RES EARCH

DRYLAND ECOLOGY they constitute 78% of rangelands worldwide

(14) and support ~1 billion people who rely on
Grazing and ecosystem service delivery in grazing by livestock as a critical source of pro-
tein and income (15). Although grazing may
global drylands have beneficial effects by reducing fuel loads
and enhancing primary production and plant
Fernando T. Maestre1,2*, Yoann Le Bagousse-Pinguet3, Manuel Delgado-Baquerizo4,5, David J. Eldridge6, diversity under certain conditions (3, 16), in-
Hugo Saiz7,8, Miguel Berdugo9,10, Beatriz Gozalo1, Victoria Ochoa1,11, Emilio Guirado1, creasing grazing pressure is also considered
Miguel García-Gómez12, Enrique Valencia13,14, Juan J. Gaitán15,16,17, Sergio Asensio1, Betty J. Mendoza13, a major driver of rangeland degradation and
César Plaza11, Paloma Díaz-Martínez11, Ana Rey18, Hang-Wei Hu19,20, Ji-Zheng He19,20, Jun-Tao Wang21,22,23, desertification across drylands worldwide (17).
Anika Lehmann24,25, Matthias C. Rillig24,25, Simone Cesarz26,27, Nico Eisenhauer26,27, These contrasting effects of grazing likely de-
Jaime Martínez-Valderrama1, Eduardo Moreno-Jiménez28, Osvaldo Sala29,30,31, Mehdi Abedi32, pend on local climate, soil conditions, and both
Negar Ahmadian32, Concepción L. Alados33, Valeria Aramayo34, Fateh Amghar35, Tulio Arredondo36, plant and soil diversity, which largely influence
Rodrigo J. Ahumada37, Khadijeh Bahalkeh32, Farah Ben Salem38, Niels Blaum39, Bazartseren Boldgiv40, dryland functioning (18, 19). However, the in-
Matthew A. Bowker41,42, Donaldo Bran34, Chongfeng Bu43,44, Rafaella Canessa45,46, teractions of these factors with grazing pres-
Andrea P. Castillo-Monroy47, Helena Castro48, Ignacio Castro49, Patricio Castro-Quezada50, sure have never, to our knowledge, been
Roukaya Chibani38, Abel A. Conceição51, Courtney M. Currier29,31, Anthony Darrouzet-Nardi52, assessed. Identifying environmental conditions
Balázs Deák53, David A. Donoso47,54, Andrew J. Dougill55, Jorge Durán48,56, Batdelger Erdenetsetseg40, and biodiversity levels under which increasing

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Carlos I. Espinosa57, Alex Fajardo58, Mohammad Farzam59, Daniela Ferrante60,61, grazing pressure will favor or detract ecosys-
Anke S. K. Frank62,63,64, Lauchlan H. Fraser65, Laureano A. Gherardi66, Aaron C. Greenville63, tem service delivery is a crucial step toward
Carlos A. Guerra26,67, Elizabeth Gusmán-Montalvan57, Rosa M. Hernández-Hernández49, achieving multiple UN Sustainable Develop-
Norbert Hölzel68, Elisabeth Huber-Sannwald36, Frederic M. Hughes51,69, Oswaldo Jadán-Maza50, ment Goals (2) and other international initia-
Florian Jeltsch25,39, Anke Jentsch70, Kudzai F. Kaseke71, Melanie Köbel72, Jessica E. Koopman73, tives related to dryland desertification and
Cintia V. Leder17,74, Anja Linstädter64,75, Peter C. le Roux76, Xinkai Li43,44, Pierre Liancourt46,77,78, restoration (20).
Jushan Liu79, Michelle A. Louw76, Gillian Maggs-Kölling80, Thulani P. Makhalanyane73, Here, we used a standardized field survey
Oumarou Malam Issa81, Antonio J. Manzaneda82,83, Eugene Marais80, Juan P. Mora58, (13) carried out at 98 sites across 25 countries
Gerardo Moreno84, Seth M. Munson85, Alice Nunes72, Gabriel Oliva60,61, Gastón R. Oñatibia86, and six continents (Fig. 1 and movie S1) to
Guadalupe Peter17,74, Marco O. D. Pivari87, Yolanda Pueyo33, R. Emiliano Quiroga37,88, assess how the effects of grazing pressure on
Soroor Rahmanian59,89, Sasha C. Reed90, Pedro J. Rey82,83, Benoit Richard91, Alexandra Rodríguez48, nine essential ecosystem services depend on
Víctor Rolo84, Juan G. Rubalcaba92, Jan C. Ruppert46, Ayman Salah93, Max A. Schuchardt70, biodiversity, climate, and soil conditions across
Sedona Spann41, Ilan Stavi94, Colton R. A. Stephens65, Anthony M. Swemmer95, Alberto L. Teixido96, global drylands. Each site included a collection
Andrew D. Thomas97, Heather L. Throop98,99, Katja Tielbörger46, Samantha Travers100, James Val101, of three or four 45-m–by–45-m plots repre-
Orsolya Valkó53, Liesbeth van den Brink46, Sergio Velasco Ayuso86, Frederike Velbert68, senting local gradients of grazing pressure
Wanyoike Wamiti102, Deli Wang79, Lixin Wang103, Glenda M. Wardle63, Laura Yahdjian86, Eli Zaady104, [from ungrazed or low grazing pressure to
Yuanming Zhang105, Xiaobing Zhou105, Brajesh K. Singh21,22, Nicolas Gross106 high grazing pressure (13)], resulting in a total
of 326 plots. These gradients were mostly
Grazing represents the most extensive use of land worldwide. Yet its impacts on ecosystem services driven by livestock (fig. S1), although wild
remain uncertain because pervasive interactions between grazing pressure, climate, soil properties, herbivores were also present in each site and
and biodiversity may occur but have never been addressed simultaneously. Using a standardized survey taken into account. In each plot, we assessed
at 98 sites across six continents, we show that interactions between grazing pressure, climate, soil, vascular plant, mammalian herbivore (ac-
and biodiversity are critical to explain the delivery of fundamental ecosystem services across drylands counting for domestic and wild herbivores),
worldwide. Increasing grazing pressure reduced ecosystem service delivery in warmer and species- and belowground (soil bacteria, fungi, pro-
poor drylands, whereas positive effects of grazing were observed in colder and species-rich areas. tists, and invertebrates) diversity as well as
Considering interactions between grazing and local abiotic and biotic factors is key for understanding multiple regulating (water regulation, soil
the fate of dryland ecosystems under climate change and increasing human pressure. carbon storage, organic matter decomposi-
tion, and erosion control), supporting (soil

G
fertility and aboveground plant biomass and
razing accounts for 77% of global agri- interactions between grazing pressure and its temporal stability), and provisioning (wood
cultural land (1), sustains billions of abiotic and biotic features, which results in quantity, forage quantity, and quality) eco-
people worldwide, and is closely linked strong context-dependent ecological impacts system services (table S1). Our survey captured
to 10 of 17 United Nations (UN) Sus- of grazing (3, 4, 10, 11). Large-scale, stan- most climatic conditions supporting livestock
tainable Development Goals (2). Despite dardized field surveys that explore how such grazing in drylands, as well as a wide range
its importance, there is no consensus on how impacts depend on above- and belowground of ecosystem types; soil properties; plant, soil,
grazing affects ecosystem services (3–6), which biodiversity, soils, and climate to drive multi- and mammalian diversities; and grazing pres-
may depend on the coevolutionary history ple ecosystem services across contrasting re- sure levels (figs. S2 to S9 and table S2). These
between vegetation and herbivores (3), graz- gions and environmental contexts are lacking distinctive features of our global study rendered
ing pressure (4), and local climatic, edaphic, at present but are sorely needed to evaluate grazing pressure largely independent of cli-
and biodiversity conditions (7, 8). Most field whether general patterns emerge beyond these mate, soil, and biodiversity attributes [table S3
assessments have focused on local to regional context dependencies (12). and (13)] and allowed us to (i) evaluate the
scales (3, 4, 6, 8), have studied a limited num- Investigating the effects of grazing pressure main and interactive effects of grazing pres-
ber of taxa—mostly plants—and single ecosys- across global abiotic and biotic gradients is sure, climate, soil properties, and biodiversity
tem services (3, 4, 9), and have not considered particularly important in drylands [areas with on ecosystem service delivery across global
domestic and wild herbivores simultaneously. an aridity index (precipitation divided by po- drylands; (ii) identify the environmental and
Another major source of uncertainty relates to tential evapotranspiration) <0.65 (13)] because biodiversity conditions under which the effects

Maestre et al., Science 378, 915–920 (2022) 25 November 2022 1 of 6

RES EARCH | REPOR T

of grazing pressure on ecosystem services are system services through direct effects (no extent, with mean annual precipitation (9% of
positive or negative; and (iii) simultaneously significant indirect effects through changes in the best-fitting models). A negative relation-
assess relationships between plant, soil, and soil properties or biodiversity were found; ship between mean annual temperature and
mammalian herbivore diversity and multiple figs. S10 and S11 and tables S4 to S12) and in- soil carbon storage, organic matter decompo-
ecosystem services. teractive effects (interactions between grazing sition, and erosion control was found under
We fitted linear mixed models to data from and climate, grazing and soil properties, or high, but not under low, grazing pressure (Fig.
all sites and grazing pressure levels and ap- grazing and biodiversity were selected in 86% 3, A to C). Our results provide an empirical
plied a multimodel inference procedure based of the best-fitting models; Fig. 2 and tables S13 validation of the importance of interactions
on Akaike information criterion (AIC) to select to S28). between climate change drivers, grazing, and
the set of best-fitting models [i.e., those with a Interactions between grazing and climate soil carbon storage that are predicted by glob-
DAIC <2 (13)]. We also considered potential in- were selected in 48% of the best-fitting models al modeling studies (21). They also indicate
direct effects of grazing through the modification (fig. S12), with grazing primarily interacting that considering grazing pressure can improve
of local biodiversity and soil parameters using with mean annual temperature (40% of the our capacity to assess soil carbon–temperature
confirmatory path analyses (13). We found best-fitting models) and rainfall seasonality feedbacks, a key process involved in climate
that increasing grazing pressure affects eco- (20% of the best-fitting models) and, to a lesser warming (22).

1
Instituto Multidisciplinar para el Estudio del Medio “Ramón Margalef,” Universidad de Alicante, Alicante, Spain. 2Departamento de Ecología, Universidad de Alicante, Alicante, Spain. 3Aix Marseille
Univ, CNRS, Avignon Université, IRD, IMBE, Aix-en-Provence, France. 4Laboratorio de Biodiversidad y Funcionamiento Ecosistémico, Instituto de Recursos Naturales y Agrobiología de Sevilla

Downloaded from https://www.science.org at Universidade Federal do Mato Grosso on November 24, 2022
(IRNAS), CSIC, Sevilla, Spain. 5Unidad Asociada CSIC-UPO (BioFun), Universidad Pablo de Olavide, Sevilla, Spain. 6Department of Planning and Environment, c/o Centre for Ecosystem Science,
School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, New South Wales, Australia. 7Departamento de Ciencias Agrarias y Medio Natural, Escuela
Politécnica Superior, Instituto Universitario de Investigación en Ciencias Ambientales de Aragón (IUCA), Universidad de Zaragoza, Huesca, Spain. 8Institute of Plant Sciences, University of Bern,
Bern, Switzerland. 9Institut de Biología Evolutiva (UPF-CSIC), Barcelona, Spain. 10Department of Environmental Systems Science, ETH Zurich, Zurich, Switzerland. 11Instituto de Ciencias
Agrarias, Consejo Superior de Investigaciones Científicas, Madrid, Spain. 12Departamento de Ingeniería y Morfología del Terreno, Escuela Técnica Superior de Ingenieros de Caminos, Canales y
Puertos, Universidad Politécnica de Madrid, Madrid, Spain. 13Departamento de Biología y Geología, Física y Química Inorgánica, Universidad Rey Juan Carlos, Móstoles, Spain. 14Departamento de
Biodiversidad, Ecología y Evolución, Facultad de Ciencias Biológicas, Universidad Complutense de Madrid, Madrid, Spain. 15Instituto Nacional de Tecnología Agropecuaria (INTA), Instituto de
Suelos-CNIA, Buenos Aires, Argentina. 16Universidad Nacional de Luján, Departamento de Tecnología, Luján, Argentina. 17Consejo Nacional de Investigaciones Científicas y Técnicas de Argentina
(CONICET), Buenos Aires, Argentina. 18Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas, Madrid, Spain. 19Key Laboratory for Humid Subtropical Eco-
geographical Processes of the Ministry of Education, School of Geographical Science, Fujian Normal University, Fuzhou, China. 20Faculty of Veterinary and Agricultural Sciences, The University of
Melbourne, Melbourne, Victoria, Australia. 21Global Centre for Land-Based Innovation, Western Sydney University, Sydney, New South Wales, Australia. 22Hawkesbury Institute for the Environment,
Western Sydney University, Sydney, New South Wales, Australia. 23State Key Laboratory of Urban and Regional Ecology, Research Center for Eco-Environmental Sciences, Chinese Academy of
Sciences, Beijing, China. 24Freie Universität Berlin, Institute of Biology, Berlin, Germany. 25Berlin-Brandenburg Institute of Advanced Biodiversity Research (BBIB), Berlin, Germany. 26German
Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, Germany. 27Leipzig University, Institute of Biology, Leipzig, Germany. 28Department of Agricultural and Food
Chemistry, Faculty of Sciences, Universidad Autónoma de Madrid, Madrid, Spain. 29School of Life Sciences, Arizona State University, Tempe, AZ, USA. 30School of Sustainability, Arizona State
University, Tempe, AZ, USA. 31Global Drylands Center, Arizona State University, Tempe, AZ, USA. 32Department of Range Management, Faculty of Natural Resources and Marine Sciences, Tarbiat
Modares University, Noor, Mazandaran Province, Iran. 33Instituto Pirenaico de Ecología (IPE, CSIC), Zaragoza, Spain. 34Instituto Nacional de Tecnología Agropecuaria (INTA), Estación Experimental
Agropecuaria Bariloche, Bariloche, Río Negro, Argentina. 35Laboratoire de Recherche: Biodiversité, Biotechnologie, Environnement et Développement Durable (BioDev), Faculté des Sciences,
Université M’hamed Bougara de Boumerdès, Boumerdès, Algérie. 36Instituto Potosino de Investigación Científica y Tecnológica, A.C., San Luis Potosí, Mexico. 37Instituto Nacional de Tecnología
Agropecuaria, Estación Experimental Agropecuaria Catamarca, Catamarca, Argentina. 38Laboratory of Range Ecology, Institut des Régions Arides (IRA), Médenine, Tunisia. 39University of Potsdam,
Plant Ecology and Conservation Biology, Potsdam, Germany. 40Laboratory of Ecological and Evolutionary Synthesis, Department of Biology, School of Arts and Sciences, National University of
Mongolia, Ulaanbaatar, Mongolia. 41School of Forestry, Northern Arizona University, Flagstaff, AZ, USA. 42Center for Ecosystem Science and Society, Northern Arizona University, Flagstaff, AZ, USA.
43
Institute of Soil and Water Conservation, Northwest A&F University, Yangling, Shaanxi, China. 44Institute of Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water
Resources, Yangling, Shaanxi, China. 45Ecological Plant Geography, Faculty of Geography, University of Marburg, Marburg, Germany. 46Plant Ecology Group, University of Tübingen, Tübingen,
Germany. 47Departamento de Biología, Escuela Politécnica Nacional, Quito, Ecuador. 48Centre for Functional Ecology, Department of Life Sciences, University of Coimbra, Coimbra, Portugal.
49
Universidad Nacional Experimental Simón Rodríguez (UNESR), Instituto de Estudios Científicos y Tecnológicos (IDECYT), Centro de Estudios de Agroecología Tropical (CEDAT), Miranda,
Venezuela. 50Universidad de Cuenca, Facultad de Ciencias Agropecuarias, Carrera de Ingeniería Agronómica, Grupo de Agroforestería, Manejo y Conservación del paisaje, Cuenca, Ecuador.
51
Universidade Estadual de Feira de Santana (UEFS), Departamento de Ciências Biológicas, Bahia, Brazil. 52Department of Biological Sciences, University of Texas at El Paso, El Paso, TX, USA.
53
Lendület Seed Ecology Research Group, Institute of Ecology and Botany, Centre for Ecological Research, Vácrátót, Hungary. 54Centro de Investigación de la Biodiversidad y Cambio Climático,
Universidad Tecnológica Indoamérica, Quito, Ecuador. 55Department of Environment and Geography, University of York, York, UK. 56Misión Biolóxica de Galicia, CSIC, Pontevedra, Spain.
57
Departamento de Ciencias Biológicas, Universidad Técnica Particular de Loja, Loja, Ecuador. 58Instituto de Investigación Interdisciplinaria (I3), Vicerrectoría Académica, Universidad de Talca,
Talca, Chile. 59Department of Range and Watershed Management, Ferdowsi University of Mashhad, Mashhad, Iran. 60Instituto Nacional de Tecnología Agropecuaria EEA Santa Cruz, Río Gallegos,
Santa Cruz, Argentina. 61Universidad Nacional de la Patagonia Austral, Río Gallegos, Santa Cruz, Argentina. 62School of Agriculture, Environmental and Veterinary Sciences, Charles Sturt
University, Port Macquarie, New South Wales, Australia. 63Desert Ecology Research Group, School of Life and Environmental Sciences, The University of Sydney, Sydney, New South Wales,
Australia. 64Institute of Crop Science and Resource Conservation, University of Bonn, Bonn, Germany. 65Department of Natural Resource Science, Thompson Rivers University, Kamloops, British
Columbia, Canada. 66Department of Environmental Science, Policy and Management, University of California, Berkeley, Berkeley, CA, USA. 67Institute of Biology, Martin-Luther University Halle
Wittenberg, Halle (Saale), Germany. 68Institute of Landscape Ecology, University of Münster, Münster, Germany. 69Instituto Nacional da Mata Atlântica (INMA), Espírito Santo, Brazil. 70Department
of Disturbance Ecology, Bayreuth Center of Ecology and Environmental Research BayCEER, University of Bayreuth, Bayreuth, Germany. 71Earth Research Institute, University of California, Santa
Barbara, Santa Barbara, CA, USA. 72Centre for Ecology, Evolution and Environmental Changes, Faculdade de Ciências, Universidade de Lisboa, Lisboa, Portugal. 73Microbiome@UP, Department of
Biochemistry, Genetics and Microbiology, University of Pretoria, Pretoria, South Africa. 74Universidad Nacional de Río Negro, Sede Atlántica, CEANPa, Río Negro, Argentina. 75Biodiversity
Research/Systematic Botany Group, Institute of Biochemistry and Biology, University of Potsdam, Potsdam, Germany. 76Department of Plant and Soil Sciences, University of Pretoria, Pretoria,
South Africa. 77Institute of Botany, Czech Academy of Sciences, Pruhonice, Czech Republic. 78Botany Department, State Museum of Natural History Stuttgart, Stuttgart, Germany. 79Key
Laboratory of Vegetation Ecology of the Ministry of Education, Jilin Songnen Grassland Ecosystem National Observation and Research Station, Institute of Grassland Science, Northeast Normal
University, Changchun, China. 80Gobabeb-Namib Research Institute, Walvis Bay, Namibia. 81Institut d’Écologie et des Sciences de l’Environnement de Paris (iEES-Paris), Sorbonne Université,
IRD, CNRS, INRAE, Université Paris Est Creteil, Université de Paris, Centre IRD de France Nord, Bondy, France. 82Instituto Interuniversitario de Investigación del Sistema Tierra en Andalucía,
Universidad de Jaén, Jaén, Spain. 83Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Jaén, Spain. 84Forestry School, INDEHESA, Universidad de Extremadura,
Plasencia, Spain. 85US Geological Survey, Southwest Biological Science Center, Flagstaff, AZ, USA. 86Cátedra de Ecología, Facultad de Agronomía, Universidad de Buenos Aires, Instituto de
Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura (IFEVA-CONICET), Ciudad Autónoma de Buenos Aires, Argentina. 87Departamento de Botânica, Universidade Federal de
Minas Gerais, Minas Gerais, Brazil. 88Cátedra de Manejo de Pastizales Naturales, Facultad de Ciencias Agrarias, Universidad Nacional de Catamarca, Catamarca, Argentina. 89Department of
Forest Engineering, Forest Management Planning and Terrestrial Measurements, Faculty of Silviculture and Forest Engineering, Transilvania University of Brasov, Brasov, Romania 90US
Geological Survey, Southwest Biological Science Center, Moab, UT, USA. 91Normandie Univ, UNIROUEN, INRAE, ECODIV, Rouen, France. 92Deparment of Biology, McGill University, Montreal,
Quebec, Canada. 93Al Quds University, Abu Dis, Palestine. 94Dead Sea and Arava Science Center, Yotvata, Israel. 95South African Environmental Observation Network (SAEON), Phalaborwa,
Kruger National Park, South Africa. 96Departamento de Botânica e Ecologia, Instituto de Biociências, Universidade Federal de Mato Grosso, Mato Grosso, Brazil. 97Department of Geography and
Earth Sciences, Aberystwyth University, Wales, UK. 98School of Earth and Space Exploration, Arizona State University, Tempe, AZ, USA. 99School of Life Sciences, Arizona State University,
Tempe, AZ, USA. 100Centre for Ecosystem Science, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, New South Wales, Australia. 101Science
Division, Department of Planning, Industry and Environment, New South Wales Government, Buronga, New South Wales, Australia. 102Zoology Department, National Museums of Kenya, Nairobi,
Kenya. 103Department of Earth Sciences, Indiana University–Purdue University Indianapolis (IUPUI), Indianapolis, IN, USA. 104Department of Natural Resources, Agricultural Research
Organization, Institute of Plant Sciences, Gilat Research Center, Mobile Post Negev, Israel. 105State Key Laboratory of Desert and Oasis Ecology, Xinjiang Institute of Ecology and Geography,
Chinese Academy of Sciences, Urumqi, China. 106Université Clermont Auvergne, INRAE, VetAgro Sup, Unité Mixte de Recherche Ecosystème Prairial, Clermont-Ferrand, France.
*Corresponding author. Email: [email protected]

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Fig. 1. Locations of the 98 study sites with examples of the local grazing for livestock in drylands; they were used in this study to create local grazing
gradients surveyed at each site. Each black dot represents a site with multiple gradients (13). The background of the map indicates the extent of dryland
45-m–by–45-m plots (white dots) surveyed in situ; a total of 326 plots were rangeland areas. The aridity index is calculated as precipitation divided by
surveyed across the 98 study sites. The inset graphics [(A) to (G)] highlight potential evapotranspiration and is strongly related to mean annual precipitation
examples of the local gradients surveyed at each site. Watering points are ponds, in our dataset [coefficient of determination (R2) = 0.82]. See (13) for the
impoundments, or drinking troughs that provide permanent sources of water aridity index and rangeland area data sources that were used.

Soil texture also regulated grazing pressure

effects on multiple ecosystem services, which
include soil fertility, wood quantity, and for-
age quality (interactions between grazing and
sand content were selected in 37% of the best-
fitting models; fig. S12). As sand content in-
creased, soil fertility declined more steeply
under high grazing pressure (Fig. 3E), wood
quantity increased under high but declined
under low grazing pressure (Fig. 3G), and
forage quality declined under high but in-
creased under low grazing pressure (Fig. 3I).
These findings illustrate how increases in
grazing pressure interact with soil properties
to either increase or reduce the delivery of
multiple ecosystem services.
Biodiversity impacts on ecosystem func-
tioning and services are typically examined
in isolation from other drivers in experimen-
tal and observational studies (23). However,
we found interactions between grazing and
biodiversity in 44% of the best-fitting models
(fig. S12). For instance, increasing grazing
pressure shifted the relationships between Fig. 2. Relative importance of predictors of ecosystem services selected in the best-fitting models.
plant species richness and water regulation Importance is quantified as the sum of the Akaike weights of all models that included the predictor (grazing pressure,
from positive to negative (Fig. 3D) and those climate, biodiversity, and soil variables, and their interactions) of interest, considering the number of models in
between plant species richness and both wood which each predictor appears. It is proportional to the number of times that a given predictor (and its interactions
quantity and aboveground plant biomass and with other predictors) was selected in the final set of best-fitting models (13). Interactions include all interactions
its temporal stability from negative to posi- between grazing pressure and climate, biodiversity, and soil variables; the importance of each interaction type
tive (Fig. 3, F and G). We also found positive is shown in fig. S12. In the case of biodiversity, predictor importance considers the number of models that include at
relationships between plant species richness least one biodiversity proxy (plant species richness, mammalian herbivore richness, or belowground diversity).
and soil carbon storage, organic matter de- Separate results for each biodiversity proxy are shown in fig. S12. Full details on model results, including the number
composition, erosion control, and both forage of best-fitting models, are available in tables S13 to S15. “Plant biomass and stability” represents aboveground
quality and quantity (Fig. 3, A to C, H, and I) and plant biomass and its temporal stability, and “grazing” represents grazing pressure. MAT, mean annual temperature;
between belowground diversity and organic RASE, rainfall seasonality; MAP, mean annual precipitation.

Maestre et al., Science 378, 915–920 (2022) 25 November 2022 3 of 6

RES EARCH | REPOR T

biodiversity conditions. They also suggest that

efforts to promote diverse grazing systems
may enhance soil carbon storage and reduce
negative impacts of increased grazing pres-
sure. To date, such results have only been
modeled or observed locally (26, 27).
The multiple interactions we observed high-
light that the effect of grazing pressure on eco-
system services can be positive or negative
depending on local climate, soil, and biodi-
versity conditions (Fig. 4). On average, in-
creasing grazing pressure had positive effects
on ecosystem services in colder sites with high
plant species richness but negative effects in
warmer sites with high rainfall seasonality
and low plant species richness (Fig. 4, E and I).
When sets of ecosystem services were consid-
ered separately, responses to grazing pres-

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sure ranged from mostly neutral to positive
(regulating and supporting services; Fig. 4, B
and C) and from negative to neutral (provi-
sioning services; Fig. 4D). These results allow
us to identify ecological conditions under which
ecosystem services are positively or negatively
associated with changes in grazing pressure
(Fig. 4 and figs. S16 to S18) and to frame new
hypotheses that explore the local context de-
pendencies of grazing impacts. For instance,
we observed negative effects of increasing
grazing pressure on ecosystem services in
plant species–poor drylands, as reported in
recent local-scale studies [e.g., (11)], whereas posi-
tive effects of grazing were mostly observed
in species-rich drylands. Thus, protecting
biodiversity in species-rich areas or restor-
ing it in species-poor areas could minimize
some of the negative effects of increasing
Fig. 3. Predicted responses of ecosystem services to changes in climate, sand content, and plant grazing pressure on ecosystem service delivery
species richness at low and high grazing pressure levels. (A to I) Predicted responses of regulating (fig. S19).
[(A) to (D)], supporting [(E) and (F)], and provisioning [(G) to (I)] ecosystem services. The lines in each panel The effects of increasing grazing pressure on
show model fits (using partial residuals) for each predictor selected in the final best-fitting models at low ecosystem services were mostly negative in
(dashed lines) and high (solid lines) grazing pressures for each service. Shading around each line represents warmer drylands (Fig. 4 and fig. S17), where
the 95% confidence interval. Panels surrounded by a border denote significant interactions between a large proportion of the human population
grazing and other predictors. Predicted responses of ecosystem services to all grazing pressure levels relies heavily on livestock for subsistence (15).
(ungrazed, low, medium, and high) and to other model predictors are presented in figs. S13 to S15. The Limiting grazing pressure through livestock
complete set of statistical results and model fits are available in tables S13 to S15. “Sand” represents sand removal is neither socially nor economically
content. PR, plant species richness. feasible in these areas (2), yet they are ex-
pected to experience high warming rates and
water shortages under most climate change
matter decomposition (fig. S13), irrespective S13), with aboveground plant biomass and scenarios (17). Our results thus suggest that
of grazing pressure. These results broaden and its temporal stability under high grazing pres- grazing pressure may interact with climate
validate previous findings on the relationship sure (fig. S14), and with forage quality under change to reduce ecosystem service delivery
between biodiversity and ecosystem functioning low grazing pressure (fig. S15). Both domestic in warmer drylands, with potentially devastat-
(18, 19) and support arguments for conserving and wild herbivore species can exhibit strong ing implications for the fate of these ecosys-
and restoring diverse plant communities to feeding niche differences (24, 25); thus, in- tems [e.g., increased land degradation and
prevent land degradation, increase forage pro- creasing their diversity can enhance ecosys- desertification (17)] and their inhabitants [e.g.,
duction, and mitigate climate change in grazed tem functioning (25). Despite a renewed in- greater poverty, migration, and/or social un-
drylands (20). terest in mixed-species grazing, studies have rest (28)]. Although dryland pastoralists have
Mammalian herbivore richness, which was been conducted at only a handful of sites or historically adopted strategies to cope with
selected in 33% of the best-fitting models with a limited suite of herbivores (25–27). Our environmental uncertainty (e.g., nomadism,
(fig. S12), was positively related to multiple findings provide empirical evidence of the po- transhumance), benefits of these strategies
ecosystem services. Greater herbivore rich- tential benefits of increasing herbivore rich- will wane if livestock concentrates in partic-
ness positively correlated with soil carbon ness to enhance the delivery of key ecosystem ular areas as a result of resource scarcity or
storage regardless of grazing pressure (fig. services across contrasting environmental and droughts (29).

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Fig. 4. Geographical variation in the effect of grazing pressure on ecosystem (lnRR) (13). Predictions were made using plant species richness, mean annual
services across global drylands. (A to L) For each of the 98 sites surveyed, temperature, and rainfall seasonality; all other parameters were fixed at their mean
the effect of grazing pressure on ecosystem services predicted by model parameters value (13). For simplicity, grazing effects were averaged across all ecosystem
is plotted along the wide climatic and plant species richness gradients that were services [(A), (E), and (I)] and across regulating [(B), (F), and (J)], supporting [(C),
evaluated. To do this, each ecosystem service at low and high grazing pressures was (G), and (K)], and provisioning [(D), (H), and (L)] services. Blue and red dots indicate
first predicted using predictor estimates of the best-fitting models (see tables S13 the most-negative and most-positive effects of grazing, respectively. See figs. S16
to S15). Then, the predicted effect of grazing at each site was calculated as the and S17 for detailed results on each service and Fig. 2 for the meaning of the
difference between high and low grazing pressure levels using a log response ratio symbols depicting each ecosystem service.

Our findings underscore the importance of bivore richness and the provision of multiple 8. J. J. Gaitán et al., Land Degrad. Dev. 29, 210–218 (2018).
accounting for interactions between grazing ecosystem services across contrasting environ- 9. P. D’Ottavio et al., Grass Forage Sci. 73, 15–25 (2018).
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ing ecosystem service delivery in drylands. and grazing pressure levels. Our work ad- 24, 2054–2064 (2021).
They also illustrate those climate change and dresses a key knowledge gap that can lead to 12. P. Manzano et al., One Earth 4, 651–665 (2021).
biodiversity loss drivers that are the most better management of drylands, the largest 13. See materials and methods.
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(2021). talent of Comunidad de Madrid (no. 2017‐T2/ AMB‐5406). M.A.B. J.J.G., S.A., B.J.M., C.P., P.D.-M., A.Re., A.Le., M.C.R., S.C., N.E.,
23. J. E. Duffy, C. M. Godwin, B. J. Cardinale, Nature 549, 261–264 acknowledges support from the School of Forestry and College of E.M.-J., O.S., M.A., N.A., C.L.A., V.A., F.A., T.A., R.J.A., K.B., F.B.S.,
(2017). the Environment, Forestry and Natural Sciences of Northern N.B., B.B., M.A.B., D.B., C.B., R.Ca., A.P.C.-M., H.C., I.C., P.C.-Q.,
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25. L. Wang et al., Proc. Natl. Acad. Sci. U.S.A. 116, 6187–6192 (2019). Nacional de Ciencia y Tecnología (SEP-CB-2015-01-251388, PN A.F., M.F., D.F., A.S.K.F., L.H.F., L.A.G., A.C.G., E.G.-M., R.M.H.-H.,
26. J. P. G. M. Cromsigt et al., Philos. Trans. R. Soc. London Ser. B 2017-5036 and PRONAII 319059). F.M.H. acknowledges support N.H., E.H.-S., F.M.H., O.J.-M., F.J., A.J., K.F.K., M.K., J.E.K., C.V.L.,
373, 20170440 (2018). from the National Council for Scientific and Technological A.Li., P.C.l.R., X.L., P.L., J.L., M.A.L., G.M.-K., T.P.M., O.M.I., A.J.M.,
27. N. Pettorelli, S. M. Durant, J. T. du Toit, Eds., Rewilding Development (CNPq - PCI/INMA) of the Brazilian Ministry of E.M., J.P.M., G.M., S.M.M., A.N., G.O., G.R.O., G.P., M.O.D.P., Y.P.,
(Cambridge Univ. Press, 2019). Science, Technology and Innovation (MCTI, processes number R.E.Q., S.R., S.C.R., P.J.R., B.R., A.Ro., V.R., J.G.R., J.C.R., A.S.,
28. C. Almer, J. Laurent-Lucchetti, M. Oechslin, J. Environ. Econ. 302381/2020-1). H.L.T. acknowledges support from the US M.A.S., S.S., I.S., C.R.A.S., A.M.S., A.L.T., A.D.T., H.L.T., K.T., S.T.,
Manage. 86, 193–209 (2017). National Science Foundation (NSF) (DEB 0953864). A.N. and M.K. J.V., O.V., L.v.d.B., S.V.A., F.V., W.W., D.W., L.W., G.M.W., L.Y., E.Z.,
29. S. A. Mousavi, M. Sarshad Ghahfarokhi, S. Soltani Koupaei, acknowledge support from the Fundação para a Ciência e a Y.Z., X.Z., N.G.; Formal analysis: N.G., Y.L.B.-P., M.B., D.J.E., E.V.,
Ecol. Indic. 110, 105946 (2020). Tecnologia (SFRH/BD/130274/2017, CEECIND/02453/2018/ C.A.G., J.-T.W., H.-W.H., J.-Z.H.; Resources: F.T.M., Y.L.B.-P., D.J.E.,
30. T. Fetzel et al., Global Biogeochem. Cycles 31, 1089–1102 (2017). CP1534/CT0001, PTDC/ASP-SIL/7743/2020 and UIDB/00329/ H.S., M.B., B.G., V.O., M.G.-G., E.V., J.J.G., S.A., B.J.M., C.P.,
31. F. T. Maestre et al., Data and R code from “Grazing and 2020). A.A.C. acknowledges support from the Coordenação de P.D.-M., A.Re., H.-W.H., J.-Z.H., J.-T.W., A.Le., M.C.R., S.C., N.E.,
ecosystem service delivery in global drylands,” figshare (2022); Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - E.M.-J., O.S., M.A., N.A., C.L.A., V.A., F.A., T.A., R.J.A., K.B., F.B.S.,
https://doi.org/10.6084/m9.figshare.14923065.v1. Finance Code 001. J.E.K. and T.P.M. acknowledge the National N.B., B.B., M.A.B., D.B., C.B., R.Ca., A.P.C.-M., H.C., I.C., P.C.-Q.,
32. B. K. Singh, J.-T. Wang, M. Delgado-Baquerizo, F. T. Maestre, Research Foundation of South Africa (grant no. 114412). F.J. and R.Ch., A.A.C., C.M.C., A.D.-N., B.D., D.A.D., A.J.D., J.D., B.E., C.I.E.,
16S and 18S data from “Grazing and ecosystem service N.B. acknowledge support from the German Federal Ministry of A.F., M.F., D.F., A.S.K.F., L.H.F., L.A.G., A.C.G., E.G.-M., R.M.H.-H.,
delivery in global drylands,” figshare (2022); Education and Research (BMBF) in the framework of the SPACES N.H., E.H.-S., F.M.H., O.J.-M., F.J., A.J., K.F.K., M.K., J.E.K., C.V.L.,
https://doi.org/10.6084/m9.figshare.20131355.v1. projects OPTIMASS (FKZ: 01LL1302A) and ORYCS (FKZ: A.Li., P.C.l.R., X.L., P.L., J.L., M.A.L., G.M.-K., T.P.M., O.M.I., A.J.M.,
01LL1804A). A.Li. and A.S.K.F. acknowledge support from the E.M., J.P.M., G.M., S.M.M., A.N., G.O., G.R.O., G.P., M.O.D.P., Y.P.,

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ACKN OW LEDG MEN TS German Federal Ministry of Education and Research (BMBF) in the R.E.Q., S.R., S.C.R., P.J.R., B.R., A.Ro., V.R., J.G.R., J.C.R., A.S.,
We acknowledge S. Undrakhbold, M. Uuganbayar, B. Byambatsogt, framework of the SPACES projects Limpopo Living Landscapes M.A.S., S.S., I.S., C.R.A.S., A.M.S., A.L.T., A.D.T., H.L.T., K.T.,
S. Khaliun, S. Solongo, B. Batchuluun, M. Sloan, J. Spence, (FKZ: 01LL1304D) and SALLnet (FKZ: 01LL1802C). L.W. S.T., J.V., O.V., L.v.d.B., S.V.A., F.V., W.W., D.W., L.W., G.M.W.,
E. Geiger, I. Souza, R. Onoo, T. Araújo, M. Mabaso, P. M. Lunga, acknowledges support from the US NSF (EAR 1554894). L.H.F. L.Y., E.Z., Y.Z., X.Z., B.K.S., N.G., M.D.-B. ; Funding acquisition:
L. Eloff, P. Eloff, J. Sebei, J. Joordan, E. Mudongo, V. Mokoka, acknowledges support from the Natural Sciences and Engineering F.T.M., N.G., N.E., Y.L.B.-P., B.B., B.E., J.D., A.Ro., C.P., M.A.B.,
B. Mokhou, T. Maphanga, F. Hoffmann, R. Peters, A. Lozada, E. Vidal, Research Council of Canada Industrial Research Chair Program in E.H.-S., F.M.H., H.L.T., A.N., M.K., J.E.K., T.P.M., F.J., N.B., A.Li., A.S.K.F.,
F. Perrona, R. Ledezma, R. Matjea, L. Kindermann, C. Goebel, Ecosystem Reclamation. S.C.R. acknowledges support from the L.W., L.H.F., S.C.R., G.M.W., K.T., A.F., C.B., A.J.; Project
B. Semple, and B. Tamayo for assistance with field work. We thank US Geological Survey Ecosystems Mission Area and the US Bureau administration: F.T.M., N.G., Y.L.B.-P., H.S., E.V., J.M.-V., V.O., B.G.;
Bush Heritage Australia; the University of Limpopo; the Ministry of of Land Management. G.M.W. acknowledges support from the Software: N.G., Y.L.B.-P.; Supervision: F.T.M.; Validation: M.B.,
Environment and Tourism (Namibia); the Ministry of Agriculture, Australian Research Council. L.v.d.B. and K.T. acknowledge support C.P.; Visualization: N.G., Y.L.B.-P., D.J.E., E.G., M.B., E.V.; Writing –
Water and Land Reform (Namibia); Corporación Nacional Forestal from the German Research Foundation (DFG) priority research original draft: F.T.M., N.G., Y.L.B.-P., D.J.E., H.S., M.D.-B.; Writing –
(CONAF) and the agricultural community Quebrada de Talca (Chile); program SPP-1803 “EarthShape: Earth Surface Shaping by Biota” review and editing: F.T.M., N.G., Y.L.B.-P., D.J.E., M.D.-B., H.S.,
and the South African military for granting research permissions (TI 338/14-1). M.D.-B. acknowledges support from the Spanish M.B., B.G., V.O., E.G., J.M.-V., M.G.-G., E.V., E.G., J.J.G., S.A., B.J.M.,
and/or granting access to their research farms and properties. We Ministry of Science and Innovation for the I+D+i project PID2020- C.P., P.D.-M., A.Re., H.-W.H., J.-Z.H., J.-T.W., A.Le., M.C.R., S.C.,
also thank B. Benito for his revisions to the R code, and C. Abel for 115813RA-I00 funded by MCIN/AEI/10.13039/501100011033. N.E., E.M.-J., O.S., M.A., N.A., C.L.A., V.A., F.A., T.A., R.J.A., K.B.,
her comments on the manuscript. Any use of trade, product, or firm M.D.-B. is also supported by a project of the Fondo Europeo de F.B.S., N.B., B.B., M.A.B., D.B., C.B., R.Ca., A.P.C.-M., H.C., I.C.,
names in this paper is for descriptive purposes only and does not Desarrollo Regional (FEDER) and the Consejería de Transformación P.C.-Q., R.Ch., A.A.C., C.M.C., A.D.-N., B.D., D.A.D., A.J.D., J.D., B.E.,
imply endorsement by the US government. Funding: This research Económica, Industria, Conocimiento y Universidades of the Junta C.I.E., A.F., M.F., D.F., A.S.K.F., L.H.F., L.A.G., A.C.G., E.G.-M.,
was funded by the European Research Council [ERC grant agreement de Andalucía (FEDER Andalucía 2014-2020 Objetivo temático R.M.H.-H., N.H., E.H.-S., F.M.H., O.J.-M., F.J., A.J., K.F.K., M.K.,
647038 (BIODESERT)] and Generalitat Valenciana (CIDEGENT/2018/ “01 - Refuerzo de la investigación, el desarrollo tecnológico y la J.E.K., C.V.L., A.Li., P.C.l.R., X.L., P.L., J.L., M.A.L., G.M.-K., T.P.M.,
041). F.T.M. acknowledges support from a Rei Jaume I Award, the innovación”) associated with the research project P20_00879 O.M.I., A.J.M., E.M., J.P.M., G.M., S.M.M., A.N., G.O., G.R.O., G.P.,
Alexander von Humboldt Foundation, and the Synthesis Center (sDiv) (ANDABIOMA). P.J.R. and A.J.M. acknowledge support from Fondo M.O.D.P., Y.P., R.E.Q., S.R., S.C.R., P.J.R., B.R., A.Ro., V.R., J.G.R.,
of the German Centre for Integrative Biodiversity Research Halle– Europeo de Desarrollo Regional through the FEDER Andalucía J.C.R., A.S., M.A.S., S.S., I.S., C.R.A.S., A.M.S., A.L.T., A.D.T.,
Jena–Leipzig (iDiv). C.A.G., S.C., and N.E. acknowledge support from operative program, FEDER-UJA 1261180 project. A.F. thanks ANID H.L.T., K.T., S.T., J.V., O.V., L.v.d.B., S.V.A., F.V., W.W., D.W., L.W.,
iDiv and the Deutsche Forschungsgemeinschaft (DFG– FZT 118, PIA/BASAL FB210006 and Millennium Science Initiative Program G.M.W., L.Y., E.Z., Y.Z., X.Z., B.K.S. Competing interests: The
202548816; Flexpool proposal 34600850). Y.L.B.-P. was supported by NCN2021-050. A.J. acknowledges support from the Bavarian authors declare that they have no competing interests. Data and
a Marie Sklodowska-Curie Actions Individual Fellowship (MSCA-IF) Research Alliance Germany (BayIntAn_UBT_2017_61). C.B. materials availability: All the data used in this article and the
within the European Program Horizon 2020 (DRYFUN Project acknowledges the National Natural Science Foundation of China R scripts used to generate the main results of the study are
656035). N.G. was supported by CAP 20-25 (16-IDEX-0001) and (grant no. 41971131). Biodiversity and ecosystem function research available through figshare (31). The raw sequence data generated
the AgreenSkills+ fellowship program, which has received funding in the B.K.S. laboratory is funded by the Australian Research in this study are also available through figshare (32). License
from the EU’s Seventh Framework Programme under grant agreement Council (DP210102081). Any use of trade, product, or firm names information: Copyright © 2022 the authors, some rights reserved;
N° FP7-609398 (AgreenSkills+ contract). B.B. and B.E. were in this paper is for descriptive purposes only and does not imply exclusive licensee American Association for the Advancement of
supported by the Taylor Family–Asia Foundation Endowed Chair in endorsement by the US government. H.S. is supported by a Science. No claim to original US government works. https://www.
Ecology and Conservation Biology. J.D., A.Ro., and H.C. acknowledge María Zambrano fellowship funded by the Ministry of Universities science.org/about/science-licenses-journal-article-reuse
support from the Fundação para a Ciência e a Tecnologia (IF/00950/ and European Union-Next Generation plan. G.P. and C.V.L.
2014 and 2020.03670.CEECIND, SFRH/BDP/108913/2015, and in the acknowledge support from Universidad Nacional de Río Negro SUPPLEMENTARY MATERIALS
scope of the framework contract foreseen in the numbers 4-6 of the (PI 40-C-873 and 654). V.R. acknowledges support from the
science.org/doi/10.1126/science.abq4062
article 23, of the Decree-Law 57/2016, August 29, changed by Law Regional Government of Extremadura (Spain) through a “Talento”
Materials and Methods
57/2017, July 19, respectively), as well as from the MCTES, FSE, UE, fellowship (TA18022). M.F. acknowledges support from the
Figs. S1 to S19
and the CFE (UIDB/04004/2020) research unit financed by Department of Range and Watershed Management, Ferdowsi
Tables S1 to S28
Fundação para a Ciência e a Tecnologia/MCTES through national University of Mashhad, Mashhad, Iran. Participation of recent
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Science, 378 (6622), • DOI: 10.1126/science.abq4062

Complex effects of livestock

Livestock grazing provides food and livelihoods for billions of people but at the cost of ecosystem degradation in many
places. Maestre et al. investigated how grazing by livestock and native herbivores affects ecosystem functions and
services and how these effects vary with climate, soil properties, and biodiversity (see the Perspective by Ganguli
and O’Rourke). Using a replicated survey at 98 dryland sites spanning six continents, the authors found that grazing
effects on ecosystem services often depend on other factors. Interactions between grazing and climate were especially
important; warmer sites had lower rates of carbon storage, organic matter deposition, and erosion control under high
(but not low) grazing pressure. —BEL

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