MASTREE+: Time-series of plant reproductive effort from six continents.

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.

Aún existen importantes vacíos en la comprensión de la respuesta reproductiva de las plantas al cambio medioambiental, en parte, porque su monitoreo en especies de plantas longevas requiere una observación directa durante muchos años, y estos conjuntos de datos rara vez han estado disponibles. Aquí presentamos a MASTREE +, una base de datos que recopila series de tiempo de la reproducción de las plantas de todo el planeta, poniendo a disposición estos datos de libre acceso para la comunidad científica. MASTREE + incluye 73.828 puntos de observación de la reproducción anual georreferenciados (ej. conteos de semillas y frutos) en poblaciones de plantas perennes en todo el mundo. Estas observaciones consisten en 5971 series temporales a nivel de población provenientes de 974 especies en 66 países. La mediana de la duración de las series de tiempo es de 10 años (media = 12.4 años) y el conjunto de datos incluye 1.122 series de al menos dos décadas (≥20 años de observaciones). Para un subconjunto de especies bien estudiadas, MASTREE +incluye un amplio conjunto de series temporales replicadas en gradientes geográficos y climáticos. Describimos el conjunto de datos de acceso abierto disponible como un archivo.csv y presentamos una aplicación web asociada para la exploración de datos. MASTREE+ proporcionará la base para mejorar la comprensión sobre la respuesta reproductiva de plantas longevas al cambio medioambiental. Además, MASTREE+ facilitará los avances en la investigación de la ecología y la evolución de las estrategias reproductivas en plantas perennes y el papel de la reproducción vegetal como determinante de la dinámica de ecosistemas.

From theory to experiments for testing the proximate mechanisms of mast seeding: an agenda for an experimental ecology.

Highly variable and synchronised production of seeds by plant populations, known as masting, is implicated in many important ecological processes, but how it arises remains poorly understood. The lack of experimental studies prevents underlying mechanisms from being explicitly tested, and thereby precludes meaningful predictions on the consequences of changing environments for plant reproductive patterns and global vegetation dynamics. Here we review the most relevant proximate drivers of masting and outline a research agenda that takes the biology of masting from a largely observational field of ecology to one rooted in mechanistic understanding. We divide the experimental framework into three main processes: resource dynamics, pollen limitation and genetic and hormonal regulation, and illustrate how specific predictions about proximate mechanisms can be tested, highlighting the few successful experiments as examples. We envision that the experiments we outline will deliver new insights into how and why masting patterns might respond to a changing environment.

Geographical adaptation prevails over species-specific determinism in trees' vulnerability to climate change at Mediterranean rear-edge forests.

Climate change may reduce forest growth and increase forest mortality, which is connected to high carbon costs through reductions in gross primary production and net ecosystem exchange. Yet, the spatiotemporal patterns of vulnerability to both short-term extreme events and gradual environmental changes are quite uncertain across the species' limits of tolerance to dryness. Such information is fundamental for defining ecologically relevant upper limits of species tolerance to drought and, hence, to predict the risk of increased forest mortality and shifts in species composition. We investigate here to what extent the impact of short- and long-term environmental changes determines vulnerability to climate change of three evergreen conifers (Scots pine, silver fir, Norway spruce) and two deciduous hardwoods (European beech, sessile oak) tree species at their southernmost limits of distribution in the Mediterranean Basin. Finally, we simulated future forest growth under RCP 2.6 and 8.5 emission scenarios using a multispecies generalized linear mixed model. Our analysis provides four key insights into the patterns of species' vulnerability to climate change. First, site climatic marginality was significantly linked to the growth trends: increasing growth was related to less climatically limited sites. Second, estimated species-specific vulnerability did not match their a priori rank in drought tolerance: Scots pine and beech seem to be the most vulnerable species among those studied despite their contrasting physiologies. Third, adaptation to site conditions prevails over species-specific determinism in forest response to climate change. And fourth, regional differences in forests vulnerability to climate change across the Mediterranean Basin are linked to the influence of summer atmospheric circulation patterns, which are not correctly represented in global climate models. Thus, projections of forest performance should reconsider the traditional classification of tree species in functional types and critically evaluate the fine-scale limitations of the climate data generated by global climate models.

Climatically controlled reproduction drives interannual growth variability in a temperate tree species.

Climatically controlled allocation to reproduction is a key mechanism by which climate influences tree growth and may explain lagged correlations between climate and growth. We used continent-wide datasets of tree-ring chronologies and annual reproductive effort in Fagus sylvatica from 1901 to 2015 to characterise relationships between climate, reproduction and growth. Results highlight that variable allocation to reproduction is a key factor for growth in this species, and that high reproductive effort ('mast years') is associated with stem growth reduction. Additionally, high reproductive effort is associated with previous summer temperature, creating lagged climate effects on growth. Consequently, understanding growth variability in forest ecosystems requires the incorporation of reproduction, which can be highly variable. Our results suggest that future response of growth dynamics to climate change in this species will be strongly influenced by the response of reproduction.

Spatial patterns and broad-scale weather cues of beech mast seeding in Europe.

Mast seeding is a crucial population process in many tree species, but its spatio-temporal patterns and drivers at the continental scale remain unknown . Using a large dataset (8000 masting observations across Europe for years 1950-2014) we analysed the spatial pattern of masting across the entire geographical range of European beech, how it is influenced by precipitation, temperature and drought, and the temporal and spatial stability of masting-weather correlations. Beech masting exhibited a general distance-dependent synchronicity and a pattern structured in three broad geographical groups consistent with continental climate regimes. Spearman's correlations and logistic regression revealed a general pattern of beech masting correlating negatively with temperature in the summer 2 yr before masting, and positively with summer temperature 1 yr before masting (i.e. 2T model). The temperature difference between the two previous summers (DeltaT model) was also a good predictor. Moving correlation analysis applied to the longest eight chronologies (74-114 yr) revealed stable correlations between temperature and masting, confirming consistency in weather cues across space and time. These results confirm widespread dependency of masting on temperature and lend robustness to the attempts to reconstruct and predict mast years using temperature data.

Two centuries of masting data for European beech and Norway spruce across the European continent.

Tree masting is one of the most intensively studied ecological processes. It affects nutrient fluxes of trees, regeneration dynamics in forests, animal population densities, and ultimately influences ecosystem services. Despite a large volume of research focused on masting, its evolutionary ecology, spatial and temporal variability, and environmental drivers are still matter of debate. Understanding the proximate and ultimate causes of masting at broad spatial and temporal scales will enable us to predict tree reproductive strategies and their response to changing environment. Here we provide broad spatial (distribution range-wide) and temporal (century) masting data for the two main masting tree species in Europe, European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) H. Karst.). We collected masting data from a total of 359 sources through an extensive literature review and from unpublished surveys. The data set has a total of 1,747 series and 18,348 yearly observations from 28 countries and covering a time span of years 1677-2016 and 1791-2016 for beech and spruce, respectively. For each record, the following information is available: identification code; species; year of observation; proxy of masting (flower, pollen, fruit, seed, dendrochronological reconstructions); statistical data type (ordinal, continuous); data value; unit of measurement (only in case of continuous data); geographical location (country, Nomenclature of Units for Territorial Statistics NUTS-1 level, municipality, coordinates); first and last record year and related length; type of data source (field survey, peer reviewed scientific literature, gray literature, personal observation); source identification code; date when data were added to the database; comments. To provide a ready-to-use masting index we harmonized ordinal data into five classes. Furthermore, we computed an additional field where continuous series with length >4 yr where converted into a five classes ordinal index. To our knowledge, this is the most comprehensive published database on species-specific masting behavior. It is useful to study spatial and temporal patterns of masting and its proximate and ultimate causes, to refine studies based on tree-ring chronologies, to understand dynamics of animal species and pests vectored by these animals affecting human health, and it may serve as calibration-validation data for dynamic forest models.

A walk on the wild side: Disturbance dynamics and the conservation and management of European mountain forest ecosystems.

Mountain forests are among the most important ecosystems in Europe as they support numerous ecological, hydrological, climatic, social, and economic functions. They are unique relatively natural ecosystems consisting of long-lived species in an otherwise densely populated human landscape. Despite this, centuries of intensive forest management in many of these forests have eclipsed evidence of natural processes, especially the role of disturbances in long-term forest dynamics. Recent trends of land abandonment and establishment of protected forests have coincided with a growing interest in managing forests in more natural states. At the same time, the importance of past disturbances highlighted in an emerging body of literature, and recent increasing disturbances due to climate change are challenging long-held views of dynamics in these ecosystems. Here, we synthesize aspects of this Special Issue on the ecology of mountain forest ecosystems in Europe in the context of broader discussions in the field, to present a new perspective on these ecosystems and their natural disturbance regimes. Most mountain forests in Europe, for which long-term data are available, show a strong and long-term effect of not only human land use but also of natural disturbances that vary by orders of magnitude in size and frequency. Although these disturbances may kill many trees, the forests themselves have not been threatened. The relative importance of natural disturbances, land use, and climate change for ecosystem dynamics varies across space and time. Across the continent, changing climate and land use are altering forest cover, forest structure, tree demography, and natural disturbances, including fires, insect outbreaks, avalanches, and wind disturbances. Projected continued increases in forest area and biomass along with continued warming are likely to further promote forest disturbances. Episodic disturbances may foster ecosystem adaptation to the effects of ongoing and future climatic change. Increasing disturbances, along with trends of less intense land use, will promote further increases in coarse woody debris, with cascading positive effects on biodiversity, edaphic conditions, biogeochemical cycles, and increased heterogeneity across a range of spatial scales. Together, this may translate to disturbance-mediated resilience of forest landscapes and increased biodiversity, as long as climate and disturbance regimes remain within the tolerance of relevant species. Understanding ecological variability, even imperfectly, is integral to anticipating vulnerabilities and promoting ecological resilience, especially under growing uncertainty. Allowing some forests to be shaped by natural processes may be congruent with multiple goals of forest management, even in densely settled and developed countries.

Active governance of agro-pastoral, forest and protected areas mitigates wildfire impacts in Italy.

Wildfire regimes affected by global change have been the cause of major concern in recent years. Both direct prevention (e.g., fuel management planning) and land governance strategies (e.g., agroforestry development) can have an indirect regulatory effect on wildfires. Herein, we tested the hypothesis that active land planning and management in Italy have mitigated wildfire impacts in terms of loss of ecosystem services and forest cover, and burned wildland-urban interface, from 2007 to 2017. At the national scale, we assessed the effect size of major potential fire drivers such as climate, weather, flammability, socio-economic descriptors, land use changes, and proxies for land governance (e.g., European funds for rural development, investments in sustainable forest management, agro-pastoral activities), including potential interactions, on fire-related impacts via Random Forest modelling and Generalized Additive Mixed Model. Agro-forest districts (i.e., aggregations of neighbouring municipalities with homogeneous forest and agricultural characteristics) were used as spatial units of analysis. Our results confirm that territories with more active land governance show lower wildfire impacts, even under severe flammability and climatic conditions. This study supports current regional, national, and European strategies towards "fire resistant and resilient landscapes" by fostering agro-forestry, rural development, and nature conservation integrated policies.

The ecology and evolution of synchronized reproduction in long-lived plants.

Populations of many long-lived plants exhibit spatially synchronized seed production that varies extensively over time, so that seed production in some years is much higher than on average, while in others, it is much lower or absent. This phenomenon termed masting or mast seeding has important consequences for plant reproductive success, ecosystem dynamics and plant-human interactions. Inspired by recent advances in the field, this special issue presents a series of articles that advance the current understanding of the ecology and evolution of masting. To provide a broad overview, we reflect on the state-of-the-art of masting research in terms of underlying proximate mechanisms, ontogeny, adaptations, phylogeny and applications to conservation. While the mechanistic drivers and fitness consequences of masting have received most attention, the evolutionary history, ontogenetic trajectory and applications to plant-human interactions are poorly understood. With increased availability of long-term datasets across broader geographical and taxonomic scales, as well as advances in molecular approaches, we expect that many mysteries of masting will be solved soon. The increased understanding of this global phenomenon will provide the foundation for predictive modelling of seed crops, which will improve our ability to manage forests and agricultural fruit and nut crops in the Anthropocene. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'.

Modes of climate variability bridge proximate and evolutionary mechanisms of masting.

There is evidence that variable and synchronous reproduction in seed plants (masting) correlates to modes of climate variability, e.g. El Niño Southern Oscillation and North Atlantic Oscillation. In this perspective, we explore the breadth of knowledge on how climate modes control reproduction in major masting species throughout Earth's biomes. We posit that intrinsic properties of climate modes (periodicity, persistence and trends) drive interannual and decadal variability of plant reproduction, as well as the spatial extent of its synchrony, aligning multiple proximate causes of masting through space and time. Moreover, climate modes force lagged but in-phase ecological processes that interact synergistically with multiple stages of plant reproductive cycles. This sets up adaptive benefits by increasing offspring fitness through either economies of scale or environmental prediction. Community-wide links between climate modes and masting across plant taxa suggest an evolutionary role of climate variability. We argue that climate modes may 'bridge' proximate and ultimate causes of masting selecting for variable and synchronous reproduction. The future of such interaction is uncertain: processes that improve reproductive fitness may remain coupled with climate modes even under changing climates, but chances are that abrupt global warming will affect Earth's climate modes so rapidly as to alter ecological and evolutionary links. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'.

Natural disturbances and masting: from mechanisms to fitness consequences.

The timing of seed production and release is highly relevant for successful plant reproduction. Ecological disturbances, if synchronized with reproductive effort, can increase the chances of seeds and seedlings to germinate and establish. This can be especially true under variable and synchronous seed production (masting). Several observational studies have reported worldwide evidence for co-occurrence of disturbances and seed bumper crops in forests. Here, we review the evidence for interaction between disturbances and masting in global plant communities; we highlight feedbacks between these two ecological processes and posit an evolutionary pathway leading to the selection of traits that allow trees to synchronize seed crops with disturbances. Finally, we highlight relevant questions to be tested on the functional and evolutionary relationship between disturbances and masting. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'.

Tackling unresolved questions in forest ecology: The past and future role of simulation models.

Understanding the processes that shape forest functioning, structure, and diversity remains challenging, although data on forest systems are being collected at a rapid pace and across scales. Forest models have a long history in bridging data with ecological knowledge and can simulate forest dynamics over spatio-temporal scales unreachable by most empirical investigations.We describe the development that different forest modelling communities have followed to underpin the leverage that simulation models offer for advancing our understanding of forest ecosystems.Using three widely applied but contrasting approaches - species distribution models, individual-based forest models, and dynamic global vegetation models - as examples, we show how scientific and technical advances have led models to transgress their initial objectives and limitations. We provide an overview of recent model applications on current important ecological topics and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.Synthesis. This overview shows that forest models, due to their complementarity and mutual enrichment, represent an invaluable toolkit to address a wide range of fundamental and applied ecological questions, hence fostering a deeper understanding of forest dynamics in the context of global change.

Forest carbon allocation modelling under climate change.

Carbon allocation plays a key role in ecosystem dynamics and plant adaptation to changing environmental conditions. Hence, proper description of this process in vegetation models is crucial for the simulations of the impact of climate change on carbon cycling in forests. Here we review how carbon allocation modelling is currently implemented in 31 contrasting models to identify the main gaps compared with our theoretical and empirical understanding of carbon allocation. A hybrid approach based on combining several principles and/or types of carbon allocation modelling prevailed in the examined models, while physiologically more sophisticated approaches were used less often than empirical ones. The analysis revealed that, although the number of carbon allocation studies over the past 10 years has substantially increased, some background processes are still insufficiently understood and some issues in models are frequently poorly represented, oversimplified or even omitted. Hence, current challenges for carbon allocation modelling in forest ecosystems are (i) to overcome remaining limits in process understanding, particularly regarding the impact of disturbances on carbon allocation, accumulation and utilization of nonstructural carbohydrates, and carbon use by symbionts, and (ii) to implement existing knowledge of carbon allocation into defence, regeneration and improved resource uptake in order to better account for changing environmental conditions.

Tree mortality submodels drive simulated long-term forest dynamics: assessing 15 models from the stand to global scale.

Models are pivotal for assessing future forest dynamics under the impacts of changing climate and management practices, incorporating representations of tree growth, mortality, and regeneration. Quantitative studies on the importance of mortality submodels are scarce. We evaluated 15 dynamic vegetation models (DVMs) regarding their sensitivity to different formulations of tree mortality under different degrees of climate change. The set of models comprised eight DVMs at the stand scale, three at the landscape scale, and four typically applied at the continental to global scale. Some incorporate empirically derived mortality models, and others are based on experimental data, whereas still others are based on theoretical reasoning. Each DVM was run with at least two alternative mortality submodels. Model behavior was evaluated against empirical time series data, and then, the models were subjected to different scenarios of climate change. Most DVMs matched empirical data quite well, irrespective of the mortality submodel that was used. However, mortality submodels that performed in a very similar manner against past data often led to sharply different trajectories of forest dynamics under future climate change. Most DVMs featured high sensitivity to the mortality submodel, with deviations of basal area and stem numbers on the order of 10-40% per century under current climate and 20-170% under climate change. The sensitivity of a given DVM to scenarios of climate change, however, was typically lower by a factor of two to three. We conclude that (1) mortality is one of the most uncertain processes when it comes to assessing forest response to climate change, and (2) more data and a better process understanding of tree mortality are needed to improve the robustness of simulated future forest dynamics. Our study highlights that comparing several alternative mortality formulations in DVMs provides valuable insights into the effects of process uncertainties on simulated future forest dynamics.

Inter-annual and decadal changes in teleconnections drive continental-scale synchronization of tree reproduction.

Climate teleconnections drive highly variable and synchronous seed production (masting) over large scales. Disentangling the effect of high-frequency (inter-annual variation) from low-frequency (decadal trends) components of climate oscillations will improve our understanding of masting as an ecosystem process. Using century-long observations on masting (the MASTREE database) and data on the Northern Atlantic Oscillation (NAO), we show that in the last 60 years both high-frequency summer and spring NAO, and low-frequency winter NAO components are highly correlated to continent-wide masting in European beech and Norway spruce. Relationships are weaker (non-stationary) in the early twentieth century. This finding improves our understanding on how climate variation affects large-scale synchronization of tree masting. Moreover, it supports the connection between proximate and ultimate causes of masting: indeed, large-scale features of atmospheric circulation coherently drive cues and resources for masting, as well as its evolutionary drivers, such as pollination efficiency, abundance of seed dispersers, and natural disturbance regimes.

Forest disturbances under climate change.

Forest disturbances are sensitive to climate. However, our understanding of disturbance dynamics in response to climatic changes remains incomplete, particularly regarding large-scale patterns, interaction effects and dampening feedbacks. Here we provide a global synthesis of climate change effects on important abiotic (fire, drought, wind, snow and ice) and biotic (insects and pathogens) disturbance agents. Warmer and drier conditions particularly facilitate fire, drought and insect disturbances, while warmer and wetter conditions increase disturbances from wind and pathogens. Widespread interactions between agents are likely to amplify disturbances, while indirect climate effects such as vegetation changes can dampen long-term disturbance sensitivities to climate. Future changes in disturbance are likely to be most pronounced in coniferous forests and the boreal biome. We conclude that both ecosystems and society should be prepared for an increasingly disturbed future of forests.

Fire severity, residuals and soil legacies affect regeneration of Scots pine in the Southern Alps.

Regeneration of non fire-adapted conifers following crown fires on the European Alps is often delayed or unsuccessful. Fire may limit establishment by eliminating seed trees, altering soil properties, or modifying microsite and soil conditions via disturbance legacies. However, the effect of soil legacies on post-fire establishment has rarely been discussed. We analyzed the abundance of Scots pine regeneration in a 257 ha wildfire in an inner-alpine forest. Our aims were (1) to model fire intensity at the soil surface and topsoil heating along a gradient of increasing fire severities; (2) to assess the differences in soil properties along the fire severity gradient; (3) to model the effect of disturbance and soil legacies on the density of pine seedlings. We reconstructed fire behavior and soil heating with the First Order Fire Effects Model (FOFEM), tested the effect of fire severity on soils by nonparametric distributional tests, and modeled seedling density as a function of site, disturbance and soil legacies by fitting a GLM following a variable selection procedure. Topsoil heating differed markedly between the moderate and high severity fires, reaching temperatures high enough to strongly and permanently alter soil properties only in the latter. High fire severity resulted in decreased soil consistency and wet aggregate stability. Burned soils had lower organic matter and cations than those unburned. Pine seedlings favored low-fertility, eroded, and chemically poor sites. Establishment was facilitated by the presence of coarse woody debris, but hampered by increasing distance from the seed source. These results suggest that in dry, inner-alpine valleys, fire residuals and soil legacies interact in determining the success of Scots pine re-establishment. High severity fire can promote favorable soil conditions, but distance from the seed source and high evaporation rates of bare soils must be mitigated in order to ensure a successful restoration.

Driving factors of a vegetation shift from Scots pine to pubescent oak in dry Alpine forests.

An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner-Alpine regions, the species composition in low elevation forests is changing: The sub-boreal Scots pine (Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub-Mediterranean pubescent oak (Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger-scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory (NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed-effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small-diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services.