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7 páginas.- 4 figuras.- 32 referencias.- Supplementary materials: science.org/doi/10.1126/science.abq4062 Materials and Methods Figs. S1 to S19 Tables S1 to S28 References (33–269) MDAR Reproducibility Checklist Movie S1.- Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and speciespoor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure. Copyright © 2022 the authors, Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species-poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. Considering interactions between grazing and local abiotic and biotic factors is key for understanding the fate of dryland ecosystems under climate change and increasing human pressure. Copyright © 2022 the authors, Funding: This research was funded by the European Research Council [ERC grant agreement 647038 (BIODESERT)] and Generalitat Valenciana (CIDEGENT/2018/ 041). F.T.M. acknowledges support from a Rei Jaume I Award, the Alexander von Humboldt Foundation, and the Synthesis Center (sDiv) of the German Centre for Integrative Biodiversity Research Halle–Jena–Leipzig (iDiv). C.A.G., S.C., and N.E. acknowledge support from iDiv and the Deutsche Forschungsgemeinschaft (DFG– FZT 118, 202548816; Flexpool proposal 34600850). Y.L.B.-P. was supported by a Marie Sklodowska-Curie Actions Individual Fellowship (MSCA-IF) within the European Program Horizon 2020 (DRYFUN Project 656035). N.G. was supported by CAP 20-25 (16-IDEX-0001) and the AgreenSkills+ fellowship program, which has received funding from the EU’s Seventh Framework Programme under grant agreement N° FP7-609398 (AgreenSkills+ contract). B.B. and B.E. were supported by the Taylor Family–Asia Foundation Endowed Chair in Ecology and Conservation Biology. J.D., A.Ro., and H.C. acknowledge support from the Fundação para a Ciência e a Tecnologia (IF/00950/ 2014 and 2020.03670.CEECIND, SFRH/BDP/108913/2015, and in the scope of the framework contract foreseen in the numbers 4-6 of the article 23, of the Decree-Law 57/2016, August 29, changed by Law 57/2017, July 19, respectively), as well as from the MCTES, FSE, UE, and the CFE (UIDB/04004/2020) research unit financed by Fundação para a Ciência e a Tecnologia/MCTES through national funds (PIDDAC). C.P. acknowledges support from the Spanish Ministry of Science and Innovation (ref. AGL201675762-R, AEI/FEDER, UE, and PID2020-116578RB-I00, MCIN/AEI/10.13039/501100011033) and the European Union’s Horizon 2020 Research and Innovation Programme under grant agreement no. 101000224. E.V. was funded by the 2017 program for attracting and retaining talent of Comunidad de Madrid (no. 2017‐T2/ AMB‐5406). M.A.B. acknowledges support from the School of Forestry and College of the Environment, Forestry and Natural Sciences of Northern Arizona University. E.H.-S. acknowledges support from the Consejo Nacional de Ciencia y Tecnología (SEP-CB-2015-01-251388, PN 2017-5036 and PRONAII 319059). F.M.H. acknowledges support from the National Council for Scientific and Technological Development (CNPq - PCI/INMA) of the Brazilian Ministry of Science, Technology and Innovation (MCTI, processes number 302381/2020-1). H.L.T. acknowledges support from the US National Science Foundation (NSF) (DEB 0953864). A.N. and M.K. acknowledge support from the Fundação para a Ciência e a Tecnologia (SFRH/BD/130274/2017, CEECIND/02453/2018/CP1534/CT0001, PTDC/ASP-SIL/7743/2020 and UIDB/00329/2020). A.A.C. acknowledges support from the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001. J.E.K. and T.P.M. acknowledge the National Research Foundation of South Africa (grant no. 114412). F.J. and N.B. acknowledge support from the German Federal Ministry of Education and Research (BMBF) in the framework of the SPACES projects OPTIMASS (FKZ: 01LL1302A) and ORYCS (FKZ:01LL1804A). A.Li. and A.S.K.F. acknowledge support from the German Federal Ministry of Education and Research (BMBF) in the framework of the SPACES projects Limpopo Living Landscapes (FKZ: 01LL1304D) and SALLnet (FKZ: 01LL1802C). L.W. acknowledges support from the US NSF (EAR 1554894). L.H.F. acknowledges support from the Natural Sciences and Engineering Research Council of Canada Industrial Research Chair Program in Ecosystem Reclamation. S.C.R. acknowledges support from the US Geological Survey Ecosystems Mission Area and the US Bureau of Land Management. G.M.W. acknowledges support from the Australian Research Council. L.v.d.B. and K.T. acknowledge support from the German Research Foundation (DFG) priority research program SPP-1803 “EarthShape: Earth Surface Shaping by Biota” (TI 338/14-1). M.D.-B. acknowledges support from the Spanish Ministry of Science and Innovation for the I+D+i project PID2020-115813RA-I00 funded by MCIN/AEI/10.13039/501100011033. M.D.-B. is also supported by a project of the Fondo Europeo de Desarrollo Regional (FEDER) and the Consejería de Transformación Económica, Industria, Conocimiento y Universidades of the Junta de Andalucía (FEDER Andalucía 2014-2020 Objetivo temático “01 - Refuerzo de la investigación, el desarrollo tecnológico y la innovación”) associated with the research project P20_00879 (ANDABIOMA). P.J.R. and A.J.M. acknowledge support from Fondo Europeo de Desarrollo Regional through the FEDER Andalucía operative program, FEDER-UJA 1261180 project. A.F. thanks ANID PIA/BASAL FB210006 and Millennium Science Initiative Program NCN2021-050. A.J. acknowledges support from the Bavarian Research Alliance Germany (BayIntAn_UBT_2017_61). C.B. acknowledges the National Natural Science Foundation of China (grant no. 41971131). Biodiversity and ecosystem function research in the B.K.S. laboratory is funded by the Australian Research Council (DP210102081). Any use of trade, product, or firm names in this paper is for descriptive purposes only and does not imply endorsement by the US government. H.S. is supported by a María Zambrano fellowship funded by the Ministry of Universities and European Union-Next Generation plan. G.P. and C.V.L. acknowledge support from Universidad Nacional de Río Negro (PI 40-C-873 and 654). V.R. acknowledges support from the Regional Government of Extremadura (Spain) through a “Talento” fellowship (TA18022). M.F. acknowledges support from the Department of Range and Watershed Management, Ferdowsi University of Mashhad, Mashhad, Iran. Participation of recent graduates in collecting field data at four sites in Namibia was supported by a capacity building grant to Gobabeb–Namib Research Institute by the Environmental Investment Fund in Namibia.

The discovery of complete ammonia oxidizers, comammox Nitrospira, represents a breakthrough in the history of nitrification research. Nitrospira inopinata, which was obtained from an aquatic ecosystem, is the only pure comammox bacterial isolate reported so far. Ammonia oxidation kinetics of N. inopinata indicated that they prefer an oligotrophic lifestyle and may directly compete with ammonia-oxidizing archaea (AOA), but representatives of comammox Nitrospira from terrestrial ecosystems are lacking. Current studies demonstrate that the functionally dominant comammox Nitrospira in terrestrial ecosystems are separated from most sequences obtained from aquatic and engineering systems. Clades A and B of comammox Nitrospira seemingly have different ecological preferences in soils, possibly due to their different ammonium uptake systems. Analyses of the evolutionary history indicate that some genes of comammox Nitrospira involved in ammonia oxidation could be laterally transferred from β-ammonia-oxidizing bacteria (AOB). Some comammox Nitrospira species may have similar ecological preferences with AOB and be functionally active in the niches where nitrification was previously considered to be dominated by AOB. Taken together, this review summarizes the recent findings of the biogeographical distribution and ecological preference of comammox Nitrospira in large-scale soil surveys, the responses of comammox Nitrospira to nitrogen application in soil, and the putative mechanisms underpinning the ecological niche of terrestrial comammox Nitrospira. These studies from terrestrial ecosystems suggest that comammox Nitrospira are not strictly oligotrophic but both oligotrophic and copiotrophic with a broader ecological niche breadth. Comammox Nitrospira in terrestrial ecosystems are abundant and active in both oligotrophic and copiotrophic environments. These findings have profoundly expanded our knowledge of the environmental niches of comammox Nitrospira and their relative contribution to nitrification in soils.