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12 páginas.- 4 figuras.- referencias.- Additional supporting information can be found online in the Supporting Information section at the end of this article.- Full Access in https://onlinelibrary.wiley.com/doi/full/10.1111/geb.13652, Aim Our understanding of the mechanisms that maintain forest diversity under changing climate can benefit from knowledge about traits that are closely linked to fitness. We tested whether the link between traits and seed number and seed size is consistent with two hypotheses, termed the leaf economics spectrum and the plant size syndrome, or whether reproduction represents an independent dimension related to a seed size–seed number trade-off. Location Most of the data come from Europe, North and Central America and East Asia. A minority of the data come from South America, Africa and Australia. Time period 1960–2022. Major taxa studied Trees. Methods We gathered 12 million observations of the number of seeds produced in 784 tree species. We estimated the number of seeds produced by individual trees and scaled it up to the species level. Next, we used principal components analysis and generalized joint attribute modelling (GJAM) to map seed number and size on the tree traits spectrum. Results Incorporating seed size and number into trait analysis while controlling for environment and phylogeny with GJAM exposes relationships in trees that might otherwise remain hidden. Production of the large total biomass of seeds [product of seed number and seed size; hereafter, species seed productivity (SSP)] is associated with high leaf area, low foliar nitrogen, low specific leaf area (SLA) and dense wood. Production of high seed numbers is associated with small seeds produced by nutrient-demanding species with softwood, small leaves and high SLA. Trait covariation is consistent with opposing strategies: one fast-growing, early successional, with high dispersal, and the other slow-growing, stress-tolerant, that recruit in shaded conditions. Main conclusions Earth system models currently assume that reproductive allocation is indifferent among plant functional types. Easily measurable seed size is a strong predictor of the seed number and species seed productivity. The connection of SSP with the functional traits can form the first basis of improved fecundity prediction across global forests., The project has been funded by grants to J.S.C. from the National Science Foundation, most re-cently DEB-1754443, and by the Belmont Forum (1854976), NASA (AIST16-0052 and AIST18-0063) and the Programme d'Investissement d'Avenir under project FORBIC (18-MPGA-0004; Make Our Planet Great Again). Jerry Franklin's data remain acces-sible through NSF LTER DEB-1440409. Data from Hubbard Brook (New Hampshire) were funded through NSF-LTER. Puerto Rico data were funded by NSF grants, most recently DEB 0963447 and LTREB 11222325. Data from the Andes Biodiversity and Ecosystem Research Group were funded by the Gordon and Betty Moore Foundation and NSF 727 LTREB 1754647. M.B. was supported by grant no. 2019/35/D/NZ8/00050 from the (Polish) National Science Centre and by Polish National Agency for Academic Exchange Bekker programme PPN/BEK/2020/1/00009/U/00001. Research by the USDA Forest Service and the USGS was funded by these agencies. Any use of trade, firm or product names does not imply endorsement by the US Government. Any use of trade, firm, or product names is for descriptive purposes only and does not imply en -dorsement by the U.S. Government

Made available in DSpace on 2022-04-28T19:52:06Z (GMT). No. of bitstreams: 0 Previous issue date: 2022-03-18 Urbanization transforms environments in ways that alter biological evolution. We examined whether urban environmental change drives parallel evolution by sampling 110,019 white clover plants from 6169 populations in 160 cities globally. Plants were assayed for a Mendelian antiherbivore defense that also affects tolerance to abiotic stressors. Urban-rural gradients were associated with the evolution of clines in defense in 47% of cities throughout the world. Variation in the strength of clines was explained by environmental changes in drought stress and vegetation cover that varied among cities. Sequencing 2074 genomes from 26 cities revealed that the evolution of urban-rural clines was best explained by adaptive evolution, but the degree of parallel adaptation varied among cities. Our results demonstrate that urbanization leads to adaptation at a global scale. Department of Biology University of Toronto Mississauga ON Centre for Urban Environments University of Toronto Mississauga ON Department of Biology University of North Carolina, Chapel Hill Department of Biology University of Louisiana LA Department of Biology Queen's University ON Department of Biology Concordia University QC Department of Biological Sciences DePaul University Department of Biology DePauw University IN Department of Urban Design and Planning, University of Washington, Seattle, WA, USA Colegio de Ciencias Biológicas y Ambientales Universidad San Francisco de Quito USFQ Department of Genetics University of Georgia Department of Ecology and Genetics Evolutionary Biology Centre Uppsala University Field Science Center for Northern Biosphere Hokkaido University Natural History Museum Zoology University of New England NSW Programa de Pós-Graduação em Geografia da UFMT campus de Rondonópolis Department of Botany and Biodiversity Research Centre University of British Columbia Graduate Program in Genome Sciences and Technology Genome Sciences Centre University of British Columbia Department of Microbiology and Immunology University of British Columbia Red de Biología Evolutiva Instituto de Ecología A. C. School of the Environment Yale University CT Departamento de Ciencias Ecológicas Universidad de Chile, Facultad de Ciencias Instituto de Ecología y Biodiversidad Universidad de Chile Department of Biology Mount Allison University Red de Ecoetología Instituto de Ecología A. C. Department of Biology University of Ottawa ON Department of Zoology University of Cambridge Department of Biology, Washington University in St. Louis, St. Louis, MO, USA Department of Biology University of Miami FL Centro de Investigación en Recursos Naturales y Sustentabilidad (CIRENYS) Universidad Bernardo O'Higgins Department of Biology, University of La Verne, La Verne, CA, USA Département des sciences du bois et de la forêt Université Laval QC Evolution & Ecology Research Centre School of Biological Earth and Environmental Sciences UNSW Sydney NSW Department of Biology Ghent University Department of Biology Monmouth University West Long Branch Centre for Ecology Evolution and Environmental Changes Faculdade de Ciências Universidade de Lisboa Lisboa Department of Biology KU Leuven School of Agriculture and Environment Wildlife and Ecology group Massey University, Palmerston North Department of Biological Sciences University of Cape Town Institute of Landscape Ecology University of Münster Gosnell School of Life Sciences, Rochester Institute of Technology, Rochester, NY, USA Department of Biological Sciences University of Alberta AB Louis Calder Center and Department of Biological Sciences, Fordham University, Armonk, NY, USA Departamento de Ecología Tropical, Universidad Autónoma de Yucatán, Mérida, Yucatán, México School of Life Sciences University of Sussex Department of Ecology Environment and Plant Sciences Stockholm University iBIOTROP Instituto de Biodiversidad Tropical Universidad San Francisco de Quito Department of Biology, San Francisco State University, San Francisco, CA, USA Unidad de Recursos Naturales, Centro de Investigación Científica de Yucatán AC, Mérida, Yucatán, México School of Ecological and Environmental Sciences East China Normal University Shanghai Engineering Research Center of Sustainable Plant Innovation Centre for Ecosystem Science School of Biological Earth and Environmental Sciences UNSW Sydney NSW Department of Ecology and Evolutionary Biology University of Michigan MI Department of Biosciences Rice University TX IFEVA Universidad de Buenos Aires Facultad de Agronomía, CONICET Biology Department Saint Mary's University NS Department of Biological Sciences, Universidad de los Andes Department of Biology and Biochemistry University of Houston TX Université de Rennes Department of Zoology and Biodiversity Research Centre University of British Columbia Department of Environmental Studies Dordt University Sioux Center IA Department of Biology Minneapolis Community and Technical College MN Department of Natural Sciences Ecology and Environment Research Centre Manchester Metropolitan University Instituto de Investigaciones en Ecosistemas y Sustentabilidad UNAM Department of Botany School of Biology Aristotle University of Thessaloniki Faculty of Biological and Environmental Science Organismal & Evolutionary Biology Research Programme University of Helsinki Institute of Biodiversity Animal Health and Comparative Medicine University of Glasgow Department of Biology Hendrix College Department of Ecological Science Vrije Universiteit Amsterdam Departamento de Ciencias Biológicas y Agropecuarias Universidad Técnica Particular de Loja Departamento de Biologia Universidade Federal de Santa Maria (UFSM) Department of Plant Sciences School of Biology College of Science University of Tehran NTNU University Museum Norwegian University of Science and Technology Red de Estudios Moleculares Avanzados Instituto de Ecología A. C. School of Biological Sciences, University of Reading, Whiteknights Park, Reading, Berkshire, UK Department of Biology Northern Arizona University AZ Department of Biological Sciences MacEwan University AB Max Planck Institute for Plant Breeding Research Departamento de Ecología Evolutiva Instituto de Ecología Universidad Nacional Autónoma de México Max Planck Institute of Molecular Plant Physiology Potsdam-Golm BIO5 Institute University of Arizona AZ Alaska Center for Conservation Science University of Alaska Anchorage Tropical Diversity, Royal Botanical Garden of Edinburgh Département de biologie Université de Moncton Department of Biological Sciences University of Manitoba MB Departments of Microbiology & Statistics University of Manitoba MB Department of Biology University of New Brunswick Department of Biology Kalamazoo College MI BioProtection Research Centre Lincoln University Departamento de Ciencias Facultad de Artes Liberales Universidad Adolfo Ibáñez Department of Ecology Evolution Behaviour University of Minnesota MN Department of Biological Sciences Brock University Department of Environmental Toxicology, University of California, Davis, CA, USA ICB - University of Talca School of Molecular and Life Science Curtin University College of Science Health Engineering and Education Murdoch University, Murdoch School of Life and Environmental Sciences University of Sydney NSW School of Biological Sciences, Monash University, Melbourne, VIC, Australia Department of Biological Sciences Wayne State University MI Department of Biology Western Oregon University, OR School of Natural Resources and the Environment University of Arizona AZ Departamento de Ecología Humana, Cinvestav Mérida Departamento de Ciencias Biológicas y Departamento de Ecología y Biodiversidad Facultad de Ciencias de la Vida, Universidad Andrés Bello Institute of Ecology and Biodiversity (IEB) Department of Biology Lund University Department of Biology Norwegian University of Science and Technology Escuela Superiro de Desarrollo Sustentable Universidad Autónoma de Guerrero -CONACYT Clarkson Secondary School Peel District School Board ON Homelands Sr. Public School Peel District School Board ON Department of Biological Sciences University of Illinois at Chicago Dufferin-Peel Catholic District School Board, St. James Catholic Global Learning Centre Department of Biosciences University of Calgary AB Ecological Processes Branch U.S. Army ERDC-CERL Department of Biology, Oberlin College, Oberlin, OH, USA Escuela Nacional de Estudios Superiores Unidad Morelia UNAM Institute of Evolution and Ecology University of Tübingen Tübingen Department of Evolutionary Biology and Environmental Studies University of Zurich, Winterthurerstrasse Urban Wildlife Institute Department of Conservation and Science, Lincoln Park Zoo Departamento de Ecología Universidad Católica de la Santísima Concepción Department of Biological Sciences University of Denver CO Department of Biological Sciences Mississippi State University MS Department of Biology Center for Computational & Integrative Biology Rutgers University-Camden Kunming Institute of Botany Chinese Academy of Sciences Department of Chemistry & Biochemistry Laurentian University ON Ministry of Education Key Laboratory for Biodiversity Science and Ecological Engineering College of Life Sciences Beijing Normal University School of BioSciences, University of Melbourne, Melbourne, VIC, Australia Posgrado en Ciencias Biológicas Universidad Nacional Autónoma de México Department of Biological Sciences, Auburn University, Auburn, AL, USA Department of Entomology and Nematology, University of California, Davis, CA, USA Department of Biology University of New Mexico NM Department of Biology University of Wisconsin - Eau Claire Agriculture Institute Iranian Research Organization for Science and Technology (IROST) Department of Biology Colby College ME Instituto de Biología Universidad de Antioquia Department of Biology University of Massachusetts Boston MA Agricultural Biology Colorado State University CO Departamento de Biología Vegetal y Ecología Facultad de Biología Universidad de Sevilla, Av. Reina Mercedes s/n Facultad de Estudios Interdisciplinarios Centro GEMA- Genómica Universidad Mayor Evolutionary Ecology Group Naturalis Biodiversity Center Department of Biology and Chemistry Nipissing University ON, North Bay Center for Ecological Research Kyoto University Bonanza Creek Long Term Ecological Research Program University of Alaska Fairbanks Department of Botany and Molecular Evolution Senckenberg Research Institute and Natural History Museum Frankfurt Departamento de Biodiversidade Instituto de Biociências Univ Estadual Paulista - UNESP Nelson Institute for Environmental Studies University of Wisconsin-Madison WI Department of Biology, California State University, Northridge, Los Angeles, CA, USA Department of Ecology Swedish University of Agricultural Sciences Facultad de Ciencias y Biotecnologia Universidad CES Department of Biology Hofstra University Faculty of Biosciences and Aquaculture Nord University, Bodø Division of Biological Sciences, University of California San Diego, San Diego, CA, USA Department of Biology University of Richmond VA Estación de Biodiversidad Tiputini Colegio de Ciencias Biológicas y Ambientales Universidad San Francisco de Quito USFQ Department of Biological Sciences Institute of Environment Florida International University FL Agronomy Department University of Almería Department of Biological Sciences and Center for Urban Ecology and Sustainability Butler University IN Department of Biological Sciences Louisiana State University LA Faculty of Biological Sciences, Goethe University Frankfurt Institute of Biology Leiden Leiden University Department of Biological and Environmental Science University of Jyväskylä Jyväskylä Department of Biology University of Louisville KY Organization for Programs on Environmental Science University of Tokyo CNRS AgroParisTech Ecologie Systématique et Evolution, Université Paris-Saclay Department of Biology, Providence College, Providence, RI, USA General Zoology Institute for Biology Martin Luther University Halle-Wittenberg International Arctic Research Center University of Alaska Fairbanks Science, Technology and Society Department, Rochester Institute of Technology, Rochester, NY, USA SLU Swedish Species Information Centre Swedish University of Agricultural Sciences Department of Biology Westfield State University MA Centre of New Technologies University of Warsaw Department of Biology, Stanford University, Stanford, CA, USA Plant Biology Department Michigan State University MI Biology Department Davidson College College of Horticulture and Forestry Sciences/ Hubei Engineering Technology Research Center for Forestry Information Huazhong Agricultural University School of Life Sciences Technical University of Munich School of Life Sciences Lanzhou University Institute of Ecology and Evolution University of Bern Department of Evolution Ecology and Behaviour University of Liverpool Departamento de Biodiversidade Instituto de Biociências Univ Estadual Paulista - UNESP

Populations of many tree species exhibit synchronous and highly temporally variable seed crops across years. This is called mast seeding, and there are two predominant hypotheses for this pattern of reproduction: pollination efficiency and seed-predator satiation. Mast seeding studies typically involve records of population-level reproduction, with less information on the characteristics of reproductive structures. Here, we use data across 6 years (2012–2017), spanning a range of population-level cone conditions, to characterize (i) white spruce (Picea glauca (Moench) Voss) cone lengths and seeds per cone, and (ii) levels of seed predation. We quantified population-level cone production and collected 1399 cones from a total of 38 trees in the Huron Mountains, Michigan, USA. Linear mixed models showed that mean and minimum cone lengths varied significantly across years; both being longest during the greatest cone production year. Larger cones had more seeds and the slopes of the relationships as well as the intercepts varied significantly across years. Generalized linear mixed models and Akaike’s information criterion model selection showed that cones with insect predation damage was greatest when population-level reproduction was the lowest, with a mean proportion of cones damaged 0.82 in that year. Our findings show that white spruce cone characteristics and losses to insect seed predation vary temporally, and follow expectations based on mast seeding hypotheses.

Ecological processes, such as migration and phenology, are strongly influenced by climate variability. Studying these processes often relies on associating observations of animals and plants with climate indices, such as the El Niño-Southern Oscillation (ENSO). A common characteristic of climate indices is the simultaneous emergence of opposite extremes of temperature and precipitation across continental scales, known as climate dipoles. The role of climate dipoles in shaping ecological and evolutionary processes has been largely overlooked. We review emerging evidence that climate dipoles can entrain species dynamics and offer a framework for identifying ecological dipoles using broad-scale biological data. Given future changes in climatic and atmospheric processes, climate and ecological dipoles are likely to shift in their intensity, distribution, and timing.

Resource pulses are rare events with a short duration and high magnitude that drive the dynamics of both plant and animal populations and communities1. Mast seeding is perhaps the most common type of resource pulse that occurs in terrestrial ecosystems2, is characterized by the synchronous and highly variable production of seed crops by a population of perennial plants3,4, is widespread both taxonomically and geographically5, and is often associated with nutrient scarcity6. The rare production of abundant seed crops (mast events) that are orders of magnitude greater than crops during low seed years leads to high reproductive success in seed consumers and has cascading impacts in ecosystems2,7. Although it has been suggested that mast seeding is potentially synchronized at continental scales8, studies are largely constrained to local areas covering tens to hundreds of kilometres. Furthermore, summer temperature, which acts as a cue for mast seeding9, shows patterns at continental scales manifested as a juxtaposition of positive and negative anomalies that have been linked to irruptive movements of boreal seed-eating birds10,11. Here, we show a breakdown in synchrony of mast seeding patterns across space, leading to asynchrony at the continental scale. In an analysis of synchrony for a transcontinental North America tree species spanning distances of greater than 5,200 km, we found that mast seeding patterns were significantly asynchronous at distances of greater than 2,000 km apart (all P < 0.05). Other studies have shown declines in synchrony across distance, but not asynchrony. Spatiotemporal variation in summer temperatures at the continental scale drives patterns of synchrony in mast seeding, and we anticipate that this affects the spatial dynamics of numerous seed-eating communities, from insects to small mammals to the large-scale migration patterns of boreal seed-eating birds. Previous studies showing that mast seeding was synchronized at large scales were constrained to hundreds of kilometres, but this continental-scale study shows that such events are asynchronous, driving spatial and temporal impacts for a wide range of species.

12 Pág., The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential., The project has been funded continuously since 1992 by National Science Foundation grants to J.S.C, most recently DEB-1754443, and by the Belmont Forum (1854976), NASA (AIST16-0052, AIST18-0063), and the Programme d’Investissement d’Avenir under project FORBIC (18-MPGA-0004)(Make Our Planet Great Again). Puerto Rico data were funded by NSF grants to M.U., most recently, DEB 0963447 and LTREB 11222325. Data from the Andes Biodiversity and Ecosystem Research Group were funded by the Gordon and Betty Moore Foundation and NSF LTREB 1754647 to M.S. Additional funding to M.Z. came from the W.Szafer Institute of Botany of the Polish Academy of Sciences and the Polish National Science Foundation (2019/33/B/NZ8/0134). M.B. was supported by the Polish National Science Centre grant no. 2019/35/D/NZ8/00050, and Polish National Agency for Academic Exchange Bekker programme PPN/BEK/2020/1/00009/U/00001. J.M.L. was supported by NSF grant DEB 1745496. Jerry Franklin’s data remain accessible through NSF LTER DEB-1440409. USDA Forest Service and USGS research was funded by those agencies.

12 páginas.- 5 figuras.- 67 referencias.- Additional supporting information may be found in the online version of the article at the publisher’s website .- The peer review history for this article is available at https://publons.com/publon/10.1111/ele.14012 .- All data and code supporting our results are archived on the Zenodo Repository at the following link: https://doi.org/10.5281/zenodo.6381799, Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics., The project has been funded by grants to JSC from the National Science Foundation, most recently DEB-1754443, and by the Belmont Forum (1854976), NASA (AIST16-0052, AIST18-0063) and the Programme d’Investissement d’Avenir under project FORBIC (18-MPGA-0004) (Make Our Planet Great Again). Jerry Franklin’s data remain accessible through NSF LTER DEB-1440409. Puerto Rico data were funded by NSF grants, most recently, DEB 0963447 and LTREB 11222325. Data from the Andes Biodiversity and Ecosystem Research Group were funded by the Gordon and Betty Moore Foundation and NSF LTREB 1754647. MB was supported by grant no. 2019/35/D/NZ8/00050 from the (Polish) National Science Centre, and Polish National Agency for Academic Exchange Bekker programme PPN/BEK/2020/1/00009/U/00001. Research by the USDA Forest Service and the USGS was funded by these agencies. Any use of trade, firm or product names does not imply endorsement by the US Government.

Significant gaps remain in understanding the response of plant reproduction to environmental change. This is partly because measuring reproduction in long-lived plants requires direct observation over many years and such datasets have rarely been made publicly available. Here we introduce MASTREE+, a data set that collates reproductive time-series data from across the globe and makes these data freely available to the community. MASTREE+ includes 73,828 georeferenced observations of annual reproduction (e.g. seed and fruit counts) in perennial plant populations worldwide. These observations consist of 5971 population-level time-series from 974 species in 66 countries. The mean and median time-series length is 12.4 and 10 years respectively, and the data set includes 1122 series that extend over at least two decades (≥20 years of observations). For a subset of well-studied species, MASTREE+ includes extensive replication of time-series across geographical and climatic gradients. Here we describe the open-access data set, available as a.csv file, and we introduce an associated web-based app for data exploration. MASTREE+ will provide the basis for improved understanding of the response of long-lived plant reproduction to environmental change. Additionally, MASTREE+ will enable investigation of the ecology and evolution of reproductive strategies in perennial plants, and the role of plant reproduction as a driver of ecosystem dynamics. EEA Santa Cruz Fil: Hacket-Pain, Andrew. University of Liverpool. School of Environmental Sciences. Department of Geography and Planning; Reino Unido Fil: Foest, Jessie J. University of Liverpool. School of Environmental Sciences. Department of Geography and Planning; Reino Unido Fil: Pearse, Ian S. U.S. Fort Collins Science Center. Geological Survey; Estados Unidos Fil: LaMontagne, Jalene M. DePaul University. Department of Biological Sciences; Estados Unidos Fil: Koenig, Walter D. University of California Berkeley. Hastings Reservation; Estados Unidos Fil: Vacchiano, Giorgio. University of Milan. Milan Department of Agricultural and Environmental Sciences; Italia. Fil: Bogdziewicz, Michał. Adam Mickiewicz University. Faculty of Biology. Institute of Environmental Biology; Polonia Fil: Bogdziewicz, Michał. University Grenoble Alpes. INRAE, LESSEM; Francia Fil: Caignard, Thomas. Université Bordeaux. INRAE, BIOGECO; Francia Fil: Celebias, Paulina. Adam Mickiewicz University. Faculty of Biology. Institute of Environmental Biology; Polonia Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina. Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina. Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Ascoli, Davide. University of Torino. Department of Agricultural. Forest and Food Sciences (DISAFA); Italia Fil: Gallo, Leonardo Ariel. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Bariloche; Argentina Fil: Gallo, Leonardo Ariel. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones Forestales y Agropecuarias Bariloche; Argentina Fil: Marchelli, Paula. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Bariloche; Argentina Fil: Marchelli, Paula. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Instituto de Investigaciones Forestales y Agropecuarias Bariloche; Argentina

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates., Nature Communications, 12, ISSN:2041-1723

Our overall objective is to synthesize mast-seeding data on North American Pinaceae to detect characteristic features of reproduction (i.e. development cycle length, serotiny, dispersal agents), and test for patterns in temporal variation based on weather variables. We use a large dataset ( n = 286 time series; mean length = 18.9 years) on crop sizes in four conifer genera ( Abies , Picea , Pinus , Tsuga ) collected between 1960 and 2014. Temporal variability in mast seeding (CVp) for 2 year genera ( Abies , Picea , Tsuga ) was higher than for Pinus (3 year), and serotinous species had lower CVp than non-serotinous species; there were no relationships of CVp with elevation or latitude. There was no difference in family-wide CVp across four tree regions of North America. Across all genera, July temperature differences between bud initiation and the prior year (Δ T ) was more strongly associated with reproduction than absolute temperature. Both CVp and Δ T remained steady over time, while absolute temperature increased by 0.09°C per decade. Our use of the Δ T model included a modification for Pinus , which initiates cone primordia 2 years before seedfall, as opposed to 1 year. These findings have implications for how mast-seeding patterns may change with future increases in temperature, and the adaptive benefits of mast seeding. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’.

International audience; Despite its importance for forest regeneration, food webs, and human economies, changes in tree fecundity with tree size and age remain largely unknown. The allometric increase with tree diameter assumed in ecological models would substantially overestimate seed contributions from large trees if fecundity eventually declines with size. Current estimates are dominated by overrepresentation of small trees in regression models. We combined global fecundity data, including a substantial representation of large trees. We compared size–fecundity relationships against traditional allometric scaling with diameter and two models based on crown architecture. All allometric models fail to describe the declining rate of increase in fecundity with diameter found for 80% of 597 species in our analysis. The strong evidence of declining fecundity, beyond what can be explained by crown architectural change, is consistent with physiological decline. A downward revision of projected fecundity of large trees can improve the next generation of forest dynamic models.

Kelly's commentary regarding our paper1, claiming that nutrient scarcity cannot select for masting behaviour in plants, initiated a fruitful discussion on traditionally settled hypotheses about the evolution of reproductive behaviour in plants. In his commentary, Prof. Kelly raises support for a contrasting hypothesis explaining our observation that temporally variable seed production is more pronounced under nutrient scarcity, namely that nutrient scarcity does not directly cause seed production variability, but instead increases variability induced by economies of scale. The commentary hinges mainly on the argument that an economy of scale (EOS) is necessary to select for highly variable seed production. It also points out that there are no mechanisms by which nutrient scarcity would select for that particular trait over generations. In reply to the stimulating comment, we 1) propose a mechanism by which nutrient scarcity may select for highly variable seed production, with larger-scale weather patterns inducing masting synchrony across populations; 2) we further discuss why wind-pollination and predator satiation, the EOS suggested by Kelly, cannot be the only selective pressures that select for highly variable reproduction.

Seed distribution and deposition patterns around parent trees are strongly affected by functional traits and therefore influence the development of plant communities. To assess the limitations of seed dispersal and the extent to which diaspore and neighbouring parental traits explain seed rain, we used a 9-year seed data set based on 150 seed traps in a 25-ha area of a temperate forest in the Changbai Mountain. Among 480,598 seeds belonging to 12 families, 17 genera, and 26 species were identified, only 54% of the species with mature trees in the community were represented in seeds collected over the 9 years, indicating a limitation in seed dispersal. Understory species were most limited; overstory species were least limited. Species with wind-dispersed seed had the least limitation, while the lowest similarity in species richness was for animal-dispersed species followed by gravity-dispersed species; fleshy-fruited species had stronger dispersal limitations than dry-fruited species. Generalized linear mixed models showed that relative basal area had a significant positive effect on seed abundance in traps, while the contribution of diaspore traits was low for nearly all groups. These results suggest that tree traits had the strongest contribution to seed dispersal and deposition for all functional groups examined here. These findings strengthen the knowledge that tree traits are key in explaining seed deposition patterns, at least at the primary dispersal stage. This improved knowledge of sources of seeds that are dispersed could facilitate greater understanding of seedling and community dynamics in temperate forests.

Mast seeding is one of the most intriguing reproductive traits in nature. Despite its potential drawbacks in terms of fitness, the widespread existence of this phenomenon suggests that it should have evolutionary advantages under certain circumstances. Using a global dataset of seed production time series for 219 plant species from all of the continents, we tested whether masting behaviour appears predominantly in species with low foliar nitrogen and phosphorus concentrations when controlling for local climate and productivity. Here, we show that masting intensity is higher in species with low foliar N and P concentrations, and especially in those with imbalanced N/P ratios, and that the evolutionary history of masting behaviour has been linked to that of nutrient economy. Our results support the hypothesis that masting is stronger in species growing under limiting conditions and suggest that this reproductive behaviour might have evolved as an adaptation to nutrient limitations and imbalances.

1.Synchronous pulses of seed masting and natural disturbance have positive feedbacks on the reproduction of masting species in disturbance‐prone ecosystems. We test the hypotheses that disturbances and proximate causes of masting are correlated, and that their large‐scale synchrony is driven by similar climate teleconnection patterns at both inter‐annual and decadal time scales. 2.Hypotheses were tested on white spruce (Picea glauca), a masting species which surprisingly persists in fire‐prone boreal forests while lacking clear fire adaptations. We built masting, drought and fire indices at regional (Alaska, Yukon, Alberta, Quebec) and subcontinental scales (western North America) spanning the second half of the 20th century. Superposed Epoch Analysis tested the temporal associations between masting events, drought and burnt area at the regional scale. At the sub‐continental scale, Superposed Epoch Analysis tested whether El Nino‐Southern Oscillation (ENSO) and its coupled effects with the Atlantic Multidecadal Oscillation (AMO) in the positive phase (AMO+/ENSO+) synchronize drought, burnt area and masting. We additionally tested the consistency of our synchronization hypotheses on a decadal temporal scale to verify whether long‐term oscillations in AMO+/ENSO + are coherent to decadal variation in drought, burnt area and masting. 3.Analyses demonstrated synchronicity between drought, fire and masting. In all regions the year before a mast event was drier and more fire‐prone than usual. During AMO+/ENSO + events sub‐continental indices of drought and burnt area experienced significant departures from mean values. The same was observed for large‐scale masting in the subsequent year, confirming 1 year lag between fire and masting. Sub‐continental indices of burnt area and masting showed in‐phase decadal fluctuations led by the AMO+/ENSO+. Results support the “Environmental prediction hypothesis” for mast seeding. 4.Synthesis. We provide evidence of large‐scale synchronicity between seed masting in Picea glauca and fire regimes in boreal forests of western North America at both inter‐annual and decadal time scales. We conclude that seed production in white spruce predicts changes in disturbance regimes by sharing the same large‐scale climate drivers with drought and fire. This gives new insides in a mechanism providing a fire‐sensitive species with higher than expected adaptability to changes in climate.

Interannual variability of seed crops (CVp) has profound consequences for plant populations and food webs, where high CVp is termed 'masting'. Here we ask: is global variation in CVp better predicted by plant or habitat differences consistent with adaptive economies of scale, in which flower and seed benefits increase disproportionately during mast years; or by passive mechanisms, in which seed production responds to variation in resource availability associated with climate variability? To address this question, we compiled a dataset for phylogenetic comparative analysis of long-term fruit/seed production for plants comprising 920 time series spanning 311 plant species. Factors associated with both adaptive benefits of CVp (wind pollination and seed dispersal) and climatic variability (variability of summer precipitation) were among the best predictors of global variation in CVp. We observed a hump-shaped relationship between CVp and latitude and intermediate phylogenetic and geographic signals in CVp. CVp is patterned nonrandomly across the globe and over the plant tree of life, where high CVp is associated with species benefiting from economies of scale of seed or flower production and with species that experience variable rainfall over summer months when seeds usually mature.

Overfishing of top predators along the western Atlantic coastline has led to a trophic cascade in salt marshes, with increases in herbivorous purple marsh crab (Sesarma reticulatum (Say, 1817)) abundances in North American estuaries leading to overgrazing of cordgrass (Spartina alterniflora Loisel.) and shoreline erosion. To evaluate potential physiological limits on the range of S. reticulatum within an estuary, we evaluated survival and physiological tolerance of S. reticulatum from the Ashepoo–Combhee–Edisto (ACE) River Basin in South Carolina, USA, to combinations of salinity (5‰ and 30‰) and pH (pH 6.6, 7.6, and 8.6) challenges, representative of estuarine extremes. Survival, haemolymph ion concentrations, and gill Na+,K+-ATPase (NKA) and vacuolar-type H+-ATPase (VHA) activity were measured after a 48 h exposure to each experimental condition. Survival was nearly 100% and osmoregulatory control was maintained across estuarine salinity and pH ranges. Sesarma reticulatum appeared to be robust to all potential combinations of salinity and pH stressors examined in this study, and therefore are likely unrestricted in their fundamental niche based on these stressors throughout an estuary.

Questions Mast seeding, synchronous highly variable seed production among years, occurs in many perennial plant species and across diverse plant communities. Two predominant ultimate hypotheses for mast seeding are pollination efficiency and predator satiation, with weather conditions as a proximate cause. Little consensus has been achieved regarding the relative impacts of ultimate selection and proximate weather on variation in seed production. Moreover, mast-seeding studies often focus on a single species or phylogenetically closely related species, while studies on plant communities consisting of tree species with a diversity of reproductive strategies are less common. Location The 25-ha Changbaishan temperate forest dynamic plot, Northeast China. Methods We used eight-years of seed rain data to characterize the patterns and evaluate underlying selective drivers of mast seeding. We employed generalized linear mixed-effects models to analyze the relationships between interannual variability of mast seeding and weather conditions. Results All 20 species in this forest community exhibited high variability in annual seed production, but the magnitude of seed production among species was generally asynchronous across years. Wind-pollinated species had higher interannual variation of seed production than animal-pollinated species, while species dispersed by seed predators and abiotic modes (e.g., wind and gravity) showed little variation. Species responded individually to weather conditions for both temperature and precipitation, and spring phenology at the same year as seed production had a larger effect among species than both the weather conditions of the same summer and season-long lags. Conclusions Our findings suggest that pollination efficiency hypothesis hada much stronger effect than predation satiation hypothesis on mast seeding, and weather conditions showed the proximate role of weather drivers in producing the community-wide mast seeding pattern. We emphasize the necessity to simultaneously assess drivers of mast seeding for multiple species within a plant community. This article is protected by copyright. All rights reserved.

The resource budget model for mast seeding hypothesizes that soil nutrients proximately influence reproduction. Plants in high soil nutrient (particularly N) areas are predicted to have lower reproductive variability over time and higher mean reproduction. While often examined theoretically, there are relatively few empirical tests of this hypothesis. We quantified cone production of 110 individual white spruce (Picea glauca) trees over seven years and quantified plant-available soil macronutrients (N, Ca, K, Mg, P, S) in natural forest conditions across three years with different cone crop conditions. Each of these plant-available soil nutrients were correlated across years (rs = 0.55–0.89; all > 0.81 for total-N); spatially, total-N availability varied 366-fold across trees. Plant-available soil nutrients did not influence variability or mean annual reproduction, contrary to nutrient perturbation experiments. We examined within-year nutrient and cone-production relationships, and observed significant positive relationships between reproduction and plant-available soil nutrients only in a low-reproduction year preceding a mast event. Both during a mast event and the following year, when overall cone production was very high or very low, there were no relationships. Both external drivers (e.g., weather) and internal resource budgets likely influence soil nutrient-reproduction relationships. These results suggest that plant-available soil nutrients may not be a large factor influencing mast-seeding patterns among individuals in this species.

Resource pulses are rare events with a short duration and high magnitude that drive the dynamics of both plant and animal populations and communities

A Correction to this paper has been published: https://doi.org/10.1038/s41467-021-22025-2

European starlings (Sturnus vulgaris) in North America often achieve high densities in urban regions and compete with other cavity-nesting species for nest sites, including the rapidly declining red-headed woodpecker (Melanerpes erythrocephalus). However, at a continental scale, European starling populations are unrelated to trends in red-headed woodpeckers. To assess whether urbanization influences population trends of cavity-nesting bird species and the relationship between European starlings and other cavity-nesting birds, we compared population patterns within the highly urbanized region of Cook County, Illinois, USA (which encompasses the city of Chicago) with data from surrounding rural counties. We used data from the Breeding Bird Survey (BBS) and the Christmas Bird Count (CBC) spanning 1970–2017, and i) quantified population trends of 19 cavity-nesting species, and ii) examined the patterns of relationships of each of these species with European starlings. Models fit to population trends over time with a variable for urban/rural location included were selected over models without urban/rural location for 14/17 species in the BBS and 14/15 species in the CBC. Starlings showed population declines in both urban and rural BBS and in urban CBC, while they increased in the rural CBC. Red-headed woodpecker populations declined across every survey set and location. Contrary to predictions, the counts of some species in surveys were significantly positively correlated with European starling population trends, while none were significantly negatively correlated. Our findings suggest that urban and rural locations add information to population trends of cavity-nesting species, and that European starlings have little negative impact on cavity-nesting birds in either rural or urban locations, suggesting that habitat loss and other factors are likely greater impacts on red-headed woodpeckers and other members of the cavity-nesting guild.

Forest habitat is important for a variety of woodpecker species, and is under pressure from urbanization. Red-headed Woodpeckers (Melanerpes erythrocephalus; RHWO) were once abundant across Eastern North America, and their populations have been declining since the 1960s. Their distribution encompasses urban centers, and since urban habitats differ from natural forest areas, our goal was to understand RHWO nest-site selection in an urban context. We addressed two main questions 1) what are the characteristics of RHWO nest selection across multiple spatial scales and 2) how do RHWO nest tree characteristics in city parks compare to those in forest preserves? This work was done in Cook County, IL, which includes Chicago, the third-largest city in the USA by population. We examined 34 RHWO nest trees used between 2010 and 2013, their surrounding habitat, and the landscape within a 1 km radius. Used trees and habitats were compared to paired unused trees and habitats, and landscape-scale characteristics were compared to random locations. Advanced decay of the nest tree, low canopy cover and increased presence of fungus on trees in the surrounding habitat were the best predictors of RHWO nesting in the area. Nests were most commonly found in forested areas outside of dense urban areas. However, we did not detect significant differences in the characteristics of the nest trees located in forest preserves and city parks. Our findings are consistent with nest selection studies in rural and natural areas, suggesting that forest habitats in metropolitan landscapes can support RHWO nesting.

Mast seeding, or masting, is the highly variable and spatially synchronous production of seeds by a population of plants. The production of variable seed crops is typically correlated with weather, so it is of considerable interest whether global climate change has altered the variability of masting or the size of masting events. We compiled 1086 datasets of plant seed production spanning 1900–2014 and from around the world, and then analysed whether the coefficient of variation (CV) in seed set, a measure of masting, increased over time. Over this 115-year period, seed set became more variable for plants as a whole and for the particularly well-studied taxa of conifers and oaks. The increase in CV corresponded with a decrease in the long-term mean of seed set of plant species. Seed set CV increased to a greater degree in plant taxa with a tendency towards masting. Seed set is becoming more variable among years, especially for plant taxa whose masting events are known to affect animal populations. Such subtle change in reproduction can have wide-ranging effects on ecosystems because seed crops provide critical resources for a wide range of taxa and have cascading effects throughout food webs.

Mast seeding, or masting, is the highly variable and spatially synchronous production of seeds by a population of plants. The production of variable seed crops is typically correlated with weather, so it is of considerable interest whether global climate change has altered the variability of masting or the size of masting events. We compiled 1086 datasets of plant seed production spanning 1900–2014 and from around the world, and then analysed whether the coefficient of variation (CV) in seed set, a measure of masting, increased over time. Over this 115-year period, seed set became more variable for plants as a whole and for the particularly well-studied taxa of conifers and oaks. The increase in CV corresponded with a decrease in the long-term mean of seed set of plant species. Seed set CV increased to a greater degree in plant taxa with a tendency towards masting. Seed set is becoming more variable among years, especially for plant taxa whose masting events are known to affect animal populations. Such subtle change in reproduction can have wide-ranging effects on ecosystems because seed crops provide critical resources for a wide range of taxa and have cascading effects throughout food webs.

Tree cavities are used by a wide variety of species for nesting, food storage, and cover. Most studies on cavity availability have been conducted in forests, and little is known about urban areas. With urbanization, species that excavate cavities may be less abundant, natural tree-decay processes are managed, and tree densities are reduced, all of which may influence tree-cavity availability. We investigated three questions: 1) What is the prevalence of tree cavities in different habitats in the Chicago area? 2) How do the characteristics of natural and woodpecker-excavated cavities and cavity-trees differ across habitats? 3) How does the urban landscape influence the prevalence of tree cavities? We tested the capacity for large urban parks and residential areas to provide tree cavities at levels similar to forested areas. We surveyed 1,545 trees in these three habitats for excavated and natural (caused by decay) cavities. Cavities were most available in forests, where the density of trees was highest. We found that a similar proportion of trees in forests and parks had excavated cavities, but excavated cavities were rare in residential areas. Trees containing cavities were larger than control trees and had more decay, and excavated cavities were in larger trees with more decay than natural cavity trees. Canopy cover was the main landscape variable influencing excavated cavity availability. Our results suggest that the prevalence of tree cavities may not be a limiting factor for urban wildlife, however that is contingent on the levels of use of natural cavities, which is currently unknown.

In territorial species, competitive asymmetries can result in the uneven distribution of food resources as high-quality individuals force young or subordinates to occupy smaller or lower-quality sites, or both. However, spatiotemporal variation in the production of resources also can influence an individual's ability to monopolize resources and, consequently, affect survival. We examined how spatial and temporal variation in food supply affects the distribution of resources among territorial food-hoarding red squirrels (Tamiasciurus hudsonicus) by mapping cone production over 62.5 ha of white spruce (Picea glauca) forest and measuring the interrelationships between territory size, cone supply, and survival during 4 years of low cone abundance in Yukon, Canada. Territory size and cone production within a territory varied 10- and 520-fold, respectively, with juvenile squirrels occupying smaller territories with fewer cones. Because of small-scale heterogeneity in the distribution of cones, territor...

Although there is a quantitative method that is commonly used for identifying mast-seeding behaviour of a plant population based on the coefficient of variation (i.e. CV is standard deviation/mean>1), there is no general quantitative method for delineating “mast” as opposed to “non-mast” years. Mast years are, however, described qualitatively as years when “large”, “unusually large” and “high” seed production occurs. The use of a consistent and generally applicable method for delineating mast years across species and plant populations is important for synthesizing knowledge of the causes and consequences of mast seeding, which could be confounded by using different methods among studies. We examine six quantitative methods for identifying mast years: four methods from the literature and two methods developed here. We use 36 seed production datasets covering a variety of species with ≥10 years of data to test the performance of these six methods. For each method, we quantify the percentage of the datasets to which the method could be successfully applied, the magnitude of the mast year relative to the mean, the frequency of mast years and the occurrence of consecutive mast years. The majority of the methods failed to meet the criteria for a suitable method. The best method used the number of standard deviates (standardized deviate method) of the annual mean seed production from the long-term mean of the dataset to identify mast-seeding years. General results from the standardized deviate method include that the occurrence of mast-seeding years is largely unrelated to plant population CV, but similar across species and data collection methods.

Summary 1 Mast seeding is the synchronous and highly variable production of seed by a population of plants. Mast seeding results from the behaviour of individuals; however, little is known about the synchrony of individuals at local scales. 2 We address two primary questions at a within-population (17–36 ha study plots) and individual level: (i) How variable is seed production between and within years? (ii) How synchronized is seed production between individuals? 3 We monitored annual cone production of 356 Picea glauca (white spruce) from 1990 to 2005 within four plots spanning a total distance of 5.3 km in the Yukon Territory, Canada. 4 Spearman correlations (rs) were conducted to test for synchrony. Overall, the trees were moderately synchronous (mean rs (± SE) of 0.52 ± 0.14), and synchrony was statistically detectable (rs > 0) over all distances. Individuals 3 km apart. There was considerable variation in cone production patterns among pairs of individuals. 5 The number of mast years per plot varied from one to three. During a mast year, many individuals within plots produced large cone crops, with more variability between individuals in low mean cone years. Individual trees had dominant endogenous cycles varying from none to 1–5 years. Forty-four per cent of trees had no significant lag, 23% a negative 1-year lag, and 20% a positive 3-year lag. Basal area did not influence lags, but trees with higher mean cone production throughout the study were more likely to have a 3-year lag compared with no lag. 6 The scale of highest synchrony coincided with the scale at which the dominant seed predator in the area, the territorial red squirrel (Tamiasciurus hudsonicus), operates. This may be the scale at which selection for synchrony occurs. 7 Based on high synchrony locally, high synchrony within a mast year, and multiple lags in cone production by individuals, both available resources and strong weather cues appear to play roles in the observed patterns.

We report the novel occurrence of North American red squirrels (Tamiasciurus hudsonicus) feeding on spruce bark beetle (Dendroctonus rufipennis) larvae, and consider the energetic and ecological implications. Although a bark beetle outbreak was 1st noted at our study site in 1994, significant feeding on them by red squirrels was not observed until 2002, after which there was significant increase in the prevalence of autumn beetle foraging by squirrels into 2003. This increase corresponded with a decrease in spruce seed availability, the squirrels' preferred food source. Spruce bark beetles currently represent an important food source, with 73% of monitored squirrels engaged in beetle feeding in autumn 2003, providing 20% of daily energy requirements. In autumn 2002 and 2003, the density of beetle-infested trees on a squirrel's territory was a significant predictor of whether it fed on beetle larvae but not the proportion of its foraging bouts involving bark beetles. Feeding on larval spruce bark beetles by red squirrels represents a short-term solution to a climate-mediated beetle outbreak that will ultimately reduce local spruce seed production and habitat suitability for red squirrels.

The number of cones produced by coniferous trees is commonly estimated by visual counts from the ground of a portion of the tree multiplied by a simple conversion factor. Linear conversion factors have been used to estimate total cone production by white spruce (Picea glauca (Moench) Voss). However, these conversion factors originate from other coniferous species and were often based on assumptions of cone visibility within the crown and not on empirical data. We propose a simple method for estimating the total number of cones produced by individual white spruce. We counted visible cones (an index of cone production, or cone index) on a total of 60 trees located in Alberta and Yukon, Canada, that were then felled and all cones were counted. We found that log(actual total cones) = 0.073 + 1.189 × log(cone index) is more accurate for estimating total cone numbers for white spruce than are other conversion factors (ranging from total cones = 1.5 × cone index to total cones = 3.35 × cone index), as determined using Akaike's information criterion with small sample bias adjustment and a validation data set. The relationship between the index of cone production and actual total cones produced is nonlinear, which is contrary to that proposed for various Pinus species.