Ecology

Shuang Gao from Bjerkens Center for Climate Research in Norway, and colleagues from Germany and the United States explored future changes in marine primary production and carbon uptake under climate scenarios using the Norwegian Earth-system model, with four river transport configurations incorporating established future economic development and nutrient-use efficiency pathways. The researchers find that riverine nutrient inputs lessen nutrient limitation under warmer conditions. In the future, the effect of increased riverine carbon may be larger than the effect of nutrient inputs on the projections of ocean carbon uptake. In the historical period, increased nutrient inputs are considered the most prominent driver of carbon uptake. The results of this study are subject to model limitations, and high-resolution models should be used to assess the future impact. More

Boudreau BP, Huettel M, Forster S, Jahnke RA, McLachlan A, Middelburg JJ, et al. Permeable marine sediments: Overturning an old paradigm. Eos, Trans Am Geophys Union. 2001;82:133–6.Article 

Google Scholar 
Cook PL, Wenzhöfer F, Rysgaard S, Galaktionov OS, Meysman FJ, Eyre BD, et al. Quantification of denitrification in permeable sediments: Insights from a two‐dimensional simulation analysis and experimental data. Limnol Oceanogr Methods. 2006;4:294–307.Article 
CAS 

Google Scholar 
Evrard V, Glud RN, Cook PL. The kinetics of denitrification in permeable sediments. Biogeochemistry. 2013;113:563–72.Article 

Google Scholar 
Huettel M, Berg P, Kostka JE. Benthic exchange and biogeochemical cycling in permeable sediments. Ann Rev Marine Sci. 2014;6:23–51.Article 

Google Scholar 
Rao AMF, McCarthy MJ, Gardner WS, Jahnke RA. Respiration and denitrification in permeable continental shelf deposits on the South Atlantic Bight: Rates of carbon and nitrogen cycling from sediment column experiments. Continental Shelf Res. 2007;27:1801–19.Article 

Google Scholar 
Billerbeck M, Werner U, Polerecky L, Walpersdorf E, de Beer D, Huettel M Surficial and deep pore water circulation governs spatial and temporal scales of nutrient recycling in intertidal sand flat sediment. Marine Ecol Progr Series. 2006;326:61–76.Jansen S, Walpersdorf E, Werner U, Billerbeck M, Böttcher ME, de Beer D. Functioning of intertidal flats inferred from temporal and spatial dynamics of O2, H2S and pH in their surface sediment. Ocean Dyn. 2009;59:317–32.Article 

Google Scholar 
de Beer D, Wenzhöfer F, Ferdelman TG, Boehme SE, Huettel M, van Beusekom JEE, et al. Transport and mineralization rates in North Sea sandy intertidal sediments, Sylt-Rømø Basin, Wadden Sea. Limnol Oceanogr. 2005;50:113–27.Article 

Google Scholar 
Gao H, Matyka M, Liu B, Khalili A, Kostka JE, Collins G, et al. Intensive and extensive nitrogen loss from intertidal permeable sediments of the Wadden Sea. Limnol Oceanogr. 2012;57:185–98.Article 
CAS 

Google Scholar 
Gao H, Schreiber F, Collins G, Jensen MM, Kostka JE, Lavik G, et al. Aerobic denitrification in permeable Wadden Sea sediments. ISME J. 2009;4:417.Article 
PubMed 

Google Scholar 
Elliott AH, Brooks NH. Transfer of nonsorbing solutes to a streambed with bed forms: Theory. Water Resour Res. 1997;33:123–36.Article 
CAS 

Google Scholar 
Precht E, Huettel M. Advective pore‐water exchange driven by surface gravity waves and its ecological implications. Limnol Oceanogr. 2003;48:1674–84.Article 

Google Scholar 
Ahmerkamp S, Marchant HK, Peng C, Probandt D, Littmann S, Kuypers MM, et al. The effect of sediment grain properties and porewater flow on microbial abundance and respiration in permeable sediments. Sci Rep. 2020;10:1–12.Article 

Google Scholar 
Ahmerkamp S, Winter C, Krämer K, Beer DD, Janssen F, Friedrich J, et al. Regulation of benthic oxygen fluxes in permeable sediments of the coastal ocean. Limnol Oceanogr. 2017;62:1935–54.Article 
CAS 

Google Scholar 
Cardenas MB, Wilson JL Dunes, turbulent eddies, and interfacial exchange with permeable sediments. Water Resour Res. 2007;43:W08412.Santos IR, Eyre BD, Huettel M. The driving forces of porewater and groundwater flow in permeable coastal sediments: a review. Estuarine, Coastal Shelf Sci. 2012;98:1–15.Article 

Google Scholar 
Probandt D, Eickhorst T, Ellrott A, Amann R, Knittel K. Microbial life on a sand grain: from bulk sediment to single grains. ISME J. 2018;12:623–33.Article 
PubMed 

Google Scholar 
Marchant HK, Ahmerkamp S, Lavik G, Tegetmeyer HE, Graf J, Klatt JM, et al. Denitrifying community in coastal sediments performs aerobic and anaerobic respiration simultaneously. ISME J. 2017;11:1799.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Marchant HK, Holtappels M, Lavik G, Ahmerkamp S, Winter C, Kuypers MMM. Coupled nitrification–denitrification leads to extensive N loss in subtidal permeable sediments. Limnol Oceanogr. 2016;61:1033–48.Article 

Google Scholar 
Marchant HK, Tegetmeyer HE, Ahmerkamp S, Holtappels M, Lavik G, Graf J, et al. Metabolic specialization of denitrifiers in permeable sediments controls N2O emissions. Environ Microbiol. 2018;20:4486–502.Article 
CAS 
PubMed 

Google Scholar 
Laverman AM, Pallud C, Abell J, Van, Cappellen P. Comparative survey of potential nitrate and sulfate reduction rates in aquatic sediments. Geochimica et Cosmochimica Acta. 2012;77:474–88.Article 
CAS 

Google Scholar 
Fenchel T, Jørgensen B. Detritus food chains of aquatic ecosystems: the role of bacteria. Adv Microb Ecol. 1977;1:1–58.Article 
CAS 

Google Scholar 
Canfield DE, Kristensen E, Thamdrup B Aquatic Geomicrobiology: Elsevier Science; 2005.Froelich PN, Klinkhammer G, Bender ML, Luedtke N, Heath GR, Cullen D, et al. Early oxidation of organic matter in pelagic sediments of the eastern equatorial Atlantic: suboxic diagenesis. Geochimica et cosmochimica Acta. 1979;43:1075–90.Article 
CAS 

Google Scholar 
Eckford RE, Fedorak PM. Chemical and microbiological changes in laboratory incubations of nitrate amendment “sour” produced waters from three western Canadian oil fields. J Ind Microbiol Biotechnol. 2002;29:243–54.Article 
CAS 
PubMed 

Google Scholar 
Grigoryan AA, Cornish SL, Buziak B, Lin S, Cavallaro A, Arensdorf JJ, et al. Competitive oxidation of volatile fatty acids by sulfate- and nitrate-reducing bacteria from an oil field in Argentina. Appl Environ Microbiol. 2008;74:4324.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Hubert C, Nemati M, Jenneman G, Voordouw G. Containment of biogenic sulfide production in continuous up-flow packed-bed bioreactors with nitrate or nitrite. Biotechnol Progr. 2003;19:338–45.Article 
CAS 

Google Scholar 
Greene EA, Hubert C, Nemati M, Jenneman GE, Voordouw G. Nitrite reductase activity of sulphate-reducing bacteria prevents their inhibition by nitrate-reducing, sulphide-oxidizing bacteria. Environ Microbiol. 2003;5:607–17.Article 
CAS 
PubMed 

Google Scholar 
Wolfe BM, Lui SM, Cowan JA. Desulfoviridin, a multimeric-dissimilatory sulfite reductase from Desulfovibrio vulgaris (Hildenborough) Purification, characterization, kinetics and EPR studies. Eur J Biochem. 1994;223:79–89.Article 
CAS 
PubMed 

Google Scholar 
Fossing H, Gallardo VA, Jørgensen BB, Hüttel M, Nielsen LP, Schulz H, et al. Concentration and transport of nitrate by the mat-forming sulphur bacterium Thioploca. Nature. 1995;374:713–15.Article 
CAS 

Google Scholar 
Jørgensen BB. Big sulfur bacteria. ISME J. 2010;4:1083.Article 
PubMed 

Google Scholar 
Marzocchi U, Trojan D, Larsen S, Louise Meyer R, Peter Revsbech N, Schramm A, et al. Electric coupling between distant nitrate reduction and sulfide oxidation in marine sediment. ISME J. 2014;8:1682.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Londry KL, Suflita JM. Use of nitrate to control sulfide generation by sulfate-reducing bacteria associated with oily waste. J Ind Microbiol Biotechnol. 1999;22:582–9.Article 
CAS 
PubMed 

Google Scholar 
McInerney MJ, Bhupathiraju VK, Sublette KL. Evaluation of a microbial method to reduce hydrogen sulfide levels in a porous rock biofilm. J Ind Microbiol. 1992;11:53–8.Article 
CAS 

Google Scholar 
Schwermer CU, Ferdelman TG, Stief P, Gieseke A, Rezakhani N, Van Rijn J, et al. Effect of nitrate on sulfur transformations in sulfidogenic sludge of a marine aquaculture biofilter. FEMS Microbiol Ecol. 2010;72:476–84.Article 
CAS 
PubMed 

Google Scholar 
Thamdrup B, Fossing H, Jørgensen BB. Manganese, iron and sulfur cycling in a coastal marine sediment, Aarhus Bay, Denmark. Geochimica et Cosmochimica Acta. 1994;58:5115–29.Article 
CAS 

Google Scholar 
Al-Raei AM, Bosselmann K, Böttcher ME, Hespenheide B, Tauber F. Seasonal dynamics of microbial sulfate reduction in temperate intertidal surface sediments: controls by temperature and organic matter. Ocean Dyn. 2009;59:351–70.Article 

Google Scholar 
Dyksma S, Pjevac P, Ovanesov K, Mussmann M. Evidence for H2 consumption by uncultured Desulfobacterales in coastal sediments. Environ Microbiol. 2018;20:450–61.Article 
CAS 
PubMed 

Google Scholar 
Musat N, Werner U, Knittel K, Kolb S, Dodenhof T, van Beusekom JEE, et al. Microbial community structure of sandy intertidal sediments in the North Sea, Sylt-Rømø Basin, Wadden Sea. Syst Appl Microbiol. 2006;29:333–48.Article 
PubMed 

Google Scholar 
Mußmann M, Ishii K, Rabus R, Amann R. Diversity and vertical distribution of cultured and uncultured Deltaproteobacteria in an intertidal mud flat of the Wadden Sea. Environ Microbiol. 2005;7:405–18.Article 
PubMed 

Google Scholar 
Dyksma S, Lenk S, Sawicka JE, Mußmann M. Uncultured gammaproteobacteria and desulfobacteraceae account for major acetate assimilation in a coastal marine sediment. Front Microbiol. 2018;9:3124Article 
PubMed 
PubMed Central 

Google Scholar 
Chen J, Hanke A, Tegetmeyer HE, Kattelmann I, Sharma R, Hamann E, et al. Impacts of chemical gradients on microbial community structure. ISME J. 2017;11:920.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Saad S, Bhatnagar S, Tegetmeyer HE, Geelhoed JS, Strous M, Ruff SE. Transient exposure to oxygen or nitrate reveals ecophysiology of fermentative and sulfate‐reducing benthic microbial populations. Environ Microbiol. 2017;19:4866–81.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Brunet RC, Garcia-Gil LJ. Sulfide-induced dissimilatory nitrate reduction to ammonia in anaerobic freshwater sediments. FEMS Microbiol Ecol. 1996;21:131–8.Article 
CAS 

Google Scholar 
Murphy AE, Bulseco AN, Ackerman R, Vineis JH, Bowen JL. Sulphide addition favours respiratory ammonification (DNRA) over complete denitrification and alters the active microbial community in salt marsh sediments. Environ Microbiol. 2020;22:2124–39.Article 
CAS 
PubMed 

Google Scholar 
Krekeler D, Cypionka H. The preferred electron acceptor of Desulfovibrio desulfuricans CSN. FEMS Microbiol Ecol. 1995;17:271–7.Article 
CAS 

Google Scholar 
Seitz H-J, Cypionka H. Chemolithotrophic growth of Desulfovibrio desulfuricans with hydrogen coupled to ammonification of nitrate or nitrite. Arch Microbiol. 1986;146:63–7.Article 
CAS 

Google Scholar 
Dalsgaard T, Bak F. Nitrate reduction in a sulfate-reducing bacterium, Desulfovibrio desulfuricans, isolated from rice paddy soil: sulfide inhibition, kinetics, and regulation. Appl Environ Microbiol. 1994;60:291–7.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Marietou A. Nitrate reduction in sulfate-reducing bacteria. FEMS Microbiol Lett. 2016;363:fnw155.Article 
PubMed 

Google Scholar 
Marietou A, Griffiths L, Cole J. Preferential reduction of the thermodynamically less favorable electron acceptor, sulfate, by a nitrate-reducing strain of the sulfate-reducing bacterium Desulfovibrio desulfuricans 27774. J Bacteriol. 2009;191:882–889.Article 
CAS 
PubMed 

Google Scholar 
Korte HL, Saini A, Trotter VV, Butland GP, Arkin AP, Wall JD. Independence of nitrate and nitrite inhibition of Desulfovibrio vulgaris Hildenborough and use of nitrite as a substrate for growth. Environ Sci Technol. 2015;49:924–931.Article 
CAS 
PubMed 

Google Scholar 
Pereira IA, LeGall J, Xavier AV, Teixeira M. Characterization of a heme c nitrite reductase from a non-ammonifying microorganism, Desulfovibrio vulgaris Hildenborough. Biochimica et Biophysica Acta (BBA)-Protein Struct Mol Enzymol. 2000;1481:119–130.Article 
CAS 

Google Scholar 
Werner U, Billerbeck M, Polerecky L, Franke U, Huettel M, van Beusekom JEE, et al. Spatial and temporal patterns of mineralization rates and oxygen distribution in a permeable intertidal sand flat (Sylt, Germany). Limnol Oceanogr. 2006;51:2549–63.Article 
CAS 

Google Scholar 
Marchant HK, Lavik G, Holtappels M, Kuypers MMM. The fate of nitrate in intertidal permeable sediments. PLOS ONE. 2014;9:e104517.Article 
PubMed 
PubMed Central 

Google Scholar 
Canfield DE. Reactive iron in marine sediments. Geochimica et cosmochimica acta. 1989;53:619–632.Article 
CAS 
PubMed 

Google Scholar 
Billerbeck M, Werner U, Bosselmann K, Walpersdorf E, Huettel M. Nutrient release from an exposed intertidal sand flat. Marine Ecol Progr Series. 2006;316:35–51.Article 
CAS 

Google Scholar 
Canfield DE, Stewart FJ, Thamdrup B, De Brabandere L, Dalsgaard T, Delong EF, et al. A cryptic sulfur cycle in oxygen-minimum–zone waters off the Chilean Coast. Science. 2010;330:1375.Article 
CAS 
PubMed 

Google Scholar 
Jørgensen BB, Findlay AJ, Pellerin A. The biogeochemical sulfur cycle of marine sediments. Front Microbiol. 2019;10:849.Article 
PubMed 
PubMed Central 

Google Scholar 
Nemati M, Mazutinec TJ, Jenneman GE, Voordouw G. Control of biogenic H2S production with nitrite and molybdate. J Ind Microbiol Biotechnol. 2001;26:350–355.Article 
CAS 
PubMed 

Google Scholar 
Haveman SA, Greene EA, Stilwell CP, Voordouw JK, Voordouw G. Physiological and Gene Expression Analysis of Inhibition of Desulfovibrio vulgaris Hildenborough by Nitrite. J Bacteriol. 2004;186:7944–7950.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Behrendt A, de Beer D, Stief P. Vertical activity distribution of dissimilatory nitrate reduction in coastal marine sediments. Biogeosciences. 2013;10:7509–23.Article 

Google Scholar 
Findlay AJ, Pellerin A, Laufer K, Jørgensen BB. Quantification of sulphide oxidation rates in marine sediment. Geochimica et Cosmochimica Acta. 2020;280:441–52.Article 
CAS 

Google Scholar 
Waite DW, Chuvochina M, Pelikan C, Parks DH, Yilmaz P, Wagner M, et al. Proposal to reclassify the proteobacterial classes Deltaproteobacteria and Oligoflexia, and the phylum Thermodesulfobacteria into four phyla reflecting major functional capabilities. Int J Syst Evolut Microbiol. 2020;70:5972–6016.Article 
CAS 

Google Scholar 
Dyksma S, Bischof K, Fuchs BM, Hoffmann K, Meier D, Meyerdierks A, et al. Ubiquitous Gammaproteobacteria dominate dark carbon fixation in coastal sediments. ISME J. 2016;10:1939–1953.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Lenk S, Arnds J, Zerjatke K, Musat N, Amann R, Mußmann M. Novel groups of Gammaproteobacteria catalyse sulfur oxidation and carbon fixation in a coastal, intertidal sediment. Environ Microbiol. 2011;13:758–774.Article 
CAS 
PubMed 

Google Scholar 
An S, Gardner W S. Dissimilatory nitrate reduction to ammonium (DNRA) as a nitrogen link, versus denitrification as a sink in a shallow estuary (Laguna Madre/Baffin Bay, Texas). Marine Ecol Progr Series. 2002;237:41–50.Article 
CAS 

Google Scholar 
Wankel SD, Ziebis W, Buchwald C, Charoenpong C, de Beer D, Dentinger J, et al. Evidence for fungal and chemodenitrification based N2O flux from nitrogen impacted coastal sediments. Nat Commun. 2017;8:15595.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Moura I, Bursakov S, Costa C, Moura JJ. Nitrate and nitrite utilization in sulfate-reducing bacteria. Anaerobe. 1997;3:279–290.Article 
CAS 
PubMed 

Google Scholar 
Song G, Liu S, Zhang J, Zhu Z, Zhang G, Marchant HK, et al. Response of benthic nitrogen cycling to estuarine hypoxia. Limnol Oceanogr. 2021;66:652–66.Article 
CAS 

Google Scholar 
Tiedje JM, Sexstone AJ, Myrold DD, Robinson JA. Denitrification: ecological niches, competition and survival. Antonie van Leeuwenhoek. 1983;48:569–583.Article 

Google Scholar 
Strohm TO, Griffin B, Zumft WG, Schink B. Growth yields in bacterial denitrification and nitrate ammonification. Appl Environ Microbiol. 2007;73:1420–1424.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Rütting T, Boeckx P, Müller C, Klemedtsson L. Assessment of the importance of dissimilatory nitrate reduction to ammonium for the terrestrial nitrogen cycle. Biogeosciences. 2011;8:1779–91.Article 

Google Scholar 
Røy H, Lee JS, Jansen S, de Beer D. Tide-driven deep pore-water flow in intertidal sand flats. Limnol Oceanogr. 2008;53:1521–30.Article 

Google Scholar 
Cline JD. Spectrophotometric determination of hydrogen sulfide in natural waters 1. Limnol Oceanogr. 1969;14:454–458.Article 
CAS 

Google Scholar 
Viollier E, Inglett P, Hunter K, Roychoudhury A, Van, Cappellen P. The ferrozine method revisited: Fe (II)/Fe (III) determination in natural waters. Appl Geochem. 2000;15:785–90.Article 
CAS 

Google Scholar 
Røy H, Weber HS, Tarpgaard IH, Ferdelman TG, Jørgensen BB. Determination of dissimilatory sulfate reduction rates in marine sediment via radioactive 35S tracer. Limnol Oceanogr Methods. 2014;12:196–211.Article 

Google Scholar 
García-Robledo E, Corzo A, Papaspyrou S. A fast and direct spectrophotometric method for the sequential determination of nitrate and nitrite at low concentrations in small volumes. Marine Chem. 2014;162:30–36.Article 

Google Scholar 
Miranda KM, Espey MG, Wink DA. A rapid, simple spectrophotometric method for simultaneous detection of nitrate and nitrite. Nitric Oxide. 2001;5:62–71.Article 
CAS 
PubMed 

Google Scholar 
Holtappels M, Lavik G, Jensen MM, Kuypers MMM Chapter ten – 15N-Labeling Experiments to Dissect the Contributions of Heterotrophic Denitrification and Anammox to Nitrogen Removal in the OMZ Waters of the Ocean. In: Klotz MG, editor. Methods in Enzymology. 486: Academic Press; 2011. p. 223-251.Preisler A, De Beer D, Lichtschlag A, Lavik G, Boetius A, Jørgensen BB. Biological and chemical sulfide oxidation in a Beggiatoa inhabited marine sediment.ISME J. 2007;1:341–353.Article 
CAS 
PubMed 

Google Scholar 
Warembourg FR 5 – Nitrogen fixation in soil and plant systems. In: Knowles R, Blackburn TH, editors. Nitrogen Isotope Techniques. San Diego: Academic Press; 1993. p. 127-156.Orellana LH, Rodriguez-R LM, Konstantinidis KT. ROCker: accurate detection and quantification of target genes in short-read metagenomic data sets by modeling sliding-window bitscores. Nucleic Acids Res. 2016;45:e14–e14.PubMed Central 

Google Scholar 
Menzel P, Ng KL, Krogh A. Fast and sensitive taxonomic classification for metagenomics with Kaiju. Nat Commun. 2016;7:11257.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Anantharaman K, Hausmann B, Jungbluth SP, Kantor RS, Lavy A, Warren LA, et al. Expanded diversity of microbial groups that shape the dissimilatory sulfur cycle. ISME J. 2018;12:1715–28.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Watanabe T, Kojima H, Fukui M. Identity of major sulfur-cycle prokaryotes in freshwater lake ecosystems revealed by a comprehensive phylogenetic study of the dissimilatory adenylylsulfate reductase. Sci Rep. 2016;6:36262.Article 
CAS 
PubMed 
PubMed Central 

Google Scholar  More

Scanning electron microscopy and energy spectrum analysisThe SEM images show the morphological structures of DA and Al-DA before and after adsorption (Fig. 1). DA and Al-DA have disk-like microstructures29 with sur-faces containing both large and small pores, that is, DA and Al-DA have unique multi-level pore structures. The main component of DA and Al-DA is silica, which has a large specific surface area, good thermal stability, and is a natural green material for use as a water treatment agent with a porous structure31. The micrographs show that before adsorption, the DA surface is smooth with a distinct pore structure, whereas modification with aluminium hydroxide makes DA coarse and loose because of the formation of amorphous aluminium hydroxide colloids32. After adsorption, the surface pore structure is covered over for DA and completely covered over for Al-DA, which indicates that F− reacts with Al3+ to form nanoscale precipitates22. The results of the EDS analysis (Fig. 2) show that the content of elemental Al increased from 3.96 to 12.74% after DA was modified with aluminium hydroxide, indicating that Al adhered effectively to the modified DA surface. After adsorption, the content of elemental Al decreased from 3.96 to 1.36% for DA and from 12.74 to 2.03% for Al-DA, which fully confirmed that fluorine preferentially combined with Al to form aluminium precipitates during adsorption, thereby decreasing the Al content.Figure 1SEM images of DA and Al-DA before and after adsorption. (A) Before DA adsorption. (B) After DA adsorption. (C) Before Al-DA adsorption. (D) After Al-DA adsorption.Full size imageFigure 2EDS graphs of DA and Al-DA before and after adsorption. (A) Before DA adsorption. (B) After DA adsorption. (C) Before Al-DA adsorption. (D) After Al-DA adsorption.Full size imageXRD analysisThe surface mineral composition and crystallinity of the materials before and after adsorption were analyzed by XRD (Fig. 3). In the DA and Al-DA patterns, the wide diffraction peaks at approximately 22.0°, 26.0°, and 50.0° mainly correspond to amorphous SiO2, and the diffraction peak at approximately 35° mainly corresponds to amorphous Al2O3, indicating that the material is polycrystalline29. It has been re-ported that amorphous materials may be good adsorbents because of a large specific surface area and numerous active sites33. Many Al(OH)3 peaks and NaCl peaks appear in the XRD pattern of Al-DA, indicating the successful modification of DA by aluminium hydroxide. After adsorption, Na3AlF6 peaks appear in the DA pattern, and Na3AlF6 and AlF3 peaks appear in the Al-DA pattern, whereas the characteristic peaks of NaCl are absent in the Al-DA pattern, which indicates the participation of NaCl in the adsorption process. It has been demonstrated that in the presence of excess sodium fluoride in the reaction solution, the generated aluminium fluoride combines with sodium fluoride to form a NaAlF4 intermediate, which is subsequently converted to cryolite complexes by further adsorption of sodium fluoride34. This result confirms the XRD mapping results.Figure 3XRD patterns of DA and Al-DA before and after adsorption.Full size imageInfrared analysisFigure 4 shows the FTIR spectra of DA and Al-DA before and after adsorption: peaks at 3418, 1635, 1096, 791, and 538 cm−1 appear in the spectrum of DA spectrum before adsorption, and peaks at 3630, 3449, 1637, 1094, 913, 793, and 538 cm−1, appear in the Al-DA spectrum before adsorption. The strong and broad band centered at 3418 cm−1 is due to the stretching vibration of the adsorbed water hydroxyl group (O–H) and the surface hydroxyl group, the vibrational peak at approximately 1635 cm−1 is probably from bound water or the surface hydroxyl group; the peaks at 1096 cm−1 and 538 cm−1 correspond to siloxane groups (Si–O–Si–) and an Al–O absorption band, respectively; and the strong oscillations at 791 cm−1 may be attributed to inorganic Al salts35,36,37. The original absorption peak in the DA spectrum is shifted in the spectrum of DA modified with aluminium hydroxide, confirming the successful modification of DA. The shift of the band at 3418 cm−1 in the DA spectrum to a higher frequency at 3623 cm−1 in the DA spectrum after fluoride absorption is caused by fluoride bonding and has been previously reported38. Another noticeable change in the spectra of DA and Al-DA before and after adsorption is the increase or decrease in the intensity of bending vibrations of specific peaks because the highly electronegative fluoride may have an inductive effect on the respective groups that leads to a blueshift, and the formation of hydrogen bonds leads to a redshift and broadening of the spectral band. The shifts and changes of these peaks indicate the interaction of fluoride with the respective groups29. The new peak at approximately 1170 cm−1 in the spectra of DA and Al-DA with adsorbed fluoride may be due to the formation of Al-F bonds6. The IR spectra show that the formation of a new bonding electronic structure by surface complexation with F− is one of the main mechanisms for the adsorption of F−.Figure 4FTIR spectra of DA and Al-DA before and after adsorption.Full size imageZeta potential analysisThe zeta potential of the material surface plays a very key role in the adsorption process, which reflects the surface charge properties of the material under different pH conditions, and also reflects the surface properties of the material. To obtain the zero charge point of the material, we studied the potential change of the material under different pH values. The results are shown in Fig. 5. In the range of pH 3–11, the zeta potential of the two adsorbents decreased linearly with the increase in pH, and the pHzpc of DA and Al-DA were 9.84 and 10.61, respectively. When pH  More

Traditional ecological footprint consumption accountsTo truly reflect the ecological footprint and ecological carrying capacity of Dongying city, according to the lifestyle and consumption of Dongying city and with reference to Shandong Province Statistical Yearbook and Dongying City Statistical Yearbook, the biologically productive land is divided into arable land, forestland, grassland, water, construction land and fossil energy land, and the main consumption items of each category are shown in Fig. 3.Figure 3Traditional ecological footprint consumption accounts in Dongying city. This paper uses the carbon footprint to improve the fossil energy footprint of the traditional ecological footprint.Full size imageNPP-based correction of ecological footprint parametersThe 30 m land use of the study area was resampled to 500 m, consistent with the resolution of MOD17A3H after pre-processing with MRT and other tools. Correction of ecological footprint parameter factors in Dongying City for 2015, 2018 and 2020 based on the annual average NPP of vegetation (Table 1). This method is faster and more accurate than other methods, and the implementation of NPP calculations from the vegetation light energy use efficiency (LUE) framework to correct ecological footprint parameters is more applicable and accurate than other methods.Table 1 Average annual net primary productivity per land type in the Yellow River Delta.Full size tableYield factorThe formula for calculating the yield factor for arable land in the Yellow River Delta refers to NFA 2016:$$left{ {begin{array}{*{20}c} {Y_{j1} = frac{{Sigma A_{W} }}{{Sigma A_{N} }}} \ {A_{N} = frac{{P_{N} }}{{Y_{N} }}} \ {A_{W} = frac{{P_{N} }}{{Y_{W} }}} \ end{array} } right.$$
(1)
In Eq. (1), ({Y}_{j1}) is the yield factor of the arable land in the study area, ({A}_{N}) is the harvested area ( culture area ) of agricultural products of category (N) in the study area, ({A}_{W}) is the area required to produce an equivalent amount of this type of agricultural product based on the world average yield, ({P}_{N}) is the production of agricultural products of category (N) under the region, ({Y}_{N}) is the average yield of agricultural products of category (N) under the region, and ({Y}_{W}) is the world average production of a category of agricultural products.The NPP products from MODIS supported by remote sensing were used as the base data to correct the yield factors of woodlands and grasslands in the study area under the ecological footprint model.$$Y_{{{text{j}}2}} = overline{{NPP_{local} }} /overline{{NPP_{global} }}$$
(2)
In Eq. (2), ({Y}_{mathrm{j}2}) is the yield factor for woodland and grassland in the study area, ({NPP}_{local}) is the average annual net primary productivity of woodland and grassland in the study area in the corresponding year, and ({NPP}_{global}) is the global average NPP of woodland and grassland in the corresponding year, referring to Amthor et al.24.In addition, most of the land for construction comes from cropland, so the yield factor for construction land is the same as that for cropland25. The yield factors for the watershed were derived from the Wackernagel and Rees26 study.Balancing factorThe NPP model for provincial hectares was applied to the municipal scale. Among them, the NPP of four biologically productive lands, namely cropland, woodland, grassland and water, was weighted and summed to obtain the annual average NPP within the city area.$$overline{NPP} = frac{{mathop sum nolimits_{j} left( {A_{j} times NPP_{j} } right)}}{{mathop sum nolimits_{j} A_{j} }}$$
(3)
In Eq. (3), (overline{NPP }) is the average net primary productivity of arable land, forestland, grassland and water in Dongying, ({A}_{j}) is the area of land in category (j), and ({NPP}_{j}) is the average annual NPP of productive land in category (j).Balancing factors for arable land, woodland, grassland and water in the Yellow River Delta.$$R_{j} = frac{{NPP_{j} }}{{overline{NPP} }}$$
(4)
In Eq. (4), ({R}_{j}) is a balancing factor.The sites for construction are located in areas suitable for agricultural cultivation or directly occupy arable land, so the potential ecological productivity of urban construction land is the same as that of arable land, and therefore the equilibrium factor for construction land is equal to that of arable land27.Ecological footprint principles and improvementsEcological footprint modelEcological footprint model includes ecological footprint, ecological carrying capacity and ecological deficit. As the study area is within the city limits and the statistics have their own characteristics, adjustments have been made to the methodology for calculating the national ecological footprint accounts28. Based on the biological consumption account, the ecological footprint can be calculated for any land use type.$$EF = frac{P}{{Y_{N} }} times R_{j} times Y_{j}$$
(5)
In Eq. (5), (P) is the number of biologically productive land harvesting consumption items in a category, and ({Y}_{N}) is the average production of consumption Item (N) in the region. The ecological footprint of the construction land is measured based on the area of human infrastructure land and is equal to its ecological carrying capacity.Ecological carrying capacity is the determination of the maximum carrying capacity of an ecosystem for human activity, expressed as the sum of the biologically productive land area available in an area.$$EC = N times ec = N times sum left( {a_{j} times R_{j} times Y_{j} } right)$$
(6)
In Eq. (6), (EC) is the ecological carrying capacity per capita, and ({a}_{j}) is the per capita area of biologically productive land of category j in the region. According to the recommendations of the World Commission on Environment and Development, 12% of the ecological carrying capacity should also be deducted for biodiversity conservation. The population figures for the study area were obtained from the statistical yearbook and the seventh national census data. According to the recommendations of the World Commission on Environment and Development, 12% of the ecological carrying capacity should also be deducted for biodiversity conservation.An ecological deficit is the interpolation of the ecological footprint and ecological carrying capacity.$$ED = EF – EC$$
(7)
When (ED >0) indicates an ecological deficit, the ecological environment has exceeded the carrying capacity. Conversely, when (ED More

Nigst, P. R. et al. Early modern human settlement of Europe north of the alps occurred 43,500 years ago in a cold steppe-type environment. Proc. Natl. Acad. Sci. U.S.A. 111, 14394–14399 (2014).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Pederzani, S. et al. Subarctic climate for the earliest Homo sapiens in Europe. Sci. Adv. 7, 1–11 (2021).Article 

Google Scholar 
Shao, Y. et al. Human-existence probability of the Aurignacian techno-complex under extreme climate conditions. Quat. Sci. Rev. 263, 106995 (2021).Article 

Google Scholar 
Slimak, L. et al. Modern human incursion into Neanderthal territories 54,000 years ago at Mandrin, France. Sci. Adv. 8, 1 (2022).Article 

Google Scholar 
Fewlass, H. et al. A 14C chronology for the Middle to Upper Palaeolithic transition at Bacho Kiro Cave, Bulgaria. Nat. Ecol. Evol. 4, 794–801 (2020).Article 
PubMed 

Google Scholar 
Hublin, J. J. et al. Initial Upper Palaeolithic Homo sapiens from Bacho Kiro Cave, Bulgaria. Nature 581, 299–302 (2020).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Benazzi, S. et al. Early dispersal of modern humans in Europe and implications for Neanderthal behaviour. Nature 479, 525–528 (2011).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Benazzi, S. et al. The makers of the Protoaurignacian and implications for Neandertal extinction. Science 348, 793–796 (2015).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Cortés-Sánchez, M. et al. An early Aurignacian arrival in southwestern Europe. Nat. Ecol. Evol. 3, 207–212 (2019).Article 
PubMed 

Google Scholar 
Vidal-Cordasco, M., Ocio, D., Hickler, T. & Marín-Arroyo, A. B. Publisher Correction: Ecosystem productivity affected the spatiotemporal disappearance of Neanderthals in Iberia. Nat. Ecol. Evol. https://doi.org/10.1038/s41559-022-01917-6 (2022).Article 
PubMed 
PubMed Central 

Google Scholar 
Fu, Q. et al. An early modern human from Romania with a recent Neanderthal ancestor. Nature 524, 216–219 (2015).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Wood, R. E. et al. The chronology of the earliest Upper Palaeolithic in northern Iberia: New insights from L’Arbreda, Labeko Koba and La Viña. J. Hum. Evol. 69, 91–109 (2014).Article 
CAS 
PubMed 

Google Scholar 
Hublin, J. J. The modern human colonization of western Eurasia: When and where? Quat. Sci. Rev. 118, 194–210 (2015).Article 
ADS 

Google Scholar 
Hublin, J. J. The earliest modern human colonization of Europe. Proc. Natl. Acad. Sci. U.S.A. 109, 13471–13472 (2012).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Marín-Arroyo, A. B. & Sanz-Royo, A. What Neanderthals and AMH ate: Reassessment of the subsistence across the Middle-Upper Palaeolithic transition in the Vasco-Cantabrian region of SW Europe. J. Quat. Sci. 37, 320–334 (2022).Article 

Google Scholar 
Semal, P. et al. New data on the late Neandertals: Direct dating of the Belgian Spy fossils. Am. J. Phys. Anthropol. 138, 421–428 (2009).Article 
PubMed 

Google Scholar 
Welker, F. et al. Palaeoproteomic evidence identifies archaic hominins associated with the Châtelperronian at the Grotte du Renne. Proc. Natl. Acad. Sci. 113, 11162–11167 (2016).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Zilhão, J. Chronostratigraphy of the Middle-to-Upper Paleolithic transition in the Iberian Peninsula. Pyrenae Rev. Prehist. i Antig. la Mediter. Occident. 37, 7–84 (2006).
Google Scholar 
Vanhaeren, M. & d’Errico, F. Aurignacian ethno-linguistic geography of Europe revealed by personal ornaments. J. Archaeol. Sci. 33, 1105–1128 (2006).Article 

Google Scholar 
Higham, T. et al. Testing models for the beginnings of the Aurignacian and the advent of figurative art and music: The radiocarbon chronology of Geißenklösterle. J. Hum. Evol. 62, 664–676 (2012).Article 
PubMed 

Google Scholar 
Broglio, A. et al. L’art aurignacien dans la décoration de la Grotte de Fumane. Anthropologie 113, 753–761 (2009).Article 

Google Scholar 
Conard, N. J. Palaeolithic ivory sculptures from southwestern Germany and the origins of figurative art. Nature 426, 830–832 (2003).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Conard, N. J. A female figurine from the basal Aurignacian of Hohle Fels Cave in southwestern Germany. Nature 459, 248–252 (2009).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Bourrillon, R. et al. A new Aurignacian engraving from Abri Blanchard, France: Implications for understanding Aurignacian graphic expression in Western and Central Europe. Quat. Int. 491, 46–64 (2018).Article 

Google Scholar 
Tejero, J. M. & Grimaldi, S. Assessing bone and antler exploitation at Riparo Mochi (Balzi Rossi, Italy): Implications for the characterization of the Aurignacian in South-western Europe. J. Archaeol. Sci. 61, 59–77 (2015).Article 

Google Scholar 
Tejero, J. M. Spanish aurignacian projectile points: An example of the First European Paleolithic hunting weapons in osseous materials. In Osseous Projectile Weaponry Towards an Understanding of Pleistocene Cultural Variability (ed. Langley, M. C.) 55–70 (Springer, 2017).
Google Scholar 
Kitagawa, K. & Conard, N. J. Split-based points from the Swabian Jura highlight Aurignacian regional signatures. PLoS ONE 15(11), e0239865 (2020).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Floss, H., Hoyer, C. T., Heckel, C. & Tartar, É. The Aurignacian in Southern Burgundy. Palethnologie 7, 1–22 (2015).
Google Scholar 
Tartar, É. Origin and development of Aurignacian Osseous Technology in Western Europe: A review of current knowledge. Palethnologie 7, 1–20 (2015).
Google Scholar 
Bar-Yosef, O. Neanderthals and modern humans: A different interpretation. In When Neanderthals and Modern Humans Met (ed. Conard, N. J.) 467–482 (Kerns Verlag, 2006).
Google Scholar 
Flas, D. Les pointes foliacées et les changements techniques autour de la transitiondu Paléolithiquemoyen au supérieur dans leNord-Ouest de l’Europe. In (eds Toussaint, M. & Di Modica, K. S. P.) 261–276 (ERAUL 128, 2011).Nigst, P. R. The Early Upper Palaeolithic of the Middle Danube Region—Human Evolution (Leiden University, 2012).
Google Scholar 
Tsanova, T. Les débuts du Paléolithique supérieur dans l’Est des Balkans. Réflexion à partir de l’étude taphonomique et techno-économique des ensembles lithiques des sites de Bacho Kiro (couche 11), Temnata (couches VI et 4) et Kozarnika (niveau VII) (2008).Bosch, M. D. et al. New chronology for Ksâr ’Akil (Lebanon) supports Levantine route of modern human dispersal into Europe. Proc. Natl. Acad. Sci. U.S.A. 112, 7683–7688 (2015).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Chu, W. et al. Aurignacian dynamics in Southeastern Europe based on spatial analysis, sediment geochemistry, raw materials, lithic analysis, and use-wear from Românești-Dumbrăvița. Sci. Rep. 12, 14152 (2022).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Conard, N. J. The timing of cultural innovations and the dispersal of modern humans in Europe. Terra Nostra 6, 82–94 (2002).
Google Scholar 
Davies, W. Re-evaluating the Aurignacian as an Expression of Modern Human Mobility and Dispersal. (eds. Mellars P, Boyle K, Bar-Yosef O, S. C.) (2001).Mellars, P. A new radiocarbon revolution and the dispersal of modern humans in Eurasia. Nature 439, 931–935 (2006).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Mellars, P. Archeology and the dispersal of modern humans in Europe: Deconstructing the ‘Aurignacian’. Evol. Anthropol. 15, 167–182 (2006).Article 

Google Scholar 
Szmidt, C. C., Normand, C., Burr, G. S., Hodgins, G. W. L. & LaMotta, S. AMS 14C dating the Protoaurignacian/Early Aurignacian of Isturitz, France. Implications for Neanderthal-modern human interaction and the timing of technical and cultural innovations in Europe. J. Archaeol. Sci. 37, 758–768 (2010).Article 

Google Scholar 
Conard, N. J. & Bolus, M. Radiocarbon dating the appearance of modern humans and timing of cultural innovations in Europe: New results and new challenges. J. Hum. Evol. 44, 331–371 (2003).Article 
PubMed 

Google Scholar 
Douka, K., Grimaldi, S., Boschian, G., del Lucchese, A. & Higham, T. F. G. A new chronostratigraphic framework for the Upper Palaeolithic of Riparo Mochi (Italy). J. Hum. Evol. 62, 286–299 (2012).Article 
PubMed 

Google Scholar 
Davies, W. & Hedges, R. E. M. Dating a type site: Fitting Szeleta Cave into its regional chronometric context. Praehistoria 9–10, 35–45 (2009).
Google Scholar 
Davies, W., White, D., Lewis, M. & Stringer, C. Evaluating the transitional mosaic: Frameworks of change from Neanderthals to Homo sapiens in eastern Europe. Quat. Sci. Rev. 118, 211–242 (2015).Article 
ADS 

Google Scholar 
Marín-Arroyo, A. B. et al. Chronological reassessment of the Middle to Upper Paleolithic transition and Early Upper Paleolithic cultures in Cantabrian Spain. PLoS ONE 13, 1–20 (2018).
Google Scholar 
Barshay-Szmidt, C., Normand, C., Flas, D. & Soulier, M. C. Radiocarbon dating the Aurignacian sequence at Isturitz (France): Implications for the timing and development of the Protoaurignacian and Early Aurignacian in western Europe. J. Archaeol. Sci. Rep. 17, 809–838 (2018).
Google Scholar 
Wood, R. et al. El Castillo (Cantabria, northern Iberia) and the Transitional Aurignacian: Using radiocarbon dating to assess site taphonomy. Quat. Int. 474, 56–70 (2018).Article 

Google Scholar 
Chu, W. The danube corridor hypothesis and the carpathian basin: Geological, environmental and archaeological approaches to characterizing aurignacian dynamics. J. World Prehist. 31, 117–178 (2018).Article 

Google Scholar 
Falcucci, A., Conard, N. J. & Peresani, M. Breaking through the aquitaine frame: A re-evaluation on the significance of regional variants during the Aurignacian as seen from a key record in southern Europe. J. Anthropol. Sci. 98, 99–140 (2020).PubMed 

Google Scholar 
Banks, W. E., d’Errico, F. & Zilhão, J. Human-climate interaction during the Early Upper Paleolithic: Testing the hypothesis of an adaptive shift between the Proto-Aurignacian and the Early Aurignacian. J. Hum. Evol. 64, 39–55 (2013).Article 
PubMed 

Google Scholar 
Badino, F. et al. An overview of Alpine and Mediterranean palaeogeography, terrestrial ecosystems and climate history during MIS 3 with focus on the Middle to Upper Palaeolithic transition. Quat. Int. 551, 7–28 (2020).Article 

Google Scholar 
Bataille, G. & Conard, N. J. Blade and bladelet production at Hohle Fels Cave, AH IV in the Swabian Jura and its importance for characterizing the technological variability of the Aurignacian in Central Europe. PLoS ONE 13, 1–47 (2018).Article 

Google Scholar 
Higham, T., Wood, R., Moreau, L., Conard, N. & Ramsey, C. B. Comments on ‘Human-climate interaction during the early Upper Paleolithic: Testing the hypothesis of an adaptive shift between the Proto-Aurignacian and the Early Aurignacian’ by Banks et al.. J. Hum. Evol. 65, 806–809 (2013).Article 
PubMed 

Google Scholar 
Teyssandier, N. & Zilhão, J. On the entity and antiquity of the Aurignacian at Willendorf (Austria): Implications for modern human emergence in Europe. J. Paleolit. Archaeol. 1, 107–138 (2018).Article 

Google Scholar 
Discamps, E., Jaubert, J. & Bachellerie, F. Human choices and environmental constraints: Deciphering the variability of large game procurement from Mousterian to Aurignacian times (MIS 5–3) in southwestern France. Quat. Sci. Rev. 30, 2755–2775 (2011).Article 
ADS 

Google Scholar 
Kuhn, S. L. & Stiner, M. C. What’s a mother to do? The division of labor among Neandertals and modern humans in Eurasia. Curr. Anthropol. 47, 953–980 (2006).Article 

Google Scholar 
Starkovich, B. M. Intensification of small game resources at Klissoura Cave 1 (Peloponnese, Greece) from the Middle Paleolithic to Mesolithic. Quat. Int. 264, 17–31 (2012).Article 

Google Scholar 
Stiner, M. Prey choice, site occupation intensity & economic diversity in the Middle–early Upper Palaeolithic at the Üçağizli Caves. Turkey. Before Farm. 3, 1–20 (2009).
Google Scholar 
Yravedra Saínz de los Terreros, J., Gómez-Castanedo, A., Aramendi-Picado, J., Montes-Barquín, R. & Sanguino-González, J. Neanderthal and Homo sapiens subsistence strategies in the Cantabrian region of northern Spain. Archaeol. Anthropol. Sci. 8, 779–803 (2016).Article 

Google Scholar 
Bertacchi, A., Starkovich, B. M. & Conard, N. J. The Zooarchaeology of Sirgenstein Cave: A Middle and Upper Paleolithic site in the Swabian Jura, SW Germany. J. Paleolit. Archaeol. 4, 7 (2021).Article 

Google Scholar 
Boscato, P. & Crezzini, J. Middle-Upper Palaeolithic transition in Southern Italy: Uluzzian macromammals from Grotta del Cavallo (Apulia). Quat. Int. 252, 90–98 (2012).Article 

Google Scholar 
Grayson, D. K. & Delpech, F. The large mammals of Roc de Combe (Lot, France): The Châtelperronian and Aurignacian assemblages. J. Anthropol. Archaeol. 27, 338–362 (2008).Article 

Google Scholar 
Morin, E. et al. New evidence of broader diets for archaic Homo populations in the northwestern Mediterranean. Sci. Adv. 5, 1–12 (2019).Article 

Google Scholar 
Münzel, S. & Conard, N. J. Change and continuity in subsistence during the Middle and Upper Palaeolithic in the Ach Valley of Swabia (South-west Germany). Int. J. Osteoarchaeol. 14, 225–243 (2004).Article 

Google Scholar 
Rendu, W. et al. Subsistence strategy changes during the Middle to Upper Paleolithic transition reveals specific adaptations of Human Populations to their environment. Sci. Rep. 9, 1–11 (2019).Article 
ADS 

Google Scholar 
Romandini, M. et al. Macromammal and bird assemblages across the late Middle to Upper Palaeolithic transition in Italy: An extended zooarchaeological review. Quat. Int. 551, 188–223 (2020).Article 

Google Scholar 
Starkovich, B. M. Paleolithic subsistence strategies and changes in site use at Klissoura Cave 1 (Peloponnese, Greece). J. Hum. Evol. 111, 63–84 (2017).Article 
PubMed 

Google Scholar 
Peresani, M. Inspecting human evolution from a cave Late Neanderthals and early sapiens at Grotta di Fumane: Present state and outlook. J. Anthropol. Sci. 100, 71–107 (2022).PubMed 

Google Scholar 
Jarvis, A. et al. Hole-Filled Seamless SRTM Data V4. https://srtm.csi.cgiar.org (International Centre for Tropical Agriculture (CIAT), 2008).Delpiano, D., Heasley, K. & Peresani, M. Assessing Neanderthal land use and lithic raw material management in discoid technology. J. Anthropol. Sci. 96, 89–110 (2018).PubMed 

Google Scholar 
Marcazzan, D., Ligouis, B., Duches, R. & Conard, N. J. Middle and Upper Paleolithic occupations of Fumane Cave (Italy): A geoarchaeological investigation of the anthropogenic features. J. Antropol. Sci. 100, 1–26 (2022).
Google Scholar 
Douka, K. et al. On the chronology of the Uluzzian. J. Hum. Evol. 68, 1–13 (2014).Article 
PubMed 

Google Scholar 
Peresani, M., Cristiani, E. & Romandini, M. The Uluzzian technology of Grotta di Fumane and its implication for reconstructing cultural dynamics in the Middle-Upper Palaeolithic transition of Western Eurasia. J. Hum. Evol. 91, 36–56 (2016).Article 
PubMed 

Google Scholar 
Peresani, M., Bertola, S., Delpiano, D., Benazzi, S. & Romandini, M. The Uluzzian in the north of Italy: Insights around the new evidence at Riparo Broion. Archaeol. Anthropol. Sci. 11, 3503–3536 (2019).Article 

Google Scholar 
Aleo, A., Duches, R., Falcucci, A., Rots, V. & Peresani, M. Scraping hide in the early Upper Paleolithic: Insights into the life and function of the Protoaurignacian endscrapers at Fumane Cave. Archaeol. Anthropol. Sci. 13, 1 (2021).Article 

Google Scholar 
Falcucci, A., Conard, N. J. & Peresani, M. A critical assessment of the Protoaurignacian lithic technology at Fumane Cave and its implications for the definition of the earliest Aurignacian. PLoS ONE 12, 1–43 (2017).Article 

Google Scholar 
Falcucci, A. & Peresani, M. A pre-Heinrich Event 3 assemblage at Fumane Cave and its contribution for understanding the beginning of the Gravettian in Italy. Quartär 66, 135–154 (2019).
Google Scholar 
Higham, T. European Middle and Upper Palaeolithic radiocarbon dates are often older than they look. Antiquity 85, 235–249 (2011).Article 

Google Scholar 
Higham, T. et al. Problems with radiocarbon dating the Middle to Upper Palaeolithic transition in Italy. Quat. Sci. Rev. 28, 1257–1267 (2009).Article 
ADS 

Google Scholar 
Cassoli, P. F. & Tagliacozzo, A. Considerazioni paleontologiche, paleoecologiche e archeologiche sui micromammiferi e gli uccelli dei livelli del Pleistocene Superiore del Riparo di Fumane (Vr) (Scavi 1988–91). Bollettino del Museo Civico di Storia Naturale di Verona 18, 349–445 (1994).
Google Scholar 
Broglio, A., De Stefani, M., Tagliacozzo, A., Gurioli, F. & Facciolo, A. Aurignacian dwelling structures, hunting strategies and seasonality in the Fumane Cave (Lessini Mountains). In Kostenki and the Early Upper Paleolithic of Eurasia: General Trends, Local Developments (eds Vasilev, S. A. et al.) 263–268 (Nestor-Historia, 2006).
Google Scholar 
Bertola, S. et al. Le territoire des chasseurs aurignaciens dans les Préalpes de la Vénétie: l’exemple de la Grotte de Fumane. In Le Concept de Territoires dans le Paléolithique Supérieur Européen (eds Djindjian, F. et al.) (BAR Intemational Series, 2009).
Google Scholar 
Jéquier, C., Livraghi, A., Romandini, M. & Peresani, M. Same but different: 20,000 years of bone retouchers from northern Italy. A diachronologic approach from neanderthals to anatomically modern humans. In The Origins of Bone Tool Technologies (eds Hutson, J. M. et al.) 269–285 (Verlag des Römisch-Germanischen Zentralmuseums, 2018).
Google Scholar 
Marín-Arroyo, A. B. A comparative study of analytic techniques for skeletal part profile interpretation at El Mirón Cave (Cantabria, Spain). Archaeofauna 18, 79–98 (2009).
Google Scholar 
Romandini, M. Analisi archeozoologica, tafonomica, paleontologica e spaziale dei livelli Uluzziani e tardo-Musteriani della Grotta di Fumane (VR). Variazioni e continuità strategico-comportamentali umane in Italia Nord Occidentale: i casi di Grotta del Col della Stria. Dip. di Biol. ed Evol. PhD thesis 505 (2012).Marean, C. W., Abe, Y., Nilssen, P. J. & Stone, E. C. Estimating the minimum number of skeletal elements (MNE) in zooarchaeology: A review and a new image-analysis GIS approach. Am. Antiq. 66, 333–348 (2001).Article 
CAS 
PubMed 

Google Scholar 
Stiner, M. C. The use of mortality patterns in archaeological studies of hominid predatory adaptations. J. Anthropol. Archaeol. 9, 305–351 (1990).Article 

Google Scholar 
Marín-Arroyo, A. B. & Morales, M. R. G. Comportamiento económico de los últimos cazadores-recolectores y primeras evidencias de domesticación en el occidente de asturias. La cueva de mazaculos II. Trab. Prehist. 66, 47–74 (2009).Article 

Google Scholar 
Simpson, E. H. Measurement of diversity. Nature 163, 688 (1949).Article 
ADS 
MATH 

Google Scholar 
Magurran, A. E. Ecological Diversity and Its Measurement (Princeton University Press, 1988).Book 

Google Scholar 
Marín-Arroyo, A. B. The use of optimal foraging theory to estimate Late Glacial site catchment areas from a central place: The case of eastern Cantabria, Spain. J. Anthropol. Archaeol. 28, 27–36 (2009).Article 

Google Scholar 
Azorit, C. Guía para la determinación de la edad del ciervo ibérico (Cervus elaphus hispanicus) a través de su dentición: Revisión metodológica y técnicas de elección. An. la Real Acad. Ciencias Vet. Andalucía Orient. 24, 235–264 (2011).
Google Scholar 
Mariezkurrena, K. Contribución al conocimiento del desarrollo de la dentición y el esqueleto poscraneal de Cervus elaphus. Munibe 35, 149–202 (1983).
Google Scholar 
Tomé, C. & Vigne, J. D. Roe deer (Capreolus capreolus) age at death estimates: New methods and modern reference data for tooth eruption and wear, and for epiphyseal fusion. Archaeofauna 12, 157–173 (2003).
Google Scholar 
Couturier, M. A. J. Le bouquetin des Alpes: Capra aegagrus ibex ibex L. (Impr. Allier, 1962).
Google Scholar 
Habermehl, K.-H. Die Altersbeurteilung beim weiblichen Steinwild (Capra ibex ibex L.) anhand der Skelettentwicklung. Anat. Histol. Embryol. J. Vet. Med. Ser. C 21, 193–198 (1992).Article 
CAS 

Google Scholar 
Pflieger, R. H. P. Le chamois, son identification et sa vie (Grand Gibier, 1982).
Google Scholar 
Binford, L. R. Nunamiut Etnoarchaeology (Academic Press, 1978).
Google Scholar 
Metcalfe, D. & Jones, K. T. A reconsideration of animal body-part utility indices. Am. Antiq. 53, 486–504 (1988).Article 

Google Scholar 
Morin, E. & Ready, E. Foraging goals and transport decisions in western Europe during the Paleolithic and Early Holocene. In Zooarchaeology and Modern Human Origins. Vertebrate Paleobiology and Paleoanthropology (eds Clark, J. L. & Speth, J. D.) 227–269 (Springer, 2013).
Google Scholar 
Lam, Y. M., Chen, X. & Pearson, O. M. Intertaxonomic variability in patterns of bone density and the differential representation of bovid, cervid, and equid elements in the archaeological record. Am. Antiq. 64, 343–362 (1999).Article 

Google Scholar 
Morin, E. Fat composition and Nunamiut decision-making: A new look at the marrow and bone grease indices. J. Archaeol. Sci. 34, 69–82 (2007).Article 

Google Scholar 
Marín-Arroyo, A. B. & Ocio, D. Disentangling faunal skeletal profiles. A new probabilistic framework. Hist. Biol. 30, 720–729 (2018).Article 

Google Scholar 
Rogers, A. R. On the value of soft bones in faunal analysis. J. Archaeol. Sci. 27, 635–639 (2000).Article 

Google Scholar 
Rogers, A. R. Analysis of bone counts by maximum likelihood. J. Archaeol. Sci. 27, 111–125 (2000).Article 

Google Scholar 
Marín-Arroyo, A. B., Ocio, D., Vidal-Cordasco, M. & Vettese, D. BaSkePro: Bayesian Model to Archaeological Faunal Skeletal Profiles. R package version 0.1.0. https://CRAN.R-project.org/package=BaSkePro (2022).Binford, L. R. Bones Ancient Men and Modern Myths (Bones (Elsevier, 1981).
Google Scholar 
Galán, A. B. & Domínguez-Rodrigo, M. An experimental study of the anatomical distribution of cut marks created by filleting and disarticulation on long bone ends. Archaeometry 55, 1132–1149 (2013).Article 

Google Scholar 
Nilssen, P. J. An Actualistic Butchery Study in South Africa and Its Implications for Reconstructing Hominid Strategies of Carcass Acquisition and Butchery in the Upper Pleistocene and Plio-Pleistocene (University of Cape Town, 2000).
Google Scholar 
Capaldo, S. D. & Blumenschine, R. J. A quantitative diagnosis of notches made by hammerstone percussion and carnivore gnawing on bovid long bones. Am. Antiq. 59, 724–748 (1994).Article 

Google Scholar 
Pickering, T. R. & Egeland, C. P. Experimental patterns of hammerstone percussion damage on bones: Implications for inferences of carcass processing by humans. J. Archaeol. Sci. 33, 459–469 (2006).Article 

Google Scholar 
Bunn, H. T. Archaeological evidence for meat-eating by Plio-Pleistocene hominids from Koobi Fora and Olduvai Gorge. Nature 291, 574–577 (1981).Article 
ADS 

Google Scholar 
Villa, P. & Mahieu, E. Breakage patterns of human long bones. J. Hum. Evol. 21, 27–48 (1991).Article 

Google Scholar 
Blumenschine, R. J. & Selvaggio, M. M. Percussion marks on bone surfaces as a new diagnostic of hominid behaviour. Nature 333, 763–765 (1988).Article 
ADS 

Google Scholar 
Galán, A. B., Rodríguez, M., de Juana, S. & Domínguez-Rodrigo, M. A new experimental study on percussion marks and notches and their bearing on the interpretation of hammerstone-broken faunal assemblages. J. Archaeol. Sci. 36, 776–784 (2009).Article 

Google Scholar 
Pickering, T. R. et al. Taphonomy of ungulate ribs and the consumption of meat and bone by 1.2-million-year-old hominins at Olduvai Gorge, Tanzania. J. Archaeol. Sci. 40, 1295–1309 (2013).Article 

Google Scholar 
Vettese, D. et al. Towards an understanding of hominin marrow extraction strategies: A proposal for a percussion mark terminology. Archaeol. Anthropol. Sci. 12, 8 (2020).Article 

Google Scholar 
Vettese, D. et al. A way to break bones? The weight of intuitiveness. PLoS ONE 16, 0259136 (2021).Article 

Google Scholar 
Coil, R., Yezzi-Woodley, K. & Tappen, M. Comparisons of impact flakes derived from hyena and hammerstone long bone breakage. J. Archaeol. Sci. 120, 105167 (2020).Article 

Google Scholar 
Stiner, M. C., Kuhn, S. L., Weiner, S. & Bar-Yosef, O. Differential burning, recrystallization, and fragmentation of archaeological bone. J. Archaeol. Sci. 22, 223–237 (1995).Article 

Google Scholar 
Mallye, J. B. et al. The Mousterian bone retouchers of Noisetier Cave: Experimentation and identification of marks. J. Archaeol. Sci. 39, 1131–1142 (2012).Article 

Google Scholar 
Blumenschine, R. J. Percussion marks, tooth marks, and experimental determinations of the timing of hominid and carnivore access to long bones at FLK Zinjanthropus, Olduvai Gorge, Tanzania. J. Hum. Evol. 29, 21–51 (1995).Article 

Google Scholar 
Domínguez-Rodrigo, M. & Piqueras, A. The use of tooth pits to identify carnivore taxa in tooth-marked archaeofaunas and their relevance to reconstruct hominid carcass processing behaviours. J. Archaeol. Sci. 30, 1385–1391 (2003).Article 

Google Scholar 
Domínguez-Rodrigo, M. & Barba, R. New estimates of tooth mark and percussion mark frequencies at the FLK Zinj site: The carnivore-hominid-carnivore hypothesis falsified. J. Hum. Evol. 50, 170–194 (2006).Article 
PubMed 

Google Scholar 
Behrensmeyer, A. K. Taphonomic and écologie information from bone weathering. Paleobiology 4, 150–162 (1978).Article 

Google Scholar 
Fisher, J. W. Bone surface modifications in zooarchaeology. J. Archaeol. Method Theory 2, 7–68 (1995).Article 

Google Scholar 
Lyman, R. L. Vertebrate Taphonomy (Cambridge University Press, 1994).Book 

Google Scholar 
Shipman, P. Life History of a Fossil: An Introduction to Taphonomy and Paleoecology (Harvard University Press, 1981).
Google Scholar 
Marín-Arroyo, A. B. et al. Archaeological implications of human-derived manganese coatings: A study of blackened bones in El Mirón Cave, Cantabrian Spain. J. Archaeol. Sci. 35, 801–813 (2008).Article 

Google Scholar 
Marín-Arroyo, A. B., Landete-Ruiz, M. D., Seva-Román, R. & Lewis, M. D. Manganese coating of the Tabun faunal assemblage: Implications for modern human behaviour in the Levantine Middle Palaeolithic. Quat. Int. 330, 10–18 (2014).Article 

Google Scholar 
Blasco, R., Rosell, J., Fernández Peris, J., Cáceres, I. & Vergès, J. M. A new element of trampling: An experimental application on the Level XII faunal record of Bolomor Cave (Valencia, Spain). J. Archaeol. Sci. 35, 1605–1618 (2008).Article 

Google Scholar 
Domínguez-Rodrigo, M., de Juana, S., Galán, A. B. & Rodríguez, M. A new protocol to differentiate trampling marks from butchery cut marks. J. Archaeol. Sci. 36, 2643–2654 (2009).Article 

Google Scholar 
Hickler, T., Prentice, I. C., Smith, B., Sykes, M. T. & Zaehle, S. Implementing plant hydraulic architecture within the LPJ dynamic global vegetation model. Glob. Ecol. Biogeogr. 15, 567–577 (2006).Article 

Google Scholar 
Zampieri, D. Segmentation and linkage of the Lessini Mountains normal faults, Southern Alps, Italy. Tectonophysics 319, 19–31 (2000).Article 
ADS 

Google Scholar 
Castiglioni, G. B. et al. The loess deposits in the Lessini plateau. In The Loess in Northern and Central Italy: A Loess Basin Between the Alps and the Mediterranean Region (ed. Cremaschi, M.) 41–59 (Centro di Studio per la Stratigrafia e Petrografia delle Alpi Centrali, 1990).
Google Scholar 
Sauro, U. Il paesaggio degli alti Lessini. Studio Geomofologico (Museo Civico di Storia Naturale, 1973).
Google Scholar 
Castiglioni, G. B. et al. Geomorphological Map of Po Plain, Scale 1:250,000 (1997).Fontana, A., Mozzi, P. & Marchetti, M. Alluvial fans and megafans along the southern side of the Alps. Sediment. Geol. 301, 150–171 (2014).Article 
ADS 

Google Scholar 
Holechek, J. L., Pieper, R. D. & Herbel, C. H. Range Management Principles and Practices 3rd edn. (Prentice-Hall, 1998).
Google Scholar 
Imhoff, M. L. et al. Global patterns in human consumption of net primary production. Nature 429, 870–873 (2004).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Sitch, S. et al. Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model. Glob. Change Biol. 9, 161–185 (2003).Article 
ADS 

Google Scholar 
Smith, B. et al. Implications of incorporating N cycling and N limitations on primary production in an individual-based dynamic vegetation model. Biogeosciences 11, 2027–2054 (2014).Article 
ADS 

Google Scholar 
Githumbi, E. N. et al. Pollen, people and place: Multidisciplinary perspectives on ecosystem change at Amboseli, Kenya. Front. Earth Sci. 5, 113 (2018).Article 
ADS 

Google Scholar 
Allen, J. R. M. et al. Global vegetation patterns of the past 140,000 years. J. Biogeogr. 47, 2073–2090 (2020).Article 

Google Scholar 
Armstrong, E., Hopcroft, P. O. & Valdes, P. J. A simulated Northern Hemisphere terrestrial climate dataset for the past 60,000 years. Sci. Data 6, 1–16 (2019).Article 

Google Scholar 
Harris, I., Osborn, T. J., Jones, P. & Lister, D. Version 4 of the CRU TS monthly high-resolution gridded multivariate climate dataset. Sci. Data 7, 1–18 (2020).Article 

Google Scholar 
Adam, M., Weitzel, N. & Rehfeld, K. Identifying global-scale patterns of vegetation change during the last deglaciation from paleoclimate networks. Paleoceanogr. Paleoclimatol. 36, 1 (2021).Article 

Google Scholar 
Beyer, R., Krapp, M. & Manica, A. An empirical evaluation of bias correction methods for palaeoclimate simulations. Clim. Past 16, 1493–1508 (2020).Article 

Google Scholar 
Lüthi, D. et al. High-resolution carbon dioxide concentration record 650,000–800,000 years before present. Nature 453, 379–382 (2008).Article 
ADS 
PubMed 

Google Scholar 
Zobler, L. A World Soil File for Global Climate Modelling. NASA Technical Memorandum 87802 (1986).Bronk Ramsey, C. Bayesian analysis of radiocarbon dates. Radiocarbon 51, 337–360 (2009).Article 

Google Scholar 
Reimer, P. J. et al. The IntCal20 northern hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon 62, 725–757 (2020).Article 
CAS 

Google Scholar 
Andersen, K. K. et al. The Greenland ice core chronology 2005, 15–42 ka. Part 1: Constructing the time scale. Quat. Sci. Rev. 25, 3246–3257 (2006).Article 
ADS 

Google Scholar 
Svensson, A. et al. A 60,000 year Greenland stratigraphic ice core chronology. Clim. Past 4, 47–57 (2008).Article 

Google Scholar 
Rasmussen, S. O. et al. A stratigraphic framework for abrupt climatic changes during the Last Glacial period based on three synchronized Greenland ice-core records: Refining and extending the INTIMATE event stratigraphy. Quat. Sci. Rev. 106, 14–28 (2014).Article 
ADS 

Google Scholar 
Higham, T. et al. The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512, 306–309 (2014).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Romandini, M., Nannini, N., Tagliacozzo, A. & Peresani, M. The ungulate assemblage from layer A9 at Grotta di Fumane, Italy: A zooarchaeological contribution to the reconstruction of Neanderthal ecology. Quat. Int. 337, 11–27 (2014).Article 

Google Scholar 
Tagliacozzo, A., Romandini, M., Fiore, I., Gala, M. & Peresani, M. 2013. Animal exploitation strategies during the Uluzzian at Grotta di Fumane (Verona, Italy). In Zooarchaeology and Modern Human Origins. Vertebrate Paleobiology and Paleoanthropology (eds Clark, J. & Speth, J.) 129–150 (Springer, 2013).
Google Scholar 
Terlato, G., Livraghi, A., Romandini, M. & Peresani, M. Large bovids on the Neanderthal menu: Exploitation of Bison priscus and Bos primigenius in northeastern Italy. J. Archaeol. Sci. Rep. 25, 129–143 (2019).
Google Scholar 
Peresani, M., Fiore, I., Gala, M., Romandini, M. & Tagliacozzo, A. Late Neandertals and the intentional removal of feathers as evidenced from bird bone taphonomy at Fumane Cave 44 ky B.P., Italy. Proc. Natl. Acad. Sci. U.S.A. 108, 3888–3893 (2011).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Fiore, I. et al. From feathers to food: Reconstructing the complete exploitation of avifaunal resources by Neanderthals at Fumane cave, unit A9. Quat. Int. 421, 134–153 (2016).Article 

Google Scholar 
Romandini, M. et al. Neanderthal scraping and manual handling of raptors wing bones: Evidence from Fumane Cave. Experimental activities and comparison. Quat. Int. 421, 154–172 (2016).Article 

Google Scholar 
López-García, J. M., dalla Valle, C., Cremaschi, M. & Peresani, M. Reconstruction of the Neanderthal and Modern Human landscape and climate from the Fumane cave sequence (Verona, Italy) using small-mammal assemblages. Quat. Sci. Rev. 128, 1–13 (2015).Article 
ADS 

Google Scholar 
Maspero, A. Ricostruzione del paesaggio vegetale attorno alla grotta di Fumane durante il Paleolitico. Annu. Stor. della Valpolicella 18, 19–26 (1998).
Google Scholar 
Malerba, G. & Giacobini, G. Analisi delle tracce di macellazione in un sito Paleolitico. L’esempio del Riparo di Fumane (Valpolicella, Verona). In Atti del: “I° Convegno Nazionale di Archeozoologia”, Rovigo 5–7 marzo 1993 97–108 (1995).Fritz, S. A. et al. Twenty-million-year relationship between mammalian diversity and primary productivity. Proc. Natl. Acad. Sci. 113, 10908–10913 (2016).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Morris, R. J. Community ecology: How green is the arctic Tundra? Curr. Biol. 18, R256–R258 (2008).Article 
CAS 
PubMed 

Google Scholar 
Holt, B. et al. The Middle-Upper Paleolithic transition in Northwest Italy: New evidence from Riparo Bombrini (Balzi Rossi, Liguria, Italy). Quat. Int. 508, 142–152 (2019).Article 

Google Scholar 
Pothier Bouchard, G., Riel-Salvatore, J., Negrino, F. & Buckley, M. Archaeozoological, taphonomic and ZooMS insights into the Protoaurignacian faunal record from Riparo Bombrini. Quat. Int. https://doi.org/10.1016/j.quaint.2020.01.007 (2020).Article 

Google Scholar 
Riel-Salvatore, J. & Negrino, F. Proto-Aurignacian Lithic technology, mobility, and human niche construction: A case study from Riparo Bombrini, Italy. In Lithic Technological Organization and Paleoenvironmental Change, Studies in Human Ecology and Adaptation (eds Robinson, E. & Sellet, F.) (Springer, 2018).
Google Scholar 
Riel-Salvatore, J. & Negrino, F. Human adaptations to climatic change in Liguria across the Middle-Upper Paleolithic transition. J. Quat. Sci. 33, 313–322 (2018).Article 

Google Scholar 
Grimaldi, S., Porraz, G. & Santaniello, F. Raw material procurement and land use in the northern Mediterranean Arc: Insight from the first Proto-Aurignacian of Riparo Mochi (Balzi Rossi, Italy). Quartar 61, 113–127 (2014).
Google Scholar 
Alaique, F. Risultati preliminari dell’analisi dei resti faunistici rinvenuti nei livelli del Paleolitico superiore di Riparo Mochi (Balzi Rossi): Scavi 1995–1996. In Atti Del 2°Convegno Nazionale Di Archeozoologia (Asti, 1997) (eds Malerba, G. et al.) 125–130 (Abaco Edizioni, 2000).
Google Scholar 
Staubwasser, M. et al. Impact of climate change on the transition of Neanderthals to modern humans in Europe. Proc. Natl. Acad. Sci. 115, 9116–9121 (2018).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Columbu, A. et al. Speleothem record attests to stable environmental conditions during Neanderthal–modern human turnover in southern Italy. Nat. Ecol. Evol. 4, 1188–1195 (2020).Article 
PubMed 

Google Scholar 
Müller, U. C. et al. The role of climate in the spread of modern humans into Europe. Quat. Sci. Rev. 30, 273–279 (2011).Article 
ADS 

Google Scholar  More

Brazil’s new government aims to achieve zero deforestation in the Amazon rainforest. This welcome initiative should be extended to the Cerrado, the world’s most biodiverse woodland savannah.
Competing Interests
The authors declare no competing interests. More

The upside-down jellyfish, Cassiopea sp. produces several hydrodynamic effects capable of altering the ecosystem which it inhabits. Not only do Cassiopea produce feeding currents capable of turning over the water column above them several times per hour3, they are also capable of releasing interstitial porewater from the benthos5. The rate of porewater release, on the order of mL h−13, is capable of increasing water column NH4 levels by almost 30% under certain conditions3. In this study, we investigated two hypothetical mechanisms for this porewater release, and found that a combination of the morphology of the bell and the pulsing behavior of the jellyfish was responsible for releasing porewater from directly below the bell via a suction-pumping mechanism.The Bernoulli hypothesis4, a low-pressure zone surrounding the animal due to a velocity gradient between the substrate boundary and the incurrent flow of the Cassiopea sp. feeding current, predicted porewater release from the substrate surface surrounding the perimeter of the animal. While porewater is entrained from the perimeter of the bell into the feeding current4 lateral expulsion of porewater due to the suction pump mechanism would produce a visually similar flow of porewater. A horizontal flow of water does occur near the bottom1, but this flow is restricted to a narrow region near the bell and velocities were low compared to the vertical excurrent jet (Fig. 4). To test the effect of Bernoulli’s principle, we measured the effect on porewater release rates of an impermeable ring-shaped barrier surrounding the animal in order to inhibit benthic-pelagic fluid flux other than directly under the animal (Fig. 2A) using labeled fluorescein per the methods of Durieux et al.3, which were adapted from those of Jantzen et al.5 (Fig. 2). If the Bernoulli mechanism contributed to porewater liberation this treatment should have reduced the porewater release rate, but the release rates observed were not significantly different from the control treatment (2.23 mL h−1 ± 1.27 s.d., Fig. 2D).The suction pumping hypothesis5, a mechanism using the exumbrellar cavity as a suction pump that draws porewater vertically upward beneath the bell and then expels it laterally, would expect to see the majority of porewater released from directly under the bell of Cassiopea sp. This mechanism is supported by bell morphology5 and the appearance of deep porewater at the benthic surface of the exumbrellar cavity5. In our, an impermeable disk was placed underneath the animal to obstruct the flow predicted by the suction pump hypothesis (Fig. 2B). Additionally, we made a 6 mm perforation in the bells of the jellyfish to interfere with the ability to form the sub-ambient pressure in the exumbrellar space necessary for suction pumping to occur (Fig. 2C). Both treatments resulted in a significant decrease in porewater liberation, with flows indistinguishable from the absence of any animal (Fig. 2D), supporting the suction-pumping hypothesis.Since the suction pumping mechanism requires pressure fluctuations in the exumbrellar space, we also directly measured the water pressure below the jellyfish. The initiation of the power stroke of bell pulsation coincides with a sudden decrease in water pressure in the exumbrellar space (Fig. 3A,B) of a mean magnitude of 43.4 Pa (± 13.6 s.d.). These pressure fluctuations appear to be unaffected by animal size (Fig. 3D,E), although the rate of porewater release is known to scale with bell diameter3. Note that the muscles responsible for bell contraction in Cassiopea sp. are roughly 2-dimensional sheets13 with a thickness of one cell14 and therefore the cross-sectional area also does not scale with diameter. Our experiments were performed on smooth acrylic rather than sand, so that the conditions here were optimal for the formation of a tight seal with the bottom. However, the magnitude of this difference is likely to be small, as Cassiopea sp. produce copious amounts of mucus, which can compensate for small-scale surface roughness. In addition, the duration of each individual bell pulse is short1, so given the fine pore size of a sand or mud substrate, it is unlikely that subambient pressure would have the opportunity to dissipate enough to affect the high suction impulse produced.While not statistically significant, bell perforation did lead to data suggesting a decrease in exumbrellar pressure fluctuations (Fig. 3C), which could explain the reduction in porewater release observed (Fig. 2C). The fact that some pressure fluctuation was seen despite a complete lack of porewater release suggests that a minimum magnitude of pressure fluctuation might be necessary for suction pumping to occur. Furthermore, the effect may have been reduced by the ability of injured Cassiopea to produce copious amounts of mucus, which could have acted to minimize the impact of bell perforation. These parallel lines of reasoning firmly suggest that suction-pumping is, in fact, the dominant mechanism by which Cassiopea sp. release porewater.The suction-pumping mechanism for the release of porewater has broad-ranging ecological implications. Release rates should increase additively with population density, and the rate of bell pulsation should correlate with the rate of porewater liberation. The additive relationship to population density is important, since Cassiopea can occur at high densities of up to 100 animals m−23. Furthermore, while the Bernoulli mechanism predicted that interstitial water movement would be limited to the upper layers of the benthos, the suction pump mechanism has the potential to release porewater from deeper sediment strata. This deep flushing should expand the oxygen penetration depth downward, affecting factors such as respiration and sediment stability15. Given the fact that Cassiopea are capable of moving along the substrate5,16 this also means that the oxygen penetration depth is likely to fluctuate over time, favoring organisms that are able to adapt their metabolism or are able to relocate themselves17.Given that porewater at the field site in Long Key, Florida, from which the animals in this study were collected, has mean ammonium concentrations of 72 μM, 160 times higher than the surrounding water column11, any benthic-pelagic coupling mechanisms in this habitat could alter nitrogen dynamics, especially given the fact that many marine primary producers preferentially take up ammonium, the most reduced state of nitrogen available, as a nitrogen source18. Cassiopea sp. animal size and population densities are known to correlate with anthropogenic disturbances, and it is suggested that this is due to an increase in nutrient availability in these areas6. In addition to prey capture, Cassiopea sp. could be supplementing their nitrogen demand through the release of nutrient-rich interstitial porewater, from which Cassiopea sp. can directly absorb ammonium and other nutrients such as phosphate and trace metals5. In fact, jellyfish presence significantly reduced porewater ammonium levels near the animal5, suggesting that nutrient-rich porewater was replaced by down-welling low-nutrient surface water. The observed benthic locomotion of Cassiopea5,16 may be a mechanism to avoid remaining in locations where they have depleted this nutrient resource3. It has been reported that Cassiopea sp. affect benthic nutrient transport on a more general level, increasing ammonium uptake and decreasing nitrate uptake of the bottom sediments19. Water column nutrient levels also varied significantly between presence and absence of Cassiopea sp., and also between light and dark treatments in the presence of Cassiopea sp.20. The addition of jellyfish increased the efflux of ammonium from the benthos during the dark treatments, but reduced ammonium concentrations in the water column during light treatments20. It is entirely possible that absorption of nutrients by Cassiopea sp.5 in order to meet daytime metabolic demand resulted in the animals reducing water column ammonium concentrations in these experiments20.In addition, Cassiopea sp. have been shown to increase spatial heterogeneity of interstitial oxygen and nutrient fluxes20, making it comparable to other biogenic processes like bioturbation. Bioturbation typically oxygenates the upper layers of substrate, increasing the nitrification zone21, and also increases 3-dimensional heterogeneity of oxygen and nutrient concentrations, allowing for more complex nutrient dynamics21. The transport of interstitial porewater from specific regions under individual jellyfish could well produce a similar effect. The porewater release rates can also be compared to that of abiotic processes, such as wind-wave driven flow over sediment wave ripples, which have been shown to liberate porewater at rates of up to 140 L m−2 day−1, or three orders of magnitude greater than diffusion alone, on shallow, exposed coastlines such as beaches22. Environmental mixing would be lower in the sheltered mangrove habitats where Cassiopea sp. are found, since at our study site wind wave height was reduced from 5.4 cm in the bay to 0.07 cm in the mangroves3. In these coastal habitats, the sediment often acts as a nutrient sink, causing certain nutrients to become limiting to primary producers. Some fringe mangrove forests along coastlines in both Florida and Belize have been shown to be N-limited, for example23,24. If these nutrients are then released back into the water column, they potentially increase primary productivity in the system occupied by Cassiopea sp. Depending on the system, this could either increase production or cause eutrophication, potentially altering productivity on a local or regional scale, as has been observed when nutrients are released from the benthos by winds25 or bioturbation26.The mechanics of suction-pumping also imply that interstitial porewater release rate will correlate with bell pulse rate. Pulse rate correlates with water temperature (Fig. 5B), which would suggest that Cassiopea sp. can release greater quantities of nutrient-rich porewater during the summer months. This was confirmed by a recent study on the related species, Cassiopea medusa from Lake Macquarie, Australia8. By extension, our model suggests that pulsing, and therefore porewater release, should cease entirely below 18ºC. In fact, at our site in Lido Key, population densities of Cassiopea sp. declined rapidly once water temperatures dropped this low (Fig. 6). This same temperature of 18 °C was determined independently to be the threshold at which Cassiopea sp. polyp feeding was inhibited10. As such, it is likely that winter minimum temperatures of 18ºC represent a limiting condition on Cassiopea sp. range expansion. Studies on Cassiopea medusa, suggested thermal stress and bell degradation at 16 °C8. As global climates warm, we can expect both a poleward shift of Cassiopea sp. Range9,27 and an increase in transport rates of porewater and its associated benthic nutrients throughout this range, leading to increased productivity, and potentially exacerbating eutrophication in some regions.We determined that a suction-pumping mechanism is responsible for the interstitial porewater release by Cassiopea, suggesting that release rates are independent of population density, but affected by pulse rate. The potential role of bell pulse rate was investigated further, and we found correlations between bell pulse rate and both animal size and water temperature. As a result, we expect that porewater liberation would demonstrate seasonal variations, with lower rates during the winter and reaching a maximum during the summer months. Cassiopea are able to release nutrient-rich porewater in the shallow quiescent habitats they inhabit, and through their feeding current mix these nutrients throughout the water column. Since this effect varies seasonally, it is likely that further study will show that these jellyfish are responsible for a complex system of nutrient dynamics in their ecosystem. More

We investigated the complex relationships between the environmental characteristics of blue spaces and visit-related mental well-being in a multi-country study including 17 bluespace types and four facets of subjective well-being. Our aim was to operationalise, and consider the utility of, the Bratman et al.9 conceptual model that links ecosystem services (ESS) with mental health.Consistent with the proposed conceptual model, mental well-being outcomes relied on a complex interplay of individual, environmental, and visit characteristics.Summary of findingsOverall, bluespace visits were associated with better subjective mental well-being outcomes if the visits took place in nearby coastal areas or rural rivers, were perceived as safe and to have good water quality, and had a long duration. They could involve a range of activities such as playing with children, socialising, or walking. The degree to which the perceived presence of wildlife predicted visit satisfaction varied depending on the bluespace type, suggesting that the importance of ecosystem features such as biodiversity may vary by the setting.We can also identify the combination of environmental and visit characteristics associated with particularly high levels of well-being for a specific outcome. For example, an optimal visit in terms of happiness might be to sandy beaches where there are high levels of perceived safety and excellent water quality; with a visit lasting at least three hours; and possibly involving playing with children, socialising, sunbathing/paddling and/or walking with a dog; and has short travel times that do not involve public transport.RQ1—natural and environmental featuresResearch question 1a—Which bluespace type(s) were associated with the highest levels of recalled visit mental well-being?Four of the five bluespace types associated with the highest levels of visit satisfaction were coastal (sea cliffs, rocky shore, sandy beaches, rural river and seaside promenade), indicating that these environments may be particularly beneficial for well-being. Visits to these environments were also associated with the lowest levels of visit anxiety, with the exception of seaside promenade and sea cliffs, which were not significantly different to the grand mean. Seaside promenade was the only urban environment in the top five.In addition, only coastal sites were associated with significantly higher levels of visit happiness (compared to the grand mean), further highlighting the potential importance of these environments. Although not explored here, coastal scenes tend to be associated with particularly high aesthetic and scenic value25,26 which may also be positively related to subjective well-being.These findings are broadly consistent with other studies from the UK17,27, but are extended here to our international sample. White et al.28 also used data from the BlueHealth International Survey (BIS) and explored visit frequency to different environments and associations with general mental health and well-being outcomes, including the World Health Organisation five-item Well-being index referring to the two weeks prior to the survey. Consistent with the results here, they found that visit frequency to “coastal blue” environments was more strongly associated with psychological well-being in general than visit frequency to “inland blue” environments. Our study adds to these more general findings by showing that these associations may come as a direct result of the recalled well-being experienced on specific visits to these locations.Confidence in our results was strengthened as we included general mental well-being in our analysis to adjust for whether happier people tend to visit sandy beaches, for example. The results for visit anxiety were not always the inverse of the trends observed in the positive measures of well-being, supporting the need to look at multiple aspects of mental well-being when considering the effects of nature contact.Research question 1b—Which bluespace qualities were associated with the highest levels of recalled visit mental well-being?Of the range of qualities that we investigated as predictors, perceived safety and ‘excellent’ water quality (vs. ‘sufficient’) consistently exhibited the strongest relationships with subjective mental well-being. Perceived safety has been found to be important when visiting blue spaces in several qualitative studies29,30,31, as well as a quantitative study with older adults in Hong Kong14. Blue spaces have particular safety issues with respect to drowning32,33, but fear of crime29,30,33 or pedestrian safety34 may also be relevant.Water quality has also been found to be important in previous economic valuation studies of recreational use and enjoyment of lakes and estuaries in the USA and Australia35,36 as well as a contingent behaviour experiment carried out as part of the BlueHealth International Survey (in European countries only)37. We recognise that here we used a metric of perceived water quality, rather than measures based on biological or toxicological sampling. Nevertheless, perceptions have been reported to positively correlate with sampled water quality parameters38, although assessments can vary systematically such as by bluespace type39. Highly visible harmful algal blooms, for instance, have also been found to affect experiences of blue spaces40.Further, and again consistent with earlier work15,41,42, the presence of facilities and wildlife, and absence of litter, were generally associated with better subjective mental well-being. Both perceived presence of wildlife and facilities were also associated with higher levels of anxiety however, indicating complexities between environmental qualities and well-being. Some wildlife may be deemed unpleasant or an ecosystem disservice, for example. The presence of good facilities may indicate the presence of more people; and visitor density in natural environments can be related to preference43. These results highlight the importance of environmental quality and not just type, consistent with other frameworks12,37.Research question 2—How is exposure, as operationalised by visit duration, related to recalled visit mental well-being?Broadly consistent with research in the green and bluespace literature14,17,44, we found that mental well-being outcomes were generally higher with greater exposure as indicated by visit duration. For decreasing visit anxiety, this was only significant when visits were longer than an hour and a half. As we did not measure pre-visit anxiety levels, we are cautious about identifying this as a potential temporal threshold for reducing anxiety at this stage.Similarly, also using the BlueHealth International Survey, White et al.28 found that well-being outcomes were higher with greater visit exposure to green and blue spaces using a metric of visit frequency. However, in contrast to this and other research which looked at overall weekly aggregated time in nature (e.g.28,45), we have no evidence of diminishing marginal returns as the effect sizes associated with specific visit duration continued to increase with increasing duration.Research question 3—What experiences in blue spaces, in terms of activities (3a) and companions (3b), are associated with the most positive recalled visit mental well-being outcomes?Although walking was the most popular activity, the activity with the highest mental well-being ratings was playing with children, especially in certain locations such as beaches (Fig. 4). However, we also find that anxiety tended to be higher when children were present. We speculate that the purpose of the visit may be important. For example, many who go to the beach with children do so in order to play. However, if children are present on more adult-oriented activities such as hiking, this may increase adult anxiety during the visit. From a representative sample of English adults, White et al.17 found that recent nature visits with children were associated with the lowest levels of well-being. Therefore, visits with children may be associated with a more complex set of emotions, being both slightly more stressful, but also potentially more rewarding and ‘meaningful’46. Ecosystem features of beaches may be particularly supportive of high well-being activities. A qualitative study in the UK, for instance, highlighted the particular opportunities for adults and children to play together at the beach, including rock-pooling and making sandcastles as well as water-based activities47.Visits with other adults were associated with higher levels of both visit satisfaction and worthwhile-ness, and socialising as an activity was associated with better visit well-being for all outcomes compared to the grand mean. This is consistent with studies using the day reconstruction method, which link activities with experiential well-being, in the USA48 and Germany49 where socialising was associated with the highest, or second highest, levels of well-being for all the activities assessed. Further, social interactions have been recognised as an important benefit by many of those visiting freshwater blue spaces in a previous study18.Research question 4—Does the relationship between wildlife presence and recalled visit well-being vary by bluespace settings?The relationship between the presence of wildlife and visit satisfaction varied with bluespace type. The strongest positive association was found for fen, marsh and bog areas, which may also be related to the purpose of visit. For instance, those who visit places such as fens, marshes and bogs, may do so for the explicit purpose of observing wildlife (often birds) and the presence of wildlife would therefore be important for satisfaction with the visit.Perceptions towards wildlife have been found to vary by location in other studies. For example, in Sweden, greater prior experience with geese at beaches was associated with a negative attitude towards geese50. Further, the species present are likely to vary across different environments. In three urban areas in the UK, green spaces correlated with the abundance and species richness of birds considered to provide cultural services (songbirds and woodpeckers), while an abundance of birds considered to provide disservices (e.g. some gull species, feral pigeons) was independent of green spaces51. Preferences for some species over others may explain some of the negative or null relationships between the presence of wildlife at different blue spaces. These examples from the literature, alongside our own results, indicate the potential for benefits from the management of wildlife for psychological ecosystem services differentially across environments, although these should be considered alongside other conservation and ESS goals.MechanismsSeveral mechanisms potentially explain the beneficial effects of visiting blue spaces on mental health and well-being12, including the provision of opportunities for physical activity52,53; social interaction18; cognitive restoration and stress reduction17,54; emotion regulation55 and connecting with nature12. Consistent with these mechanisms, we found that respondents were using blue spaces for both physical activity and social interaction; and that playing with children and socialising were associated with particularly high levels of well-being.In addition to the positive association we find between some ESS and well-being, including presence of wildlife and water quality, additional bluespace ESS not considered here, may also affect mental health and well-being12. For example, the provision of a cooling effect56 and air pollution mitigation57.Strengths and limitationsA key strength is our operationalisation of the Bratman et al.9 conceptual model for mental health using data from a large, 18 country survey that included 17 different bluespace types, five quality metrics and four subjective mental well-being outcomes. The relatively high explanatory power of our models suggests all the variables we explored were important for subjective well-being.Despite the strengths, however, there were also several limitations. The survey was cross-sectional and causality cannot be inferred. For example, happier people may choose to visit a beach rather than another location, although we also controlled for general levels of subjective mental well-being in an attempt to control for this possibility (See Supplemental Materials). Further, although the majority of respondents (53%) recalled a visit within the last 7 days, some were recalling visits up to a month ago, with potential memory biases increasing in line with length of recall.Although our data were collected by an international market research company to be representative of age, gender and region within country, our online sample may not be fully representative across more characteristics and any country-level conclusions need to be treated with caution. We also acknowledge that there were no results from Africa, the Middle East or South America; and Hong Kong was the only representative from Asia. This suggests far more research is needed in other regions to better understand how bluespace ecosystems interact with subjective well-being globally.There may also be socioeconomic confounds that we did not include in our models which may account for some of the effects. Not everyone visits nature for recreation58, including about 4000 people here who did not visit a bluespace in the four weeks prior to responding to the survey. Some groups may therefore have been under-represented; and we should be careful in assuming that our findings generalise to all sub-population groups.Nevertheless, our visit sub-sample distributions were generally similar to that of the weighted percentages in the full sample, with the exception of age where those aged over 60 were under-represented (Table S2); therefore, we suspect these issues were not too influential for the overall results, although care needs to be extended to inferences with respect to older adults.A further limitation was that we only considered the qualities of places where people reported making recreational visits, with respondents presumably less likely to visit places where they feel really unsafe or lacking in facilities29. Further research may want to study responses to a broader range of bluespace settings, including those that are less visited, to determine the generalisability of the generally positive results found here. Such studies could use pre-existing tools to objectively assess the quality of blue spaces59.ImplicationsOur finding that coastal environments are particularly beneficial adds to the body of evidence linking coastal environments with health and well-being and suggests this is consistent across many countries. Previous research has found that greater proximity to blue spaces, especially coastal settings, predicts visit frequency14,60,61 as well as other health outcomes—e.g. reduced risk of mortality and better general health, well-being and physical activity53,62. Here, we found that shorter travel times also predict visit well-being, highlighting the importance of having equitable access to good quality natural environments near to people’s homes.We also identified that different types of coastal and inland blue spaces (e.g. seaside promande, rural rivers), with different qualities (e.g. wildlife present), involving particular types of activities in specific social configurations (e.g. playing with children), were especially good at promoting well-being. This moves beyond a simple location-based assessment of benefit to one that recognises the complex interplay between location, behavioural and social processes. Numerous commentators63 (including Bratman et al.9 on which we have based this paper) have argued that we need to go beyond the determinate effects of green and blue spaces and develop a far richer, more nuanced understanding. The approach we have taken here is intended as a step in this direction.In terms of policy applications, these results provide support for the potential health benefits of efforts to improve equitable access to high quality environments, such as the English Coast Path (https://englandcoastpath.co.uk/) and the creation of beaches in Barcelona with the Olympic project in 199264. Our results also hint at the importance of high-level legislation, such as the EU’s Bathing Waters Directive65 for mental well-being37. If conducted on a more fine-grained geographical level, results could have the potential to leverage public support for more localised conservation initiatives. Furthermore, such results could be used as a basis for integration into more systematic conservation planning66.Further researchAlthough we incorporate a range of variables in our analysis, and our pseudo-R2 values are relatively high for a social research context, considerable variation remains unexplained. Although other individual characteristics may be important, such as nature connectedness67 and memories68, further research could explore the specific ecosystem features and social contexts associated with the particular positive results from coastal spaces, which would be of interest to policy makers and environmental managers. We also speculated that purpose of visit may explain some of our findings. Further research could explore the interactions between motivations and location, experience, and well-being outcomes.The presence of wildlife was differentially important across bluespace types and further research could unpack this. Exploring similar possibilities for the other quality metrics, as well as considering additional ecosystem characteristics, would also be informative. For example, identifying which factors are important in perceptions of safety in blue spaces. Bratman et al.9 also considered effect modification by visitor characteristics and further research could include interactions, or sub-group analysis, by socio-demographic factors.Further research could also explore longer-term benefits of these features over repeated visits; the potential for ecosystem disservices, such as the relationships we find between an interaction of wildlife and ice rinks and well-being; the potential for negative outcomes associated with ecosystem degradation69; and the potential for positive mental health outcomes from ecological restoration70.We have demonstrated some of the complexities involved in the human-nature relationship and that many factors are related to the outcome from a visit. The conceptual model applied allows the investigation of a wide range of variables including natural features and other environmental qualities, and characteristics of the exposure and experience, as well as individual parameters. We suggest that other researchers can apply this conceptual model and design data collection accordingly to target specific research questions and hypotheses (as opposed to where we have fitted already collected data). More