Integrated global assessment of the natural forest carbon potential.

Forests are a substantial terrestrial carbon sink, but anthropogenic changes in land use and climate have considerably reduced the scale of this system1. Remote-sensing estimates to quantify carbon losses from global forests2-5 are characterized by considerable uncertainty and we lack a comprehensive ground-sourced evaluation to benchmark these estimates. Here we combine several ground-sourced6 and satellite-derived approaches2,7,8 to evaluate the scale of the global forest carbon potential outside agricultural and urban lands. Despite regional variation, the predictions demonstrated remarkable consistency at a global scale, with only a 12% difference between the ground-sourced and satellite-derived estimates. At present, global forest carbon storage is markedly under the natural potential, with a total deficit of 226 Gt (model range = 151-363 Gt) in areas with low human footprint. Most (61%, 139 Gt C) of this potential is in areas with existing forests, in which ecosystem protection can allow forests to recover to maturity. The remaining 39% (87 Gt C) of potential lies in regions in which forests have been removed or fragmented. Although forests cannot be a substitute for emissions reductions, our results support the idea2,3,9 that the conservation, restoration and sustainable management of diverse forests offer valuable contributions to meeting global climate and biodiversity targets.

Poleward shifts in the maximum of spring phenological responsiveness of Ginkgo biloba to temperature in China.

Global warming is advancing the timing of spring leaf-out in temperate and boreal plants, affecting biological interactions and global biogeochemical cycles. However, spatial variation in spring phenological responsiveness to climate change within species remains poorly understood. Here, we investigated variation in the responsiveness of spring phenology to temperature (RSP; days to leaf-out at a given temperature) in 2754 Ginkgo biloba twigs of trees distributed across subtropical and temperate regions in China from 24°N to 44°N. We found a nonlinear effect of mean annual temperature on spatial variation in RSP, with the highest response rate at c. 12°C and lower response rates at warmer or colder temperatures due to declines in winter chilling accumulation. We then predicted the spatial maxima in RSP under current and future climate scenarios, and found that trees are currently most responsive in central China, which corresponds to the species' main distribution area. Under a high-emission scenario, we predict a 4-degree latitude shift in the responsiveness maximum toward higher latitudes over the rest of the century. The identification of the nonlinear responsiveness of spring phenology to climate gradients and the spatial shifts in phenological responsiveness expected under climate change represent new mechanistic insights that can inform models of spring phenology and ecosystem functioning.

Native diversity buffers against severity of non-native tree invasions.

Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species1,2. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies3,4. Here, leveraging global tree databases5-7, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions.

Effect of climate warming on the timing of autumn leaf senescence reverses after the summer solstice.

Climate change is shifting the growing seasons of plants, affecting species performance and biogeochemical cycles. Yet how the timing of autumn leaf senescence in Northern Hemisphere forests will change remains uncertain. Using satellite, ground, carbon flux, and experimental data, we show that early-season and late-season warming have opposite effects on leaf senescence, with a reversal occurring after the year's longest day (the summer solstice). Across 84% of the northern forest area, increased temperature and vegetation activity before the solstice led to an earlier senescence onset of, on average, 1.9 ± 0.1 days per °C, whereas warmer post-solstice temperatures extended senescence duration by 2.6 ± 0.1 days per °C. The current trajectories toward an earlier onset and slowed progression of senescence affect Northern Hemisphere-wide trends in growing-season length and forest productivity.

Acclimation of phenology relieves leaf longevity constraints in deciduous forests.

Leaf phenology is key for regulating total growing-season mass and energy fluxes. Long-term temporal trends towards earlier leaf unfolding are observed across Northern Hemisphere forests. Phenological dates also vary between years, whereby end-of-season (EOS) dates correlate positively with start-of-season (SOS) dates and negatively with growing-season total net CO2 assimilation (Anet). These associations have been interpreted as the effect of a constrained leaf longevity or of premature carbon (C) sink saturation-with far-reaching consequences for long-term phenology projections under climate change and rising CO2. Here, we use multidecadal ground and remote-sensing observations to show that the relationships between Anet and EOS are opposite at the interannual and the decadal time scales. A decadal trend towards later EOS persists in parallel with a trend towards increasing Anet-in spite of the negative Anet-EOS relationship at the interannual scale. This finding is robust against the use of diverse observations and models. Results indicate that acclimation of phenology has enabled plants to transcend a constrained leaf longevity or premature C sink saturation over the course of several decades, leading to a more effective use of available light and a sustained extension of the vegetation CO2 uptake season over time.

Global warming is increasing the discrepancy between green (actual) and thermal (potential) seasons of temperate trees.

Over the past decades, global warming has led to a lengthening of the time window during which temperatures remain favorable for carbon assimilation and tree growth, resulting in a lengthening of the green season. The extent to which forest green seasons have tracked the lengthening of this favorable period under climate warming, however, has not been quantified to date. Here, we used remote sensing data and long-term ground observations of leaf-out and coloration for six dominant species of European trees at 1773 sites, for a total of 6060 species-site combinations, during 1980-2016 and found that actual green season extensions (GS: 3.1 ± 0.1 day decade-1 ) lag four times behind extensions of the potential thermal season (TS: 12.6 ± 0.1 day decade-1 ). Similar but less pronounced differences were obtained using satellite-derived vegetation phenology observations, that is, a lengthening of 4.4 ± 0.13 and 7.5 ± 0.13 day decade-1 for GS and TS, respectively. This difference was mainly driven by the larger advance in the onset of the thermal season compared to the actual advance of leaf-out dates (spring mismatch: 7.2 ± 0.1 day decade-1 ), but to a less extent caused by a phenological mismatch between GS and TS in autumn (2.4 ± 0.1 day decade-1 ). Our results showed that forest trees do not linearly track the new thermal window extension, indicating more complex interactions between winter and spring temperatures and photoperiod and a justification of demonstrating that using more sophisticated models that include the influence of chilling and photoperiod is needed to accurately predict spring phenological changes under warmer climate. They urge caution if such mechanisms are omitted to predict, for example, how vegetative health and growth, species distribution and crop yields will change in the future.

Links across ecological scales: Plant biomass responses to elevated CO2.

The degree to which elevated CO2 concentrations (e[CO2 ]) increase the amount of carbon (C) assimilated by vegetation plays a key role in climate change. However, due to the short-term nature of CO2 enrichment experiments and the lack of reconciliation between different ecological scales, the effect of e[CO2 ] on plant biomass stocks remains a major uncertainty in future climate projections. Here, we review the effect of e[CO2 ] on plant biomass across multiple levels of ecological organization, scaling from physiological responses to changes in population-, community-, ecosystem-, and global-scale dynamics. We find that evidence for a sustained biomass response to e[CO2 ] varies across ecological scales, leading to diverging conclusions about the responses of individuals, populations, communities, and ecosystems. While the distinct focus of every scale reveals new mechanisms driving biomass accumulation under e[CO2 ], none of them provides a full picture of all relevant processes. For example, while physiological evidence suggests a possible long-term basis for increased biomass accumulation under e[CO2 ] through sustained photosynthetic stimulation, population-scale evidence indicates that a possible e[CO2 ]-induced increase in mortality rates might potentially outweigh the effect of increases in plant growth rates on biomass levels. Evidence at the global scale may indicate that e[CO2 ] has contributed to increased biomass cover over recent decades, but due to the difficulty to disentangle the effect of e[CO2 ] from a variety of climatic and land-use-related drivers of plant biomass stocks, it remains unclear whether nutrient limitations or other ecological mechanisms operating at finer scales will dampen the e[CO2 ] effect over time. By exploring these discrepancies, we identify key research gaps in our understanding of the effect of e[CO2 ] on plant biomass and highlight the need to integrate knowledge across scales of ecological organization so that large-scale modeling can represent the finer-scale mechanisms needed to constrain our understanding of future terrestrial C storage.

Direct and indirect impacts of urbanization on vegetation growth across the world's cities.

Urban environments, regarded as "harbingers" of future global change, may exert positive or negative impacts on urban vegetation growth. Because of limited ground-based experiments, the responses of vegetation to urbanization and its associated controlling factors at the global scale remain poorly understood. Here, we use satellite observations from 2001 to 2018 to quantify direct and indirect impacts of urbanization on vegetation growth in 672 worldwide cities. After controlling for the negative direct impact of urbanization on vegetation growth, we find a widespread positive indirect effect that has been increasing over time. These indirect effects depend on urban development intensity, population density, and background climate, with more pronounced positive effects in cities with cold and arid environments. We further show that vegetation responses to urbanization are modulated by a cities' developmental status. Our findings have important implications for understanding urbanization-induced impacts on vegetation and future sustainable urban development.

Global relationships in tree functional traits.

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide.

Carbon Source Reduction Postpones Autumn Leaf Senescence in a Widespread Deciduous Tree.

The growing-season length of temperate and boreal trees has a strong effect on the global carbon cycle. Yet, a poor understanding of the drivers of phenological processes, such as autumn leaf senescence in deciduous trees, limits our capacity to estimate growing-season lengths under climate change. While temperature has been shown to be an important driver of autumn leaf senescence, carbon source-sink dynamics have been proposed as a mechanism that could help explain variation of this important process. According to the carbon sink limitation hypothesis, senescence is regulated by the interplay between plant carbon source and sink dynamics, so that senescence occurs later upon low carbon inputs (source) and earlier upon low carbon demand (sink). Here, we manipulated carbon source-sink dynamics in birch saplings (Betula pendula) to test the relevance of carbon sink limitation for autumn leaf senescence and photosynthetic decline in a widespread deciduous tree. Specifically, we conducted a gradient of leaf and bud removal treatments and monitored the effects on autumnal declines in net photosynthesis and the timing of leaf senescence. In line with the carbon sink limitation hypothesis, we observed that leaf removal tended to increase total leaf-level autumn photosynthesis and delayed the timing of senescence. Conversely, we did not observe an effect of bud removal on either photosynthesis or senescence, which was likely caused by the fact that our bud removal treatment did not considerably affect the plant carbon sink. While we cannot fully rule out that the observed effect of leaf removal was influenced by possible treatment-level differences in leaf age or soil resource availability, our results provide support for the hypothesis of carbon sink limitation as a driver of growing-season length and move the scientific field closer to narrowing the uncertainty in climate change predictions.

Trees growing in Eastern North America experience higher autumn solar irradiation than their European relatives, but is nitrogen limitation another factor explaining anthocyanin-red autumn leaves?: A comment on Peña-Novas and Marchetti 2021 (https://doi.org/10.1111/jeb.13903).

Trees growing at a particular latitude in Eastern North America (ENA) receive more autumn solar irradiation than do trees growing at the same latitude in Europe, a difference that could partly explain the higher percentage of anthocyanin-producing deciduous species in ENA compared with European floras. A proposed link between autumn light intensity and the production of anthocyanins is their function in photoprotection, which enables plants with red leaves to resorb nutrients for a longer time than can yellow or brown leaves. The innately 4-week-shorter foliage period of ENA trees compared with European ones may also play a role, as may stronger nitrogen limitation in poorer soils in ENA. We here test for a correlation between nitrogen access and fall anthocyanin in 126 species (55 genera, 22 families) from different temperate forests, using a Hierarchical Bayesian model and accounting for phylogenetic structure in the data. None of 81 species with nitrogen-fixing symbionts produce autumn anthocyanin, whereas 42% of non-nitrogen-fixers do. Thus, when ample nitrogen is available from symbionts, the benefits of anthocyanin-derived photoprotection apparently do not outweigh the costs of anthocyanin production. If nitrogen limitation indeed plays a role in the dominance of red-autumn-colouring trees in ENA floras - while European floras predominantly produce yellow autumn leaves - there might also be continental differences in the frequency or abundance of nitrogen-fixing trees and shrubs, a new hypothesis that deserves testing.

Impact of microclimatic conditions and resource availability on spring and autumn phenology of temperate tree seedlings.

Microclimatic effects (light, temperature) are often neglected in phenological studies and little information is known about the impact of resource availability (nutrient and water) on tree's phenological cycles. Here we experimentally studied spring and autumn phenology in four temperate trees in response to changes in bud albedo (white-painted vs black-painted buds), light conditions (nonshaded vs c. 70% shaded), water availability (irrigated, control and reduced precipitation) and nutrients (low vs high availability). We found that higher bud albedo or shade delayed budburst (up to +12 d), indicating that temperature is sensed locally within each bud. Leaf senescence was delayed by high nutrient availability (up to +7 d) and shade conditions (up to +39 d) in all species, except oak. Autumn phenological responses to summer droughts depended on species, with a delay for cherry (+7 d) and an advance for beech (-7 d). The strong phenological effects of bud albedo and light exposure reveal an important role of microclimatic variation on phenology. In addition to the temperature and photoperiod effects, our results suggest a tight interplay between source and sink processes in regulating the end of the seasonal vegetation cycle, which can be largely influenced by resource availability (light, water and nutrients).

The global distribution and environmental drivers of aboveground versus belowground plant biomass.

A poor understanding of the fraction of global plant biomass occurring belowground as roots limits our understanding of present and future ecosystem function and carbon pools. Here we create a database of root-mass fractions (RMFs), an index of plant below- versus aboveground biomass distributions, and generate quantitative, spatially explicit global maps of RMFs in trees, shrubs and grasses. Our analyses reveal large gradients in RMFs both across and within vegetation types that can be attributed to resource availability. High RMFs occur in cold and dry ecosystems, while low RMFs dominate in warm and wet regions. Across all vegetation types, the directional effect of temperature on RMFs depends on water availability, suggesting feedbacks between heat, water and nutrient supply. By integrating our RMF maps with existing aboveground plant biomass information, we estimate that in forests, shrublands and grasslands, respectively, 22%, 47% and 67% of plant biomass exists belowground, with a total global belowground fraction of 24% (20-28%), that is, 113 (90-135) Gt carbon. By documenting the environmental correlates of root biomass allocation, our results can inform model projections of global vegetation dynamics under current and future climate scenarios.

Climate data and flowering times for 450 species from 1844 deepen the record of phenological change in southern Germany.

PREMISE: State-sponsored weather stations became ubiquitous by the 1880s, yet many old climate data and phenological observations still need to be digitized and made accessible. METHODS: We here make available flowering times for 450 species of herbs and shrubs gathered in 1844 by Carl Friedrich Philipp von Martius (1794-1868), director of the Munich Botanical Garden. The data formed part of the world's third-oldest phenological monitoring network as we explain in a brief overview of the history of such networks. Using data from one of the world's oldest continuously functioning weather stations, Hohenpeißenberg, we relate temperature to flowering in three species with short flowering times and herbarium collections made since 1844 within the city's perimeter, namely, Anemone patens, A. pulsatilla, and Arum maculatum. RESULTS: Mean advances in flowering dates were 1.3-2.1 days/decade or 3.2-4.2 days/1°C warming. These advances are in keeping with similar advances in other European herbs during more recent periods. CONCLUSIONS: Future studies might use the 1844 flowering data made available here as a source of information on the availability of particular flowers for specialized pollinators including insects looking for oviposition sites, such as the Psychoda flies that become trapped in Arum inflorescences. Another use of Martius's 1844 data would be their incorporation into larger-scale analyses of flowering in southern-central Europe.

Response to Comment on "Increased growing-season productivity drives earlier autumn leaf senescence in temperate trees".

Our study showed that increases in seasonal productivity drive earlier autumn senescence of temperate trees. Norby argues that this finding is contradicted by observations from free-air CO2 enrichment (FACE) experiments, where elevated CO2 has been found to delay senescence in some cases. We provide a detailed answer showing that the results from FACE studies are in agreement with our conclusions.

Chilled to be forced: the best dose to wake up buds from winter dormancy.

Over the last decades, spring leaf-out of temperate and boreal trees has substantially advanced in response to global warming, affecting terrestrial biogeochemical fluxes and the Earth's climate system. However, it remains unclear whether leaf-out will continue to advance with further warming because species' effective chilling temperatures, as well as the amount of chilling time required to break dormancy, are still largely unknown for most forest tree species. Here, we assessed the progress of winter dormancy and quantified the efficiency of different chilling temperatures in six dominant temperate European tree species by exposing 1170 twig cuttings to a range of temperatures from -2°C to 10°C for 1, 3, 6 or 12 wk. We found that freezing temperatures were most effective for half of the species or as effective as chilling temperatures up to 10°C, that is, leading to minimum thermal time to and maximum success of budburst. Interestingly, chilling duration had a much larger effect on dormancy release than absolute chilling temperature. Our experimental results challenge the common assumption that optimal chilling temperatures range c. 4-6°C, instead revealing strong sensitivity to a large range of temperatures. These findings are valuable for improving phenological models and predicting future spring phenology in a warming world.

Increased growing-season productivity drives earlier autumn leaf senescence in temperate trees.

Changes in the growing-season lengths of temperate trees greatly affect biotic interactions and global carbon balance. Yet future growing-season trajectories remain highly uncertain because the environmental drivers of autumn leaf senescence are poorly understood. Using experiments and long-term observations, we show that increases in spring and summer productivity due to elevated carbon dioxide, temperature, or light levels drive earlier senescence. Accounting for this effect improved the accuracy of senescence predictions by 27 to 42% and reversed future predictions from a previously expected 2- to 3-week delay over the rest of the century to an advance of 3 to 6 days. These findings demonstrate the critical role of sink limitation in governing the end of seasonal activity and reveal important constraints on future growing-season lengths and carbon uptake of trees.

Late-spring frost risk between 1959 and 2017 decreased in North America but increased in Europe and Asia.

Late-spring frosts (LSFs) affect the performance of plants and animals across the world's temperate and boreal zones, but despite their ecological and economic impact on agriculture and forestry, the geographic distribution and evolutionary impact of these frost events are poorly understood. Here, we analyze LSFs between 1959 and 2017 and the resistance strategies of Northern Hemisphere woody species to infer trees' adaptations for minimizing frost damage to their leaves and to forecast forest vulnerability under the ongoing changes in frost frequencies. Trait values on leaf-out and leaf-freezing resistance come from up to 1,500 temperate and boreal woody species cultivated in common gardens. We find that areas in which LSFs are common, such as eastern North America, harbor tree species with cautious (late-leafing) leaf-out strategies. Areas in which LSFs used to be unlikely, such as broad-leaved forests and shrublands in Europe and Asia, instead harbor opportunistic tree species (quickly reacting to warming air temperatures). LSFs in the latter regions are currently increasing, and given species' innate resistance strategies, we estimate that ∼35% of the European and ∼26% of the Asian temperate forest area, but only ∼10% of the North American, will experience increasing late-frost damage in the future. Our findings reveal region-specific changes in the spring-frost risk that can inform decision-making in land management, forestry, agriculture, and insurance policy.