés, E., Manzanera, M. & Camp, J. Coastal inshore waters in the NW Mediterranean: Physicochemical and biological characterization and management implications. Estuar. Coast. Shelf Sci. 93, 279–289 (2011).CAS 

Google Scholar 
Davies, P. S. Short-term growth measurements of corals using an accurate buoyant weighing technique. Mar. Biol. 101, 389–395 (1989).
Google Scholar 
R Core Team. R: A Language and Environment for Statistical Computing. (R Core Team, 2018).Bradford, M. M. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal. Biochem. 72, 248–254 (1976).CAS 
PubMed 

Google Scholar 
Thérond, P., Auger, J., Legrand, A. & Jouannet, P. α-tocopherol in human spermatozoa and seminal plasma: Relationships with motility, antioxidant enzymes and leukocytes. Mol. Hum. Reprod. 2, 739–744 (1996).PubMed 

Google Scholar 
Beers, R. F. & Sizer, I. W. A spectrophotometric method for measuring the breakdown of hydrogen peroxide by catalase. J. Biol. Chem. 195, 133–140 (1952).CAS 
PubMed 

Google Scholar 
Kalghatgi, S. et al. Bactericidal antibiotics induce mitochondrial dysfunction and oxidative damage in mammalian cells. Sci. Transl. Med. 5, 1–10 (2013).
Google Scholar  More

Becker, J., Manske, C. & Randl, S. Green chemistry and sustainability metrics in the pharmaceutical manufacturing sector. Curr. Opin. Green Sustain. Chem. https://doi.org/10.1016/j.cogsc.2021.100562 (2022).Article 

Google Scholar 
Rajasekharreddy, P., Rani, P. U. & Sreedhar, B. Qualitative assessment of silver and gold nanoparticle synthesis in various plants: A photobiological approach. J. Nanoparticle Res. 12, 25 (2010).Article 

Google Scholar 
Mahmoud, A. E. D. Eco-friendly reduction of graphene oxide via agricultural byproducts or aquatic macrophytes. Mater. Chem. Phys. 253, 123336 (2020).Article 
CAS 

Google Scholar 
Mahmoud, A. E. D., Stolle, A. & Stelter, M. Sustainable synthesis of high-surface-area graphite oxide via dry ball milling. ACS Sustain. Chem. Eng. 6, 25 (2018).Article 

Google Scholar 
Mellinas, C., Jiménez, A. & del Carmen Garrigós, M. Microwave-assisted green synthesis and antioxidant activity of selenium nanoparticles using theobroma cacao. l. bean shell extract. Molecules 24, 25 (2019).Article 

Google Scholar 
Ahmed, S., Saifullah, A. M., Swami, B. L. & Ikram, S. Green synthesis of silver nanoparticles using Azadirachta indica aqueous leaf extract. J. Radiat. Res. Appl. Sci. 9, 25 (2016).
Google Scholar 
Ebadi, M. et al. A bio-inspired strategy for the synthesis of zinc oxide nanoparticles (ZnO NPs) using the cell extract of cyanobacterium: Nostoc sp EA03: From biological function to toxicity evaluation. RSC Adv. 9, 25 (2019).Article 

Google Scholar 
Mahmoud, A. E. D. & Fawzy, M. Nanosensors and nanobiosensors for monitoring the environmental pollutants. Top. Min. Metallurg. Mater. Eng. https://doi.org/10.1007/978-3-030-68031-2_9 (2021).Article 

Google Scholar 
Mousavi, S. M. et al. Green synthesis of silver nanoparticles toward bio and medical applications: Review study. Artif. Cells Nanomed. Biotechnol. 46, 3. https://doi.org/10.1080/21691401.2018.1517769 (2018).Article 
CAS 

Google Scholar 
Hussain, I., Singh, N. B., Singh, A., Singh, H. & Singh, S. C. Green synthesis of nanoparticles and its potential application. Biotechnol. Lett. 38, 25. https://doi.org/10.1007/s10529-015-2026-7 (2016).Article 
CAS 

Google Scholar 
Singh, P., Kim, Y. J., Zhang, D. & Yang, D. C. Biological synthesis of nanoparticles from plants and microorganisms. Trends Biotechnol. 34, 25. https://doi.org/10.1016/j.tibtech.2016.02.006 (2016).Article 
CAS 

Google Scholar 
Nilavukkarasi, M., Vijayakumar, S. & Prathipkumar, S. Capparis zeylanica mediated bio-synthesized ZnO nanoparticles as antimicrobial, photocatalytic and anti-cancer applications. Mater. Sci. Energy Technol. 3, 25 (2020).
Google Scholar 
Hussain, A. et al. Biogenesis of ZnO nanoparticles using: Pandanus odorifer leaf extract: Anticancer and antimicrobial activities. RSC Adv. 9, 25 (2019).Article 

Google Scholar 
Mohamed Isa, E. D., Shameli, K., Che Jusoh, N. W., Mohamad Sukri, S. N. A. & Ismail, N. A. Variation of green synthesis techniques in fabrication of zinc oxide nanoparticles—a mini review. IOP Conf. Ser. Mater. Sci. Eng. 1051, 25 (2021).Article 

Google Scholar 
Loganathan, S., Shivakumar, M. S., Karthi, S., Nathan, S. S. & Selvam, K. Metal oxide nanoparticle synthesis (ZnO-NPs) of Knoxia sumatrensis (Retz.) DC. Aqueous leaf extract and It’s evaluation of their antioxidant, anti-proliferative and larvicidal activities. Toxicol. Rep. 8, 25 (2021).
Google Scholar 
Mahmoud, A. E. D., El-Maghrabi, N., Hosny, M. & Fawzy, M. Biogenic synthesis of reduced graphene oxide from Ziziphus spina-christi (Christ’s thorn jujube) extracts for catalytic, antimicrobial, and antioxidant potentialities. Environ. Sci. Pollut. Res. 20, 1–16 (2022).
Google Scholar 
Ahmar Rauf, M., Oves, M., Ur Rehman, F., Rauf Khan, A. & Husain, N. Bougainvillea flower extract mediated zinc oxide’s nanomaterials for antimicrobial and anticancer activity. Biomed. Pharmacother. 116, 25 (2019).Article 

Google Scholar 
Chabattula, S. C. et al. Anticancer therapeutic efficacy of biogenic Am-ZnO nanoparticles on 2D and 3D tumor models. Mater. Today Chem. 22, 25 (2021).
Google Scholar 
Berehu, H. M. et al. Cytotoxic potential of biogenic zinc oxide nanoparticles synthesized from swertia chirayita leaf extract on colorectal cancer cells. Front. Bioeng. Biotechnol. 9, 25 (2021).Article 

Google Scholar 
Khezri, K., Saeedi, M. & Maleki Dizaj, S. Application of nanoparticles in percutaneous delivery of active ingredients in cosmetic preparations. Biomed. Pharmacother. 106, 25. https://doi.org/10.1016/j.biopha.2018.07.084 (2018).Article 
CAS 

Google Scholar 
Smijs, T. G. & Pavel, S. Titanium dioxide and zinc oxide nanoparticles in sunscreens: Focus on their safety and effectiveness. Nanotechnol. Sci. Appl. 4, 25. https://doi.org/10.2147/nsa.s19419 (2011).Article 

Google Scholar 
Nasrollahzadeh, M. S. et al. Zinc oxide nanoparticles as a potential agent for antiviral drug delivery development: A systematic literature review. Curr. Nanosci. 18, 25 (2021).
Google Scholar 
Perera, W. P. T. D. et al. Albumin grafted coaxial electrosparyed polycaprolactone-zinc oxide nanoparticle for sustained release and activity enhanced antibacterial drug delivery. RSC Adv. 12, 25 (2022).Article 

Google Scholar 
Shalaby, M. A., Anwar, M. M. & Saeed, H. Nanomaterials for application in wound healing: Current state-of-the-art and future perspectives. J. Polym. Res. 29, 25. https://doi.org/10.1007/s10965-021-02870-x (2022).Article 
CAS 

Google Scholar 
Kaushik, M. et al. Investigations on the antimicrobial activity and wound healing potential of ZnO nanoparticles. Appl. Surf. Sci. 479, 25 (2019).Article 

Google Scholar 
Espitia, P. J. P., Otoni, C. G. & Soares, N. F. F. Zinc oxide nanoparticles for food packaging applications. Antimicrob. Food Packag. https://doi.org/10.1016/B978-0-12-800723-5.00034-6.4 (2016).Article 

Google Scholar 
Doan Thi, T. U. et al. Green synthesis of ZnO nanoparticles using orange fruit peel extract for antibacterial activities. RSC Adv. 10, 25 (2020).Article 

Google Scholar 
Shobha, N. et al. Synthesis and characterization of Zinc oxide nanoparticles utilizing seed source of Ricinus communis and study of its antioxidant, antifungal and anticancer activity. Mater. Sci. Eng. C 97, 25 (2019).Article 

Google Scholar 
Zahran, M. A. & Willis, A. J. The vegetation of Egypt. Plant Veget. 2, 25 (2009).
Google Scholar 
El-Borady, O. M., Fawzy, M. & Hosny, M. Antioxidant, anticancer and enhanced photocatalytic potentials of gold nanoparticles biosynthesized by common reed leaf extract. Appl. Nanosci. (Switzerland) https://doi.org/10.1007/s13204-021-01776-w (2021).Article 

Google Scholar 
Hosny, M., Fawzy, M., Abdelfatah, A. M., Fawzy, E. E. & Eltaweil, A. S. Comparative study on the potentialities of two halophytic species in the green synthesis of gold nanoparticles and their anticancer, antioxidant and catalytic efficiencies. Adv. Powder Technol. 32, 25 (2021).Article 

Google Scholar 
Vijayakumar, S. et al. Acalypha fruticosa L. Leaf extract mediated synthesis of ZnO nanoparticles: Characterization and antimicrobial activities. Mater. Today Proc. 23, 25 (2019).
Google Scholar 
Fatimah, I., Pradita, R. Y. & Nurfalinda, A. Plant extract mediated of ZnO nanoparticles by using ethanol extract of mimosa pudica leaves and coffee powder. Proced. Eng. 148, 25 (2016).Article 

Google Scholar 
Heneidy, S. Z. & Bidak, L. M. Potential uses of plant species of the coastal mediterranean region, Egypt. Pak. J. Biol. Sci. 7, 1010–1023 (2004).Article 

Google Scholar 
Manousaki, E. & Kalogerakis, N. Halophytes present new opportunities in phytoremediation of heavy metals and saline soils. Ind. Eng. Chem. Res. 50, 25 (2011).Article 

Google Scholar 
Zengin, G., Aumeeruddy-Elalfi, Z., Mollica, A., Yilmaz, M. A. & Mahomoodally, M. F. In vitro and in silico perspectives on biological and phytochemical profile of three halophyte species—a source of innovative phytopharmaceuticals from nature. Phytomedicine 38, 35–44 (2018).Article 
CAS 
PubMed 

Google Scholar 
Xin, P. et al. Surface water and groundwater interactions in salt marshes and their impact on plant ecology and coastal biogeochemistry. Rev. Geophys. 60, 5. https://doi.org/10.1029/2021RG000740 (2022).Article 

Google Scholar 
International Union for Conservation of Nature. International Union for Conservation of Nature Natural Resources IUCN Red List Categories and Criteria (IUCN, 2001).
Google Scholar 
Boulos, L. Flora of Egypt Vol 417 21–22 (Al Hadara Publishing, 1999).
Google Scholar 
Safawo, T., Sandeep, B. V., Pola, S. & Tadesse, A. Synthesis and characterization of zinc oxide nanoparticles using tuber extract of anchote (Coccinia abyssinica (Lam.) Cong.) for antimicrobial and antioxidant activity assessment. Open Nano 3, 25 (2018).
Google Scholar 
Soltanian, S. et al. Biosynthesis of zinc oxide nanoparticles using hertia intermedia and evaluation of its cytotoxic and antimicrobial activities. https://doi.org/10.1007/s12668-020-00816-z/Published.Ogbole, O. O., Segun, P. A. & Adeniji, A. J. In vitro cytotoxic activity of medicinal plants from Nigeria ethnomedicine on Rhabdomyosarcoma cancer cell line and HPLC analysis of active extracts. BMC Complement Altern. Med. 17, 25 (2017).Article 

Google Scholar 
Slater, T. F., Sawyer, B. & Sträuli, U. Studies on succinate-tetrazolium reductase systems. III Points of coupling of four different tetrazolium salts. Biochim. Biophys. Acta 77, 25 (1963).Article 

Google Scholar 
Alley, M. C. et al. Feasibility of drug screening with panels of human tumor cell lines using a microculture tetrazolium assay. Cancer Res. 48, 25 (1988).
Google Scholar 
van de Loosdrecht, A. A., Beelen, R. H. J., Ossenkoppele, G. J., Broekhoven, M. G. & Langenhuijsen, M. M. A. C. A tetrazolium-based colorimetric MTT assay to quantitate human monocyte mediated cytotoxicity against leukemic cells from cell lines and patients with acute myeloid leukemia. J. Immunol. Methods 174, 25 (1994).
Google Scholar 
Gonelimali, F. D. et al. Antimicrobial properties and mechanism of action of some plant extracts against food pathogens and spoilage microorganisms. Front. Microbiol. 9, 25 (2018).Article 

Google Scholar 
Aldalbahi, A. et al. Greener synthesis of zinc oxide nanoparticles: Characterization and multifaceted applications. Molecules 25, 25 (2020).Article 

Google Scholar 
López-Cuenca, S. et al. High-yield synthesis of zinc oxide nanoparticles from bicontinuous microemulsions. J. Nanomater. 2011, 25 (2011).Article 

Google Scholar 
Sajadi, S. M. et al. Natural iron ore as a novel substrate for the biosynthesis of bioactive-stable ZnO@CuO@iron ore NCs: A magnetically recyclable and reusable superior nanocatalyst for the degradation of organic dyes, reduction of Cr(vi) and adsorption of crude oil aromatic compounds, including PAHs. RSC Adv. 8, 62. https://doi.org/10.1039/c8ra06028b (2018).Article 
CAS 

Google Scholar 
Meena, P. L., Poswal, K. & Surela, A. K. Facile synthesis of ZnO nanoparticles for the effective photodegradation of malachite green dye in aqueous solution. Water Environ. J. 36, 25 (2022).Article 

Google Scholar 
El-Belely, E. F. et al. Green synthesis of zinc oxide nanoparticles (Zno-nps) using arthrospira platensis (class: Cyanophyceae) and evaluation of their biomedical activities. Nanomaterials 11, 25 (2021).Article 

Google Scholar 
Faye, G., Jebessa, T. & Wubalem, T. Biosynthesis, characterisation and antimicrobial activity of zinc oxide and nickel doped zinc oxide nanoparticles using Euphorbia abyssinica bark extract (2021). https://doi.org/10.1049/nbt2.12072.Dulta, K., Koşarsoy Ağçeli, G., Chauhan, P., Jasrotia, R. & Chauhan, P. K. A novel approach of synthesis zinc oxide nanoparticles by Bergenia ciliata rhizome extract: Antibacterial and anticancer potential. J. Inorg. Organomet. Polym. Mater. 31, 25 (2021).Article 

Google Scholar 
Faisal, S. et al. Green synthesis of zinc oxide (ZnO) nanoparticles using aqueous fruit extracts of Myristica fragrans: Their characterizations and biological and environmental applications. ACS Omega 6, 25 (2021).Article 

Google Scholar 
Adams, R. P. Identification of essential oil components by gas chromatography/mass spectrometry. J. Am. Soc. Mass Spectrometry 8, 25 (2007).
Google Scholar 
VStein, S., Mirokhin, D., Tchekhovskoi, D., & Nist, G. M. The NIST Mass Spectral Search Program for the NIST/EPA/NIH Mass Spectra Library. Gaithersburg, MD: Standard Reference Data Program of the National Institute of Standards and Technology (2002).Mahmoud, A. E. D., Hosny, M., El-Maghrabi, N. & Fawzy, M. Facile synthesis of reduced graphene oxide by Tecoma stans extracts for efficient removal of Ni (II) from water: Batch experiments and response surface methodology. Sustain. Environ. Res. 32, 25 (2022).Article 

Google Scholar 
Balasubramani, G. et al. GC-MS analysis of bioactive components and synthesis of gold nanoparticle using Chloroxylon swietenia DC leaf extract and its larvicidal activity. J. Photochem. Photobiol. B 148, 25 (2015).Article 

Google Scholar 
Barzinjy, A. A. & Azeez, H. H. Green synthesis and characterization of zinc oxide nanoparticles using Eucalyptus globulus Labill. leaf extract and zinc nitrate hexahydrate salt. SN Appl. Sci. 2, 25 (2020).Article 

Google Scholar 
Anitha, R., Ramesh, K. V., Ravishankar, T. N., Sudheer Kumar, K. H. & Ramakrishnappa, T. Cytotoxicity, antibacterial and antifungal activities of ZnO nanoparticles prepared by the Artocarpus gomezianus fruit mediated facile green combustion method. J. Sci. Adv. Mater. Devices 3, 25 (2018).
Google Scholar 
Chandra, H., Patel, D., Kumari, P., Jangwan, J. S. & Yadav, S. Phyto-mediated synthesis of zinc oxide nanoparticles of Berberis aristata: Characterization, antioxidant activity and antibacterial activity with special reference to urinary tract pathogens. Mater. Sci. Eng. C 102, 25 (2019).Article 

Google Scholar 
Majeed, S., Danish, M., Ismail, M. H., Ansari, M. T. & Ibrahim, M. N. M. Anticancer and apoptotic activity of biologically synthesized zinc oxide nanoparticles against human colon cancer HCT-116 cell line- in vitro study. Sustain. Chem. Pharm. 14, 25 (2019).
Google Scholar 
Miri, A., Khatami, M., Ebrahimy, O. & Sarani, M. Cytotoxic and antifungal studies of biosynthesized zinc oxide nanoparticles using extract of Prosopis farcta fruit. Green Chem. Lett. Rev. 13, 25. https://doi.org/10.1080/17518253.2020.1717005 (2020).Article 
CAS 

Google Scholar 
Ahamed, M., Akhtar, M. J., Khan, M. A. M. & Alhadlaq, H. A. Enhanced anticancer performance of eco-friendly-prepared Mo-ZnO/RGO nanocomposites: Role of oxidative stress and apoptosis. ACS Omega 7, 25 (2022).Article 

Google Scholar 
Al-Mohaimeed, A. M., Al-Onazi, W. A. & El-Tohamy, M. F. Multifunctional eco-friendly synthesis of ZnO nanoparticles in biomedical applications. Molecules 27, 25 (2022).Article 

Google Scholar 
Schreyer, M., Guo, L., Thirunahari, S., Gao, F. & Garland, M. Simultaneous determination of several crystal structures from powder mixtures: The combination of powder X-ray diffraction, band-target entropy minimization and Rietveld methods. J. Appl. Crystallogr. 47, 25 (2014).Article 

Google Scholar 
Pu, Y., Niu, Y., Wang, Y., Liu, S. & Zhang, B. Statistical morphological identification of low-dimensional nanomaterials by using TEM. Particuology 61, 11–17 (2022).Article 
CAS 

Google Scholar 
Wu, C. M., Baltrusaitis, J., Gillan, E. G. & Grassian, V. H. Sulfur dioxide adsorption on ZnO nanoparticles and nanorods. J. Phys. Chem. C 115, 10164–10172 (2011).Article 
CAS 

Google Scholar 
Saranya, S., Vijayaranai, K., Pavithra, S., Raihana, N. & Kumanan, K. In vitro cytotoxicity of zinc oxide, iron oxide and copper nanopowders prepared by green synthesis. Toxicol. Rep. 4, 25 (2017).
Google Scholar 
Chelladurai, M. et al. Anti-skin cancer activity of Alpinia calcarata ZnO nanoparticles: Characterization and potential antimicrobial effects. J Drug Deliv. Sci. Technol. 61, 102180 (2021).Article 
CAS 

Google Scholar 
Lingaraju, K., Naika, H. R., Nagabhushana, H. & Nagaraju, G. Euphorbia heterophylla (L.) mediated fabrication of ZnO NPs: Characterization and evaluation of antibacterial and anticancer properties. Biocatal. Agric. Biotechnol. 18, 25 (2019).Article 

Google Scholar 
Sana, S. S. et al. Crotalaria verrucosa leaf extract mediated synthesis of zinc oxide nanoparticles: Assessment of antimicrobial and anticancer activity. Molecules 25, 25 (2020).Article 

Google Scholar 
Bisht, G. & Rayamajhi, S. ZnO nanoparticles: A promising anticancer agent. Nanobiomedicine https://doi.org/10.5772/63437 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Bharath, B., Perinbam, K., Devanesan, S., AlSalhi, M. S. & Saravanan, M. Evaluation of the anticancer potential of Hexadecanoic acid from brown algae Turbinaria ornata on HT–29 colon cancer cells. J. Mol. Struct. 1235, 25 (2021).Article 

Google Scholar 
Selim, Y. A., Azb, M. A., Ragab, I., Abd El-Azim, H. M. & M.,. Green synthesis of zinc oxide nanoparticles using aqueous extract of Deverra tortuosa and their cytotoxic activities. Sci. Rep. 10, 25 (2020).Article 

Google Scholar 
Medini, F. et al. Phytochemical analysis, antioxidant, anti-inflammatory, and anticancer activities of the halophyte Limonium densiflorum extracts on human cell lines and murine macrophages. South Afr. J. Bot. 99, 25 (2015).Article 

Google Scholar 
Pan, M. H., Ghai, G. & Ho, C. T. Food bioactives, apoptosis, and cancer. Mol. Nutr. Food Res. 52, 20. https://doi.org/10.1002/mnfr.200700380 (2008).Article 
CAS 

Google Scholar 
Abdallah, H. M. & Ezzat, S. M. Effect of the method of preparation on the composition and cytotoxic activity of the essential oil of Pituranthos tortuosus. Z. Nat. Sect. C J. Biosci. 66 C, 25 (2011).
Google Scholar 
Iqbal, J. et al. Green synthesis of zinc oxide nanoparticles using Elaeagnus angustifolia L. leaf extracts and their multiple in vitro biological applications. Sci. Rep. 11, 25 (2021).Article 

Google Scholar 
Norouzi Jobie, F., Ranjbar, M., Hajizadeh Moghaddam, A. & Kiani, M. Green synthesis of zinc oxide nanoparticles using Amygdalus scoparia Spach stem bark extract and their applications as an alternative antimicrobial, anticancer, and anti-diabetic agent. Adv. Powder Technol. 32, 21 (2021).Article 

Google Scholar 
Chen, F. C., Huang, C. M., Yu, X. W. & Chen, Y. Y. Effect of nano zinc oxide on proliferation and toxicity of human gingival cells. Hum. Exp. Toxicol. 41, 15 (2022).Article 

Google Scholar 
Sajjad, A. et al. Photoinduced fabrication of zinc oxide nanoparticles: Transformation of morphological and biological response on light irradiance. ACS Omega 6, 75 (2021).Article 

Google Scholar 
Sohail, M. F. et al. Green synthesis of zinc oxide nanoparticles by neem extract as multi-facet therapeutic agents. J. Drug Deliv. Sci. Technol. 59, 15 (2020).
Google Scholar 
Lopes, M., Sanches-Silva, A., Castilho, M., Cavaleiro, C. & Ramos, F. Halophytes as source of bioactive phenolic compounds and their potential applications. Crit. Rev. Food Sci. Nutr. 20, 20. https://doi.org/10.1080/10408398.2021.1959295 (2021).Article 
CAS 

Google Scholar 
Bouarab-Chibane, L. et al. Antibacterial properties of polyphenols: Characterization and QSAR (quantitative structure-activity relationship) models. Front. Microbiol. 10, 77 (2019).Article 

Google Scholar 
Guimarães, A. C. et al. Antibacterial activity of terpenes and terpenoids present in essential oils. Molecules 24, 11 (2019).Article 

Google Scholar 
Sirelkhatim, A. et al. Review on zinc oxide nanoparticles: Antibacterial activity and toxicity mechanism. Nano-Micro Lett. 7, 219–242. https://doi.org/10.1007/s40820-015-0040-x (2015).Article 
CAS 

Google Scholar 
Singh, T. A. et al. A state of the art review on the synthesis, antibacterial, antioxidant, antidiabetic and tissue regeneration activities of zinc oxide nanoparticles. Adv. Coll. Interface Sci. 295, 25. https://doi.org/10.1016/j.cis.2021.102495 (2021).Article 
CAS 

Google Scholar 
Gao, Y. et al. Biofabrication of zinc oxide nanoparticles from Aspergillus niger, their antioxidant, antimicrobial and anticancer activity. J. Clust. Sci. 30, 11 (2019).Article 

Google Scholar 
Luo, Q. et al. Terpenoid composition and antioxidant activity of extracts from four chemotypes of Cinnamomum camphora and their main antioxidant agents. Biofuels Bioprod. Biorefin. 16, 510–522 (2022).Article 
CAS 

Google Scholar 
Bose, J., Rodrigo-Moreno, A. & Shabala, S. ROS homeostasis in halophytes in the context of salinity stress tolerance. J. Exp. Bot. 65, 25. https://doi.org/10.1093/jxb/ert430 (2014).Article 
CAS 

Google Scholar  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