The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands.

Bent Creek Experimental Forest USDA Forest Service Asheville NC USA

Bilogo Dpto Conservacin y Manejo Parque Nacional Lanin Elordi y Perito Moreno San Marten de los Andes Neuqun Argentina

CEFE Universite Montpellier CNRS EPHE IRD Montpellier France

Centre d'Ecologie Fonctionnelle et Evolutive Montpellier France

Centre de Recerca Ecologica i Aplicacions Forestals Bellaterra Catalunya Spain

Centro de Investigacion Forestal Madrid Spain

CREA Research Ventre for Forestry and Wood Arezzo Italy

Department of Agricultural and Environmental Sciences Production Territory Agroenergy University of Milan Milano Italy

Department of Agriculture Forest and Food Sciences University of Torino Grugliasco Torino Italy

Department of Biodiversity Ecology and Evolution Complutense University of Madrid Madrid Spain

Department of Biologia Vegetal y Ecologia Universidad de Sevilla Sevilla Spain

Department of Biological Environment Akita Prefectural University Akita Japan

Department of Biological Sciences and Center for Adaptive Western Landscapes Flagstaff AZ USA

Department of Biological Sciences DePaul University Chicago IL USA

Department of Biological Sciences National Sun Yat sen University Kaohsiung Taiwan

Department of Biological Sciences Northern Arizona University Flagstaff AZ USA

Department of Biology Colby College Waterville ME USA

Department of Biology University of New Mexico Albuquerque NM USA

Department of Biology Wake Forest University Winston Salem NC USA

Department of Biology Washington University in St Louis St Louis MO USA

Department of Biology Wilkes University Wilkes Barre PA USA

Department of Ecology Evolution and Environmental Biology Columbia University New York NY USA

Department of Ecology Instituto de Investigaciones en Biodiversidad y Medioambiente Bariloche Argentina

Department of Ecosystem Science and Management Pennsylvania State University University Park PA USA

Department of Environmental Science and Ecology State University of New York Brockport Brockport NY USA

Department of Environmental Sciences University of Puerto Rico Rio Piedras PR USA

Department of Environmental Studies University of California Santa Cruz CA USA

Department of Forest and Rangeland Stewardship Colorado State University Fort Collins CO USA

Department of Forest Ecology Silva Tarouca Research Institute Brno Czech Republic

Department of Forest Vegetation Forestry and Forest Products Research Institute Tsukuba Japan Ibaraki

Department of Forestry and Renewable Forest Resources Biotechnical Faculty University of Ljubljana Ljubljana Slovenia

Department of Geography and Planning School of Environmental Sciences University of Liverpool Liverpool UK

Department of Natural Sciences Manchester Metropolitan University Manchester UK

Department of Plant Ecology Forestry and Forest Products Research Institute Tsukuba Ibaraki Japan

Department of Systematic Zoology Faculty of Biology Adam Mickiewicz University Poznan Poland

Department of Wildland Resources and the Ecology Center Utah State University Logan UT USA

Earth and Environment Boston University Boston MA USA

Eastern Forest Environmental Threat Assessment Center USDA Forest Service Southern Research Station Research Triangle Park NC USA

Ecological Restoration Institute Northern Arizona University Flagstaff AZ USA

Facultad de Ciencias Forestales y de la Conservacion de la Naturaleza Universidad de Chile La Pintana Santiago Chile

Forest Research Institute University of Quebec in Abitibi Temiscamingue Rouyn Noranda Quebec Canada

Forest Resources University of Washington Seattle WA USA

Health and Environmental Sciences Department Xian Jiaotong Liverpool University Suzhou China

INRAE Bordeaux Sciences Agro Villenave d'Ornon France

INRAE UR EFNO Nogent sur Vernisson France

Institut National de Recherche pour Agriculture Alimentation et Environnement Ecologie des Forets Mediterranennes Avignon France

Institute of Arctic Biology University of Alaska Fairbanks AK USA

Institute of Forest Ecology Department of Forest and Soil Sciences University of Natural Resources and Life Sciences Wien Austria

Instituto de Recursos Naturales y Agrobiologia de Sevilla Consejo Superior de Investigaciones Cientificas Seville Andalucia Spain

Instituto Pirenaico de Ecologla Consejo Superior de Investigaciones Cientificas Zaragoza Spain

Laboratoire Evolution et Diversite Biologique Toulouse France

Mammal Research Institute Polish Academy of Sciences Bialowieza Poland

Natural Resources Cornell University Ithaca NY USA

Nicholas School of the Environment Duke University Durham NC USA

NRAE Aix Marseille University UMR RECOVER Aix en Provence France

Pacific Forestry Centre Victoria British Columbia Canada

Research Centre for Forestry and Wood Arezzo Italy

School for Environment and Sustainability University of Michigan Ann Arbor MI USA

School of Life Sciences Keele University Staffordshire UK

School of Natural Sciences UC Merced Merced CA USA

Smithsonian Tropical Research Institute Balboa Republic of Panama

Tohoku Research Center Forestry and Forest Products Research Institute Morioka Iwate Japan

U S Geological Survey Fort Collins Science Center Fort Collins CO USA

U S Geological Survey Western Ecological Research Center Three Rivers CA USA

Universite Bordeaux Institut National de Recherche pour Agriculture Alimentation et Environnement Pessac France

Universite Grenoble Alpes Institut National de Recherche pour Agriculture Alimentation et Environnement St Martin d'Heres France

Universite Paris Saclay Centre national de la recherche scientifique AgroParisTech Ecologie Systematique et Evolution Orsay France

USDA Forest Service Southern Research Station Stoneville MS USA

Valles Caldera National Preserve National Park Service Jemez Springs NM USA

W Szafer Institute of Botany Polish Academy of Sciences Krakow Poland

Washington University in Saint Louis Center for Conservation and Sustainable Development Missouri Botanical Garden St Louis MO USA

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