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.

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.

Changes in above- versus belowground biomass distribution in permafrost regions in response to climate warming.

Permafrost regions contain approximately half of the carbon stored in land ecosystems and have warmed at least twice as much as any other biome. This warming has influenced vegetation activity, leading to changes in plant composition, physiology, and biomass storage in aboveground and belowground components, ultimately impacting ecosystem carbon balance. Yet, little is known about the causes and magnitude of long-term changes in the above- to belowground biomass ratio of plants (η). Here, we analyzed η values using 3,013 plots and 26,337 species-specific measurements across eight sites on the Tibetan Plateau from 1995 to 2021. Our analysis revealed distinct temporal trends in η for three vegetation types: a 17% increase in alpine wetlands, and a decrease of 26% and 48% in alpine meadows and alpine steppes, respectively. These trends were primarily driven by temperature-induced growth preferences rather than shifts in plant species composition. Our findings indicate that in wetter ecosystems, climate warming promotes aboveground plant growth, while in drier ecosystems, such as alpine meadows and alpine steppes, plants allocate more biomass belowground. Furthermore, we observed a threefold strengthening of the warming effect on η over the past 27 y. Soil moisture was found to modulate the sensitivity of η to soil temperature in alpine meadows and alpine steppes, but not in alpine wetlands. Our results contribute to a better understanding of the processes driving the response of biomass distribution to climate warming, which is crucial for predicting the future carbon trajectory of permafrost ecosystems and climate feedback.

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.

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.

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.

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.

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.

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.

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).

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.

Interactive climate factors restrict future increases in spring productivity of temperate and boreal trees.

Climate warming is currently advancing spring leaf-out of temperate and boreal trees, enhancing net primary productivity (NPP) of forests. However, it remains unclear whether this trend will continue, preventing for accurate projections of ecosystem functioning and climate feedbacks. Several ecophysiological mechanisms have been proposed to regulate the timing of leaf emergence in response to changing environmental cues, but the relative importance of those mechanisms remains unclear. Here, we use 727,401 direct phenological observations of common European forest trees to examine the dominant controls on leaf-out. Using the emerging mechanisms, we forecast future trajectories of spring arrival and evaluate the consequences for forest carbon dynamics. By representing hypothesized relationships with autumn temperature, winter chilling, and the timing of spring onset, we accurately predicted reductions in the advance of leaf-out. There was a strong consensus between our empirical model and existing process-based models, revealing that the advance in leaf-out will not exceed 2 weeks over the rest of the century. We further estimate that, under a 'business-as-usual' climate scenario, earlier spring arrival will enhance NPP of temperate and boreal forests by ~0.2 Gt per year at the end of the century. In contrast, previous estimates based on a simple degree-day model range around 0.8 Gt. As such, the expected NPP is drastically reduced in our updated model relative to previous estimates-by a total of ~25 Gt over the rest of the century. These findings reveal important environmental constraints on the productivity of broad-leaved deciduous trees and highlight that shifting spring phenology is unlikely to slow the rate of warming by offsetting anthropogenic carbon emissions.

Narrow habitat breadth and late-summer emergence increases extinction vulnerability in Central European bees.

Evaluating intrinsic and extrinsic traits that predispose species to local extinction is important for targeting conservation efforts. Among the species of special concern in Europe are bees, which, along with butterflies, are the best monitored insects. Bees are most species-rich in Mediterranean-type climates with short winters, warm springs, and dry summers. In Central Europe, climate warming per se is, therefore, expected to benefit most bee species, while pesticides and the loss of habitats and plant diversity should constitute threats. Here, we use the bee fauna of Germany, which has been monitored for Red Lists for over 40 years, to analyse the effects of habitat breadth, pollen specialization, body size, nesting sites, sociality, duration of flight activity, and time of emergence during the season. We tested each factor's predictive power against changes in commonness and Red List status, using phylogenetically informed hierarchical Bayesian (HB) models. Extinction vulnerability is strongly increased in bees flying in late summer, with a statistical model that included flight time, habitat preference, and duration of activity correctly predicting the vulnerability status of 85% of the species. Conversely, spring emergence and occurrence in urban areas each reduce vulnerability, pointing to intensive land use especially harming summer-active bees, with the combination of these factors currently shifting Germany's bee diversity towards warm-adapted, spring-flying, city-dwelling species.

The occurrence of red and yellow autumn leaves explained by regional differences in insolation and temperature.

Red or yellow autumn leaves have long fascinated biologists, but their geographical concentration in trees in Eastern North America (ENA) has defied evolutionary explanations. In this review, anthocyanins and xanthophylls are discussed in relation to their occurrence in different regions of the Northern Hemisphere, phylogenetic distribution and photoprotective function during the breakdown of chlorophylls. Pigments in senescing leaves that intercept incident light and dissipate the absorbed energy extend the time available for nutrient resorption. Experiments with Arabidopsis have revealed greatest anthocyanin photoprotective function at low temperatures and high light intensities, and high-resolution solar irradiation maps reveal that ENA and Asia receive higher irradiation than does Europe. In addition, ENA experiences higher temperature fluctuations in autumn, resulting in cold snaps during leaf senescence. Under common garden conditions, chlorophyll degradation occurs earlier in ENA species than in their European and East Asian relatives. In combination, strong solar irradiation, temperature fluctuations and, on average, 3-wk shorter vegetation periods of ENA species favour investment in pigments to extend the time for nutrient resorption before abscission, explaining the higher frequency of coloured species in ENA compared to Europe. We end by outlining research that could test this new explanation of bright New England autumns.

Increased autumn productivity permits temperate trees to compensate for spring frost damage.

Climate warming is leading to earlier budburst and therefore an increased risk of spring frost injury to young leaves. But to what extent are second-cohort leaves, which trees put out after leaf-killing frosts, able to compensate incurred losses? To investigate whether second-cohort leaves behave differently from first-cohort leaves, we exposed saplings of beech (Fagus sylvatica), oak (Quercus robur), and honeysuckle (Lonicera xylosteum) to experimental treatments mimicking either a warm spring or a warm spring with a leaf-killing frost. Refoliation took 48, 43, and 36 d for beech, oak and honeysuckle, respectively. In beech and oak, autumn Chl content and photosynthesis rates were higher in second- than in first-cohort leaves, senescence in second-cohort leaves occurred c. 2-wk-later, and autumn bud growth in beech was elevated 66% in frost-damaged plants compared with the warm spring treatment. No differences in autumn phenology and growth were observed for honeysuckle. Overall, in beech and oak, delayed Chl breakdown in second-cohort leaves mitigated 31% and 25%, respectively, of the deficit in growing-season length incurred by spring frost damage. These results reveal an unexpected ability of second-cohort leaves of beech and oak to compensate for spring frost damage, and demonstrate that long-lived trees vary their autumnal phenology depending on preceding productivity.

Shortened temperature-relevant period of spring leaf-out in temperate-zone trees.

Temperature during a particular period prior to spring leaf-out, the temperature-relevant period (TRP), is a strong determinant of the leaf-out date in temperate-zone trees. Climatic warming has substantially advanced leaf-out dates in temperate biomes worldwide, but its effect on the beginning and length of the TRP has not yet been explored, despite its direct relevance for phenology modeling. Using 1,551 species-site combinations of long-term (1951-2016) in situ observations on six tree species (namely, Aesculus hippocastanum, Alnus glutinosa, Betula pendula, Fagus sylvatica, Fraxinus excelsior, and Quercus robur) in central Europe, we found that the advancing leaf-out was accompanied by a shortening of the TRP. On average across all species and sites, the length of the TRP significantly decreased by 23% (p < .05), from 60 ± 4 days during 1951-1965 to 47 ± 4 days during 2002-2016. Importantly, the average start date of the TRP did not vary significantly over the study period (March 2-5, DOY = 61-64), which could be explained by sufficient chilling over the study period in the regions considered. The advanced leaf-out date with unchanged beginning of the TRP can be explained by the faster accumulation of the required heat due to climatic warming, which overcompensated for the retarding effect of shortening daylength on bud development. This study shows that climate warming has not yet affected the mean TRP starting date in the study region, implying that phenology modules in global land surface models might be reliable assuming a fixed TRP starting date at least for the temperate central Europe. Field warming experiments do, however, remain necessary to test to what extent the length of TRP will continue to shorten and whether the starting date will remain stable under future climate conditions.

Examining the support-supply and bud-packing hypotheses for the increase in toothed leaf margins in northern deciduous floras.

PREMISE: The proportion of woody dicots with toothed leaves increases toward colder regions, a relationship used to reconstruct past mean annual temperatures. Recent hypotheses explaining this relationship are that (1) leaves in colder regions are thinner, requiring thick veins for support and water supply, with the resulting craspedodromous venation leading to marginal teeth (support-supply hypothesis) or that (2) teeth are associated with the packing of leaf primordia in winter buds (bud-packing hypothesis). METHODS: We addressed these hypotheses by examining leaf thickness, number of primordia in buds, growing season length (mean annual temperature, MAT), and other traits in 151 deciduous woody species using georeferenced occurrences and a Bayesian model controlling for phylogeny. We excluded evergreen species because longer leaf life spans correlate with higher leaf mass per area, precluding the detection of independent effects of leaf thickness on leaf-margin type. RESULTS: The best model predicted toothed leaves with 94% accuracy, with growing season length the strongest predictor. Neither leaf thickness nor number of leaves preformed in buds significantly influenced margin type, rejecting the support-supply and bud-packing hypotheses. CONCLUSIONS: A direct selective benefit of leaf teeth via a carbon gain early in the spring as proposed by Royer and Wilf (2006) would match the strong correlation between toothed species occurrence and short growing season found here using Bayesian hierarchical models. Efforts should be directed to physiological work quantifying seasonal photosynthate production in toothed and nontoothed leaves.

Ongoing seasonally uneven climate warming leads to earlier autumn growth cessation in deciduous trees.

Ongoing global warming is causing phenological shifts that affect photosynthesis and growth rates in temperate woody species. However, the effects of seasonally uneven climate warming-as is occurring in much of Europe, where the winter/spring months are warming twice as fast than the summer/autumn months-on autumn growth cessation (completion of overwintering buds) and leaf senescence, and possible carry-over effects between phenophases, remain under-investigated. We conducted experiments in which we exposed saplings of canopy and understory species to 4 °C warming in winter/spring, summer/autumn, or all year to disentangle how the timing of bud break, bud set completion, and leaf senescence is affected by seasonally uneven warming. All-year warming led to significantly delayed leaf senescence, but advanced bud set completion; summer/autumn warming only delayed leaf senescence; and winter/spring warming advanced both bud set and senescence. The non-parallel effects of warming on bud completion and leaf senescence show that leaf senescence alone is an inadequate proxy for autumn growth cessation in trees and counterintuitively suggest that continued uneven seasonal warming will advance cessation of primary growth in autumn, even when leaf senescence is delayed. Phenological responses to warming treatments (earlier spring onset, later autumn senescence) were more than twice as high in understory species than in canopy species, which can partly be explained by the absence of carry-over effects among phenophases in the former group. This underscores the need to consider differences among plant functional types when forecasting the future behaviour of ecosystems.