Tree diversity enhances predation by birds but not by arthropods across climate gradients.

Tree diversity can promote both predator abundance and diversity. However, whether this translates into increased predation and top-down control of herbivores across predator taxonomic groups and contrasting environmental conditions remains unresolved. We used a global network of tree diversity experiments (TreeDivNet) spread across three continents and three biomes to test the effects of tree species richness on predation across varying climatic conditions of temperature and precipitation. We recorded bird and arthropod predation attempts on plasticine caterpillars in monocultures and tree species mixtures. Both tree species richness and temperature increased predation by birds but not by arthropods. Furthermore, the effects of tree species richness on predation were consistent across the studied climatic gradient. Our findings provide evidence that tree diversity strengthens top-down control of insect herbivores by birds, underscoring the need to implement conservation strategies that safeguard tree diversity to sustain ecosystem services provided by natural enemies in forests.

Factors affecting composition of fatty acids in wild-growing forest mushrooms.

The importance of mushrooms as a food source is continually increasing. To investigate how environmental factors affect the nutritional value of mushrooms, we harvested them in eastern Poland, south-central Germany, and northwestern Belgium in plots with similar environmental conditions but varying in tree species composition and richness. We used gas chromatography-mass spectrometry (GC-MS) to analyze the fatty acid (FA) content of the mushrooms. Fungal species identity explained the largest part (40%) of the total variation in FA concentration and composition. Environmental factors accounted for 1-12% of variation. The concentration of FA, especially saturated fatty acids, decreased with increasing understory cover and increasing nitrogen concentration in the topsoil. The effect of tree species richness or tree species identity was negligible. Our results suggest that the nutritional value of mushrooms depends mainly on the species identity of fungi, but that their FA content is slightly higher in forests with less undergrowth and in nitrogen-poor soils.

Unpunctual in diversity: The effect of stand species richness on spring phenology of deciduous tree stands varies among species and years.

Climatic drivers alone do not adequately explain the regional variation in budburst timing in deciduous forests across Europe. Stand-level factors, such as tree species richness, might affect budburst timing by creating different microclimates under the same site macroclimate. We assessed different phases of the spring phenology (start, midpoint, end, and overall duration of the budburst period) of four important European tree species (Betula pendula, Fagus sylvatica, Quercus robur and Tilia cordata) in monocultures and four-species mixture stands of a common garden tree biodiversity experiment in Belgium (FORBIO) in 2021 and 2022. Microclimatic differences between the stands in terms of bud chilling, temperature forcing, and soil temperature were considerable, with four-species mixtures being generally colder than monocultures in spring, but not in winter. In the colder spring of 2021, at the stand level, the end of the budburst period was advanced, and its overall duration shortened, in the four-species mixtures. At species level, this response was significant for F. sylvatica. In the warmer spring of 2022, advances in spring phenology in four-species stands were observed again in F. sylvatica and, less markedly, in B. pendula but without a general response at the stand level. Q. robur showed specific patterns with delayed budburst start in 2021 in the four-species mixtures and very short budburst duration for all stands in 2022. Phenological differences between monocultures and four-species mixtures were linked to microclimatic differences in light availability rather than in temperature as even comparatively colder microclimates showed an advanced phenology. Compared to weather conditions, tree species richness had a lower impact on budburst timing, but this impact can be of importance for key species like F. sylvatica and colder springs. These results indicate that forest biodiversity can affect budburst phenology, with wider implications, especially for forest- and land surface models.

Effects of plant diversity on productivity strengthen over time due to trait-dependent shifts in species overyielding.

Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems.

Trade-offs in biodiversity and ecosystem services between edges and interiors in European forests.

Forest biodiversity and ecosystem services are hitherto predominantly quantified in forest interiors, well away from edges. However, these edges also represent a substantial proportion of the global forest cover. Here we quantified plant biodiversity and ecosystem service indicators in 225 plots along forest edge-to-interior transects across Europe. We found strong trade-offs: phylogenetic diversity (evolutionary measure of biodiversity), proportion of forest specialists, decomposition and heatwave buffering increased towards the interior, whereas species richness, nectar production potential, stemwood biomass and tree regeneration decreased. These trade-offs were mainly driven by edge-to-interior structural differences. As fragmentation continues, recognizing the role of forest edges is crucial for integrating biodiversity and ecosystem service considerations into sustainable forest management and policy.

Limited effects of population age on the genetic structure of spatially isolated forest herb populations in temperate Europe.

Due to multiple land-cover changes, forest herb populations residing in forest patches embedded in agricultural landscapes display different ages and, thus, experience differences in genetic exchange, mutation accumulation and genetic drift. The extent of divergence in present-day population genetic structure among these populations of different ages remains unclear, considering their diverse breeding systems and associated pollinators. Answering this question is essential to understand these species' persistence, maintenance of evolutionary potential and adaptability to changing environments. We applied a multi-landscape setup to compare the genetic structure of forest herb populations across forest patches of different ages (18-338 years). We studied the impact on three common slow-colonizer herb species with distinct breeding systems and associated pollinators: Polygonatum multiflorum (outcrossing, long-distance pollinators), Anemone nemorosa (outcrossing, short-distance pollinators) and Oxalis acetosella (mixed breeding). We aimed to assess if in general older populations displayed higher genetic diversity and lower differentiation than younger ones. We also anticipated that P. multiflorum would show the smallest while O. acetosella the largest difference, between old and young populations. We found that older populations had a higher observed heterozygosity (H o) but a similar level of allelic richness (A r) and expected heterozygosity (H e) as younger populations, except for A. nemorosa, which exhibited higher A r and H e in younger populations. As populations aged, their pairwise genetic differentiation measured by D PS decreased independent of species identity while the other two genetic differentiation measures showed either comparable levels between old and young populations (G" ST) or inconsistency among three species (cGD). The age difference of the two populations did not explain their genetic differentiation. Synthesis: We found restricted evidence that forest herb populations with different ages differ in their genetic structure, indicating that populations of different ages can reach a similar genetic structure within decades and thus persist in the long term after habitat disturbance. Despite their distinct breeding systems and associated pollinators, the three studied species exhibited partly similar genetic patterns, suggesting that their common characteristics, such as being slow colonizers or their ability to propagate vegetatively, are important in determining their long-term response to land-cover change.

Recreational hazard: Vegetation and host habitat use correlate with changes in tick-borne disease hazard at infrastructure within forest stands.

Studies on density and pathogen prevalence of Ixodes ricinus indicate that vegetation and local host community drive much of their variation between green spaces. Contrarily, micro-geographic variation is understudied, although its understanding could reduce disease risk. We studied the density of infectious nymphal Ixodes sp. ("DIN", proxy for disease hazard), density of questing nymphs ("DON") and nymphal infection prevalence ("NIP") near recreational forest infrastructure. Drag sampling within forest stands and at adjacent benches and trails was combined with vegetation surveys, camera trapping hosts and pathogen screening of ticks. We analysed Borrelia burgdorferi s.l. and its genospecies, with complementary analyses on Rickettsia sp., Anaplasma phagocytophilum, Neoehrlichia mikurensis and Borrelia miyamotoi. DIN was highest in forest interior and at trails enclosed by forest. Lower disease hazard was observed at benches and trails at forest edges. This infrastructure effect can be attributed to variation in vegetation characteristics and the habitat use of tick hosts, specifically roe deer, rodents and songbirds. DON is the main driver of DIN at micro-geographic scale and negatively affected by infrastructure and forest edges. A positive association with vegetation cover in understorey and canopy was observed, as were positive trends for local rodent and songbird abundance. NIP of different pathogens was affected by different drivers. Lower B. burgdorferi s.l. prevalence in the interior of forest stands, driven by its most prevalent genospecies B. afzelii, points towards higher density of uninfected hosts there. B. afzelii was positively associated with understorey containing tall species and with high canopy cover, whereas local bird community composition predicts B. garinii prevalence. A positive effect of songbird abundance and a negative effect of pigeons were observed. Our findings support amplification and inhibition mechanisms within forest stands and highlight that the effect of established drivers of DIN may differ based on the considered spatial scale.

Combining multiple investigative approaches to unravel functional responses to global change in the understorey of temperate forests.

Plant communities are being exposed to changing environmental conditions all around the globe, leading to alterations in plant diversity, community composition, and ecosystem functioning. For herbaceous understorey communities in temperate forests, responses to global change are postulated to be complex, due to the presence of a tree layer that modulates understorey responses to external pressures such as climate change and changes in atmospheric nitrogen deposition rates. Multiple investigative approaches have been put forward as tools to detect, quantify and predict understorey responses to these global-change drivers, including, among others, distributed resurvey studies and manipulative experiments. These investigative approaches are generally designed and reported upon in isolation, while integration across investigative approaches is rarely considered. In this study, we integrate three investigative approaches (two complementary resurvey approaches and one experimental approach) to investigate how climate warming and changes in nitrogen deposition affect the functional composition of the understorey and how functional responses in the understorey are modulated by canopy disturbance, that is, changes in overstorey canopy openness over time. Our resurvey data reveal that most changes in understorey functional characteristics represent responses to changes in canopy openness with shifts in macroclimate temperature and aerial nitrogen deposition playing secondary roles. Contrary to expectations, we found little evidence that these drivers interact. In addition, experimental findings deviated from the observational findings, suggesting that the forces driving understorey change at the regional scale differ from those driving change at the forest floor (i.e., the experimental treatments). Our study demonstrates that different approaches need to be integrated to acquire a full picture of how understorey communities respond to global change.

Forest understorey flowering phenology responses to experimental warming and illumination.

Species are altering their phenology to track warming temperatures. In forests, understorey plants experience tree canopy shading resulting in light and temperature conditions, which strongly deviate from open habitats. Yet, little is known about understorey phenology responses to forest microclimates. We recorded flowering onset, peak, end and duration of 10 temperate forest understorey plant species in two mesocosm experiments to understand how phenology is affected by sub-canopy warming and how this response is modulated by illumination, which is related to canopy change. Furthermore, we investigated whether phenological sensitivities can be explained by species' characteristics, such as thermal niche. We found a mean advance of flowering onset of 7.1 d per 1°C warming, more than previously reported in studies not accounting for microclimatic buffering. Warm-adapted species exhibited greater advances. Temperature sensitivity did not differ between early- and later-flowering species. Experimental illumination did not significantly affect species' phenological temperature sensitivities, but slightly delayed flowering phenology independent from warming. Our study suggests that integrating sub-canopy temperature and light availability will help us better understand future understorey phenology responses. Climate warming together with intensifying canopy disturbances will continue to drive phenological shifts and potentially disrupt understorey communities, thereby affecting forest biodiversity and functioning.

Evaluating plant lineage losses and gains in temperate forest understories: a phylogenetic perspective on climate change and nitrogen deposition.

Global change has accelerated local species extinctions and colonizations, often resulting in losses and gains of evolutionary lineages with unique features. Do these losses and gains occur randomly across the phylogeny? We quantified: temporal changes in plant phylogenetic diversity (PD); and the phylogenetic relatedness (PR) of lost and gained species in 2672 semi-permanent vegetation plots in European temperate forest understories resurveyed over an average period of 40 yr. Controlling for differences in species richness, PD increased slightly over time and across plots. Moreover, lost species within plots exhibited a higher degree of PR than gained species. This implies that gained species originated from a more diverse set of evolutionary lineages than lost species. Certain lineages also lost and gained more species than expected by chance, with Ericaceae, Fabaceae, and Orchidaceae experiencing losses and Amaranthaceae, Cyperaceae, and Rosaceae showing gains. Species losses and gains displayed no significant phylogenetic signal in response to changes in macroclimatic conditions and nitrogen deposition. As anthropogenic global change intensifies, temperate forest understories experience losses and gains in specific phylogenetic branches and ecological strategies, while the overall mean PD remains relatively stable.

Les changements globaux accélèrent les processus de colonisation et d'extinction locales d'espèces, aboutissant à des gains ou à des pertes de lignées évolutives uniques. Ces gains et pertes se produisent-ils de manière aléatoire dans l'arbre phylogénétique ? Nous avons mesuré: les changements de diversité phylogénétique; et la parenté phylogénétique des espèces végétales gagnées ou perdues dans 2672 placettes semi-permanentes disposées dans le sous-bois de forêts tempérées d'Europe sur une période moyenne de 40 ans. Une fois corrigée par la richesse spécifique, la diversité phylogénétique a légèrement augmenté au cours du temps dans les différentes placettes. Les espèces perdues ont une plus grande parenté phylogénétique que les espèces gagnées. Les espèces gagnées sont donc issues d'un plus grand nombre de lignées évolutives que les espèces perdues. Certaines lignées ont gagné ou perdu davantage d'espèces que ce qui est prédit par le hasard : les Ericaceae, les Fabaceae et les Orchidaceae ayant davantage perdu, tandis que les Amaranthaceae, les Cyperaceae, et les Rosaceae ont plus gagné. Il n'y a pas de signal phylogénétique des gains ou pertes d'espèces en réponse aux changements de conditions macroclimatiques ou des dépôts atmosphériques d'azote. Alors que les changements globaux d'origine anthropique s'intensifient, les sous-bois des forêts tempérées connaissent des gains et des pertes de certaines lignées évolutives et de certaines stratégies écologiques, sans que la diversité phylogénétique moyenne ne s'en trouve véritablement affectée.

El cambio global ha acelerado las extinciones y colonizaciones a escala local, lo que a menudo ha supuesto pérdidas y ganancias de linajes evolutivos con características únicas. Ahora bien, ¿estas pérdidas y ganancias ocurren aleatoriamente a lo largo de la filogenia? Cuantificamos: los cambios temporales en la diversidad filogenética de las plantas; y la relación filogenética de las especies perdidas y ganadas en 2.672 parcelas de vegetación semipermanente en sotobosques templados europeos y re-muestreadas durante un período promedio de 40 años. Al controlar por las diferencias en la riqueza de especies, la diversidad filogenética aumentó ligeramente con el tiempo y entre parcelas. Además, las especies perdidas dentro de las parcelas exhibieron un mayor grado de relación filogenética que las especies ganadas. Esto implica que las especies ganadas se originaron en un conjunto de linajes evolutivos más diversos que las especies perdidas. Ciertos linajes también perdieron y ganaron más especies de las esperadas aleatoriamente: Ericaceae, Fabaceae y Orchidaceae experimentaron pérdidas y Amaranthaceae, Cyperaceae y Rosaceae mostraron ganancias. Las pérdidas y ganancias de especies no mostraron ninguna señal filogenética significativa en respuesta a los cambios en las condiciones macro-climáticas y la deposición de nitrógeno. A medida que se intensifica el cambio global antropogénico, los sotobosques temperados experimentan pérdidas y ganancias en ramas filogenéticas y estrategias ecológicas específicas, mientras que la diversidad filogenética media general permanece relativamente estable.

Impact of patch age and size on forest soil characteristics in European agricultural landscapes.

Many landscapes worldwide are characterized by the presence of a mosaic of forest patches with contrasting age and size embedded in a matrix of agricultural land. However, our understanding of the effects of these key forest patch features on the soil nutrient status (in terms of nitrogen, carbon, and phosphorus) and soil pH is still limited due to a lack of large-scale data. To address this research gap, we analyzed 830 soil samples from nearly 200 forest patches varying in age (recent versus ancient forests) and size (small versus larger patches) along a 2500-km latitudinal gradient across Europe. We also considered environmental covariates at multiple scales to increase the generality of our research, including variation in macroclimate, nitrogen deposition rates, forest cover in a buffer zone, basal area and soil type. Multiple linear mixed-effects models were performed to test the combined effects of patch features and environmental covariates on soil nutrients and pH. Recent patches had higher total soil phosphorus concentrations and stocks in the mineral soil layer, along with a lower nitrogen to phosphorus ratio within that layer. Small patches generally had a higher mineral soil pH. Mineral soil nitrogen stocks were lower in forest patches with older age and larger size, as a result of a significant interactive effect. Additionally, environmental covariates had significant effects on soil nutrients, including carbon, nitrogen, phosphorus, and their stoichiometry, depending on the specific covariates. In some cases, the effect of patch age on mineral soil phosphorus stocks was greater than that of environmental covariates. Our findings underpin the important roles of forest patch age and size for the forest soil nutrient status. Long-term studies assessing edge effects and soil development in post-agricultural forests are needed, especially in a context of changing land use and climate.

Usefulness of cultivar-level calibration of AquaCrop for vegetables depends on the crop and data availability.

As a result of climate change, climatic extremes are expected to increase. For high-value crops like vegetables, irrigation is a potentially economically viable adaptation measure in western Europe. To optimally schedule irrigation, decision support systems based on crop models like AquaCrop are increasingly used by farmers. High value vegetable crops like cauliflower or spinach are grown in two distinct growth cycles per year and, additionally, have a high turnover rate of new varieties. To successfully deploy the AquaCrop model in a decision support system, it requires a robust calibration. However, it is not known whether parameters can be conserved over both growth periods, nor whether a cultivar dependent model calibration is always required. Furthermore, when data are collected from farmers' fields, there are constraints in data availability and uncertainty. We collected data from commercial cauliflower and spinach fields in Belgium in 2019, 2020 and 2021 during different growing periods and of different cultivars. With the use of a Bayesian calibration, we confirmed the need for a condition or cultivar specific calibration for cauliflower, while for spinach, splitting the data per cultivar or pooling the data together did not improve uncertainty on the model simulations. However, due to uncertainties arising from field specific soil and weather conditions, or measurement errors from calibration data, real time field specific adjustments are advised to simulations when using AquaCrop as decision support tool. Remotely sensed or in situ ground data may be invaluable information to reduce uncertainty on model simulations.

Disentangling drivers of litter decomposition in a multi-continent network of tree diversity experiments.

Litter decomposition is a key ecosystem function in forests and varies in response to a range of climatic, edaphic, and local stand characteristics. Disentangling the relative contribution of these factors is challenging, especially along large environmental gradients. In particular, knowledge of the effect of management options, such as tree planting density and species composition, on litter decomposition would be highly valuable in forestry. In this study, we made use of 15 tree diversity experiments spread over eight countries and three continents within the global TreeDivNet network. We evaluated the effects of overstory composition (tree identity, species/mixture composition and species richness), plantation conditions (density and age), and climate (temperature and precipitation) on mass loss (after 3 months and 1 year) of two standardized litters: high-quality green tea and low-quality rooibos tea. Across continents, we found that early-stage decomposition of the low-quality rooibos tea was influenced locally by overstory tree identity. Mass loss of rooibos litter was higher under young gymnosperm overstories compared to angiosperm overstories, but this trend reversed with age of the experiment. Tree species richness did not influence decomposition and explained almost no variation in our multi-continent dataset. Hence, in the young plantations of our study, overstory composition effects on decomposition were mainly driven by tree species identity on decomposer communities and forest microclimates. After 12 months of incubation, mass loss of the high-quality green tea litter was mainly influenced by temperature whereas the low-quality rooibos tea litter decomposition showed stronger relationships with overstory composition and stand age. Our findings highlight that decomposition dynamics are not only affected by climate but also by management options, via litter quality of the identity of planted trees but also by overstory composition and structure.

Divergent roles of herbivory in eutrophying forests.

Ungulate populations are increasing across Europe with important implications for forest plant communities. Concurrently, atmospheric nitrogen (N) deposition continues to eutrophicate forests, threatening many rare, often more nutrient-efficient, plant species. These pressures may critically interact to shape biodiversity as in grassland and tundra systems, yet any potential interactions in forests remain poorly understood. Here, we combined vegetation resurveys from 52 sites across 13 European countries to test how changes in ungulate herbivory and eutrophication drive long-term changes in forest understorey communities. Increases in herbivory were associated with elevated temporal species turnover, however, identities of winner and loser species depended on N levels. Under low levels of N-deposition, herbivory favored threatened and small-ranged species while reducing the proportion of non-native and nutrient-demanding species. Yet all these trends were reversed under high levels of N-deposition. Herbivores also reduced shrub cover, likely exacerbating N effects by increasing light levels in the understorey. Eutrophication levels may therefore determine whether herbivory acts as a catalyst for the "N time bomb" or as a conservation tool in temperate forests.

Land use and soil characteristics affect soil organisms differently from above-ground assemblages.

BACKGROUND: Land-use is a major driver of changes in biodiversity worldwide, but studies have overwhelmingly focused on above-ground taxa: the effects on soil biodiversity are less well known, despite the importance of soil organisms in ecosystem functioning. We modelled data from a global biodiversity database to compare how the abundance of soil-dwelling and above-ground organisms responded to land use and soil properties. RESULTS: We found that land use affects overall abundance differently in soil and above-ground assemblages. The abundance of soil organisms was markedly lower in cropland and plantation habitats than in primary vegetation and pasture. Soil properties influenced the abundance of soil biota in ways that differed among land uses, suggesting they shape both abundance and its response to land use. CONCLUSIONS: Our results caution against assuming models or indicators derived from above-ground data can apply to soil assemblages and highlight the potential value of incorporating soil properties into biodiversity models.

Zoonotic pathogens linked with hedgehog diphtheric disease.

Hedgehog diphtheric disease (HDD), an ulcerative skin disease with a high fatality rate, is an emerging threat to European hedgehogs (Erinaceus europaeus). We explored the potential role of a panel of zoonotic pathogens in the presumed multifactorial nature of HDD in 188 hedgehogs from 3 wildlife rescue centres in Belgium. As expected, and with a prevalence of 67% in 57 hedgehogs with skin lesions, characteristic of HDD, the occurrence of Corynebacterium ulcerans was strongly associated with the disease. Remarkably, with a prevalence of 42% in affected animals, infections with Borrelia burgdorferi sensu lato were 3.92 times more likely to be detected in HDD (95% confidence interval: 1.650-9.880; p = .0024). Overall, 40 hedgehogs tested positive for the B. burgdorferi sensu lato complex, including Borrelia afzelii (n = 30), Borrelia bavariensis (n = 7) and Borrelia spielmanii (n = 7). Other widely occurring pathogens included Salmonella (prevalence of 19%, with three pulsed-field gel electrophoresis profiles) and Leptospira sp. (prevalence of 11%, including Leptospira interrogans and Leptospira borgpetersenii), but these were not associated with the occurrence of HDD. These findings show that hedgehogs in Belgium represent a significant reservoir of multiple zoonotic bacteria, of which toxigenic C. ulcerans and B. burgdorferi sensu lato are associated with widespread hedgehog skin pathology and mortality.

Survival in the understorey: Testing direct and indirect effects of microclimatological changes on Ixodes ricinus.

The distribution of ticks in the Ixodes ricinus species complex is partly driven by climate, with temperature and relative humidity affecting survival. These variables are driven by macroclimate, but vary locally due to microclimate buffering. This buffering has been suggested to be one of the driving forces behind variation in tick survival and density in time and space. In order to understand the role of the herb layer with respect to this variation, we deployed I. ricinus within an existing experimental setup studying the response of forest understorey to micrometeorological changes. This allowed for the analysis of both direct effects of warming on tick survival in controlled field conditions, as well as indirect effects through changes in herb layer biomass. Herb layer biomass estimates were observed to be higher in plots that had been experimentally warmed, with a trend towards higher survival in these warmed plots. This marginal increase in survival rate may be due to increased microclimate buffering. Comparing our results to literature implies that canopy and shrub layer vegetation have a larger effect on climate buffering, and therefore also on tick survival. Since the herb layer biomass is expected to increase due to global warming and increased frequency of disturbance-induced canopy gaps, survival in forested habitats may increase in the future. This would increase the difference in survival compared to that in open habitats.

Forest structure and composition alleviate human thermal stress.

Current climate change aggravates human health hazards posed by heat stress. Forests can locally mitigate this by acting as strong thermal buffers, yet potential mediation by forest ecological characteristics remains underexplored. We report over 14 months of hourly microclimate data from 131 forest plots across four European countries and compare these to open-field controls using physiologically equivalent temperature (PET) to reflect human thermal perception. Forests slightly tempered cold extremes, but the strongest buffering occurred under very hot conditions (PET >35°C), where forests reduced strong to extreme heat stress day occurrence by 84.1%. Mature forests cooled the microclimate by 12.1 to 14.5°C PET under, respectively, strong and extreme heat stress conditions. Even young plantations reduced those conditions by 10°C PET. Forest structure strongly modulated the buffering capacity, which was enhanced by increasing stand density, canopy height and canopy closure. Tree species composition had a more modest yet significant influence: that is, strongly shade-casting, small-leaved evergreen species amplified cooling. Tree diversity had little direct influences, though indirect effects through stand structure remain possible. Forests in general, both young and mature, are thus strong thermal stress reducers, but their cooling potential can be even further amplified, given targeted (urban) forest management that considers these new insights.

Soil geochemistry - and not topography - as a major driver of carbon allocation, stocks, and dynamics in forests and soils of African tropical montane ecosystems.

The lack of field-based data in the tropics limits our mechanistic understanding of the drivers of net primary productivity (NPP) and allocation. Specifically, the role of local edaphic factors - such as soil parent material and topography controlling soil fertility as well as water and nutrient fluxes - remains unclear and introduces substantial uncertainty in understanding net ecosystem productivity and carbon (C) stocks. Using a combination of vegetation growth monitoring and soil geochemical properties, we found that soil fertility parameters reflecting the local parent material are the main drivers of NPP and C allocation patterns in tropical montane forests, resulting in significant differences in below- to aboveground biomass components across geochemical (soil) regions. Topography did not constrain the variability in C allocation and NPP. Soil organic C stocks showed no relation to C input in tropical forests. Instead, plant C input seemingly exceeded the maximum potential of these soils to stabilize C. We conclude that, even after many millennia of weathering and the presence of deeply developed soils, above- and belowground C allocation in tropical forests, as well as soil C stocks, vary substantially due to the geochemical properties that soils inherit from parent material.

Increasing calcium scarcity along Afrotropical forest succession.

Secondary forests constitute an increasingly important component of tropical forests worldwide. Although cycling of essential nutrients affects recovery trajectories of secondary forests, the effect of nutrient limitation on forest regrowth is poorly constrained. Here we use three lines of evidence from secondary forest succession sequences in central Africa to identify potential nutrient limitation in regrowing forests. First, we show that atmospheric phosphorus supply exceeds demand along forest succession, whereas forests rely on soil stocks to meet their base cation demands. Second, soil nutrient metrics indicate that available phosphorus increases along the succession, whereas available cations decrease. Finally, fine root, foliar and litter stoichiometry show that tissue calcium concentrations decline relative to those of nitrogen and phosphorus during succession. Taken together, these observations suggest that calcium becomes an increasingly scarce resource in central African forests during secondary succession. Furthermore, ecosystem calcium storage shifts from soil to woody biomass over succession, making it a vulnerable nutrient in the wake of land-use change scenarios that involve woody biomass export. Our results thus call for a broadened focus on elements other than nitrogen and phosphorus regarding tropical forest biogeochemical cycles and identify calcium as a scarce and potentially limiting nutrient in an increasingly disturbed and dynamic tropical forest landscape.