Tree Demography Research Articles

Intraseasonal interactive effects of successive typhoons characterize canopy damage of forests in Taiwan: A remote sensing-based assessment

Wind gusts and rainfall from tropical cyclones can heavily damage forest canopies, leading to abrupt changes in forest structure and tree demography. Although many studies have shown that successive tropical cyclones can interact with each other through residual effects, the role of past disturbances is unclear because they may lead to damage amplification of the second cyclone because of weakened forest structure, or damage reduction of the second cyclone because of previous damage to susceptible trees. We investigated the interaction between consecutive cyclones between 2001 and 2017 for five well-conserved forests in Taiwan, which experiences an average of 1.75 typhoons annually. Using MODIS imagery, we computed the typhoon-induced change of a canopy vegetation index, the Normalized Difference Infrared Index (NDII). The effects of successive typhoons were assessed separately for typhoons occurring within a single year (annual analysis) and within two consecutive years (biennial analysis). We used mixed effect models of reductions in NDII, a measurement of canopy damage, in relation to target and past typhoon characteristics and damage magnitude. NDII reduction induced by preceding typhoons was slightly more important and statistically significant in explaining the variation in NDII reduction associated with the target typhoon in the annual than in the biennial analysis, where the effect was non-significant. Canopy damage did not always decrease across typhoons occurring within the same season, however, for most successive typhoons in the biennial analysis, the second cyclone caused equal or less canopy damage (16 out of 21 typhoon pairs). These results support the idea that residual interactive effects of previous typhoons decrease quickly over time and rarely last for several typhoon seasons for Taiwanese forests, contributing to their high resistance to frequent typhoon disturbance.

Ecological Dynamics and Regeneration Expansion of Treeline Ecotones in Response to Climate Change in Northern Bhutan Himalayas

The alpine treeline ecotones are an early indicator of vegetation’s response to changes in climate, and the advancement of diffuse treeline ecotones has been associated with mean annual warming temperatures. However, the knowledge of how tree demographic size, age and population distribution, and regeneration decrease with increasing elevation and mean annual temperature remain fragmentary in Bhutan. There was no explanation of how treelines migrate in response to the climate. Therefore, the objectives of this study were to investigate tree demographic size and age and population distribution, as well as the regeneration expansion of treeline ecotones of Abies densa trees in response to climate change. Demographic data from thirty transect bands from treeline ecotones and reconstructed mean annual temperatures from tree-rings were used. Regression analysis was used to establish a relationship between elevation/temperature and demographic tree size and age, as well as to determine recruitment frequency distributions and whether these could be driven by climate change. The tree demography indicated that the treeline ecotone in our sampling site is temperature limited. Hence, cooler temperatures at higher elevations should drive decreases in basal diameter, age and recruitment frequencies. From the dendroecological analysis, the diffuse treeline ecotones appear to be climbing on average 1.00 m per year in Northern Bhutan. We also found that the recruitment frequency has increased over recent years (1850–2017), as temperatures continue to rise. The thermal treeline ecotones will be likely to serve as a line of bioclimatic reference against which other zones of bioclimate can be defined. With documented responses of treeline ecotones toward mean annual temperatures, the expectation is that additional warming will continue to influence regeneration expansion in the future. This dynamic response of treeline ecotones towards the climate acts as an indicator of climate change. Information about climbing treelines and altered ecotones should be a vital part of the material for decision makers to consider, to assess impacts and threats to Himalayan alpine biota.

Demographic consequences of heterogeneity in conspecific density dependence among mast-fruiting tropical trees.

The role of conspecific density dependence (CDD) in the maintenance of species richness is a central focus of tropical forest ecology. However, tests of CDD often ignore the integrated effects of CDD over multiple life stages and their long-term impacts on population demography. We combined a 10-year time series of seed production, seedling recruitment and sapling and tree demography of three dominant Southeast Asian tree species that adopt a mast-fruiting phenology. We used these data to construct individual-based models that examine the effects of CDD on population growth rates (λ) across life-history stages. Recruitment was driven by positive CDD for all species, supporting the predator satiation hypothesis, while negative CDD affected seedling and sapling growth of two species, significantly reducing λ. This negative CDD on juvenile growth overshadowed the positive CDD of recruitment, suggesting the cumulative effects of CDD during seedling and sapling development has greater importance than the positive CDD during infrequent masting events. Overall, CDD varied among positive, neutral and negative effects across life-history stages for all species, suggesting that assessments of CDD on transitions between just two stages (e.g. seeds seedlings or juveniles mature trees) probably misrepresent the importance of CDD on population growth and stability.

Effects of fire disturbance on species and functional compositions vary with tree sizes in a tropical dry forest.

BackgroundDisturbances are crucial in determining forest biodiversity, dynamics, and ecosystem functions. Surface fire is a significant disturbance in tropical forests, but research on the effect of surface fire on structuring species and functional composition in a community through time remains scarce. Using a 20-year dataset of tree demography in a seasonal evergreen tropical forest in Thailand, we specifically addressed two essential questions: (1) What is the pattern of temporal turnover in species and functional composition in a community with frequent fire disturbance? (2) How did the temporal turnover vary with tree size?MethodsWe analyzed species compositional and functional temporal turnovers in four different tree size classes among five tree censuses. We quantified species turnover by calculating Bray-Curtis dissimilarity, and investigated its underlying mechanisms by comparing pairwise dissimilarity of functional traits with simulations from null models. If fire disturbances contribute more to a stochastic process, the functional composition would display a random pattern. However, if they contribute more towards a deterministic process, the functional composition should reveal a non-random pattern.ResultsOver 20 years (1994–2014), we observed changes in species composition, whereas functional composition remained relatively stable. The temporal turnover patterns of species and functional compositions varied with tree sizes. In particular, temporal functional turnover shifted very little for large trees, suggesting that changes in species composition of larger trees are contributed by species with similar functional traits through time. The temporal functional composition turnovers of smaller trees (DBH ≤ 5 cm) were mostly at random. We detected a higher functional turnover than expected by null models in some quadrats throughout the 50-ha study plot, and their observed turnover varied with diameter classes.ConclusionsSpecies compositional changes were caused by changes in the abundance of species with similar functional traits through time. Temporal functional turnover in small trees was random in most quadrats, suggesting that the recruits came from the equal proportions of surviving trees and new individuals of fast-growing species, which increased rapidly after fires. On the other hand, functional composition in big trees was more likely determined by surviving trees which maintained higher functional similarities than small trees through time. Fire disturbance is important for ecosystem functions, as changing forest fire frequency may alter forest turnover, particularly in functional composition in the new recruits of this forest.

Sapling growth gradients interact with homogeneous disturbance regimes to explain savanna tree cover discontinuities

AbstractSavanna tree cover often exhibits sudden discontinuities across space. It has been proposed that local spatial processes imposed by variation in tree cover itself (as opposed to by external drivers such as edaphic variation) can reinforce such discontinuities. Despite this, we generally lack data on tree demography and the environmental drivers affecting the former as a function of tree neighborhoods in these systems. Given the importance of disturbance traps in savannas, spatial processes affecting the likelihood of escape from the seedling/sapling stage to the adult tree stage are likely to be critical. In a longitudinal survey of 800 saplings distributed along eight 1‐km transects spanning woodland–grassland transitions in Serengeti National Park, we found a positive association between tree cover and sapling growth and survival, but no relationship with sapling abundance, maximum tree height, disturbance, or topkill. Based on microclimate and soil moisture dynamics data, we found no evidence to suggest that tree cover itself drives variation in growth. Based on a prior analysis of soil properties along these transects, we hypothesized that underlying edaphic conditions may be responsible for variation in growth. Regardless of the underlying mechanism, we used simulations to show that subtle growth rate gradients interacted with intense disturbance regimes to produce sharp discontinuities in tree cover, with strong demographic bottlenecks where growth is slowest, explaining the observed patterns of tree cover along the transects. Our results indicated that disturbance and herbivory are equally intense in areas of high and low tree cover, and that although trees have the potential to successfully establish and reach adulthood in open, grassy sites, they grow too slowly to escape disturbance traps there. Importantly, we showed that although herbivory and fire are fundamental for explaining savanna structural patterns, their effects are not necessarily reinforced by tree cover itself.

Effect of tree demography and flexible root water uptake for modeling the carbon and water cycles of Amazonia

Amazonian forest plays a crucial role in regulating the carbon and water cycles in the global climate system. However, the representation of biogeochemical fluxes and forest structure in dynamic global vegetation models (DGVMs) remains challenging. This situation has considerable implications to simulate the state and dynamics of Amazonian forest. This study aims at simulating the dynamic of the evapotranspiration (ET), productivity (GPP), biomass (AGB) and forest structure of wet tropical forests in the Amazon basin using the updated ORCHIDEE land surface model. The latter is improved for two processes: stand structure and demography, and plant water uptake by roots. Stand structure is simulated by adapting the CAN version of ORCHIDEE, originally developed for temperate forests. Here, we account for the permanent recruitment of young individual trees, the distribution of stand level growth into 20 different cohorts of variable diameter classes, and mortality due to asymmetric competition for light. Plant water uptake is simulated by including soil-to-root hydraulic resistance (RS). To evaluate the effect of the soil resistance alone, we performed factorial simulations with demography only (CAN) and both demography and resistance (CAN-RS). AGB, ET and GPP outputs of CAN-RS are also compared with the standard version of ORCHIDEE (TRUNK) for which eco-hydrological parameters were tuned globally to fit GPP and evapotranspiration at flux tower sites. All the model versions are benchmarked against in situ and regional datasets. We show that CAN-RS correctly reproduce stand level structural variables (as CAN) like diameter classes and tree densities when validated using in-situ data. Besides offering the key advantage to simulate forest's structure, it also correctly simulates ET and GPP and improves fluxes spatial patterns when compared to TRUNK. With the new formulation of soil water uptake, which is driven by soil water availability rather than root-biomass, the simulated trees preferentially use water in the deepest soil layers during the dry seasons. This improves the seasonality of ET and GPP compared to CAN, especially on clay soils for which the soil moisture potential drops rapidly in the dry season. Nevertheless, since demography parameters in CAN-RS are constant for all evergreen tropical forests, spatial variability of AGB and basal area across the Amazon remains too uniform compared to observations, and are very comparable to the TRUNK. Additional processes such as climate driven mortality and phosphorus limitation on growth leading to the prevalence of species with different functional traits across the Amazon need to be included in the future development of this model.

Twenty‐five years of tree demography in a frequently burned oak woodland: implications for savanna restoration

AbstractDue to decades of fire suppression, much of the Upper Midwest savanna habitat has converted to oak woodland. In efforts to restore oak savanna habitat, fire has been re‐introduced in many of these woodlands. A primary purpose of these burns is to kill the fire‐sensitive mesophytic tree species, which had established themselves during the decades of fire suppression, reduce the number of understory trees, and preserve the larger more widely spaced oaks. It is clear from ongoing efforts that restoring oak savannas will require frequent fires over decades, but frequent fires over the long term can also threaten the desirable oaks. Long‐term demographic studies at savanna restoration sites experiencing frequent fires are necessary to determine the extent to the frequent burns are supporting and/or confounding restoration goals. Results presented here are from a twenty‐five‐year demographic study of an Upper Midwest bur oak (Quercus macrocarpa) savanna/woodland experiencing frequent fire, during which both the survival and growth of more than 9000 trees were documented. Survival was assessed annually and growth every five years. In the face of frequent fires, stem survival was found to be strongly associated with tree species, stem size, and stem growth. In turn, stem growth was found to be related to tree species and stem size. Decades of frequent burning in this oak woodland have substantially reduced the abundance of unwanted trees, specifically mesophytic species and Quercus ellipsoidalis, the latter which outcompetes Q. macrocarpa in the absence of fire. While Q. macrocarpa mid‐sized (10–25 cm dbh) and large (≥25 cm dbh) trees are quite resistant to fire and now dominate the savanna landscape, they are not immune from fire‐induced mortality. It is recommended that the number and density of these trees should be re‐evaluated every few years to ensure that desirable numbers remain. If necessary, fires should be suspended for a period of time. This will give smaller Q. macrocarpa trees time to grow larger and become more fire‐resistant, thereby ensuring successive generations of Q. macrocarpa.

Intermediate disturbances are a key driver of long-term tree demography across old-growth temperate forests.

Disentangling the relative influence of background versus disturbance related mortality on forest demography is crucial for understanding long‐term dynamics and predicting the influence of global change on forests. Quantifying the rates and drivers of tree demography requires direct observations of tree populations over multiple decades, yet such studies are rare in old‐growth forest, particularly in the temperate zone of Europe. We use multi‐decade (1980–2020) monitoring of permanent plots, including observations of mode of mortality and disturbance events, to quantify rates and drivers of tree demography across a network of old‐growth remnants in temperate mountain forests of Slovenia. Annual rates of mortality and recruitment varied markedly among sites and over time; census intervals that captured intermediate severity canopy disturbances caused subtle peaks in annual mortality (e.g., >2%/year), while rates of background mortality in non‐disturbed intervals averaged about 1%/year. Roughly half of the trees died from modes of mortality associated with disturbance (i.e., uprooting or snapped‐alive). Results of a Bayesian multilevel model indicate that beech (Fagus sylvatica) had a higher likelihood of disturbance related mortality compared to fir (Abies alba), which mainly died standing, and there was a notable increase in the odds of disturbance mortality with increasing diameter for all species. Annual recruitment rates were consistently low at sites (<0.5%) that lacked evidence of disturbance, but often exceeded 3% on sites with higher levels of past canopy mortality. Recruitment was dominated by beech on sites with more diffuse background mortality, while the less shade tolerant maple (Acer pseudoplatanus) recruited following known disturbance events. Our study highlights the important role of stand‐scale, partial canopy disturbance for long‐term forest demography. These results suggest that subtle climate‐driven changes in the regime of intermediate severity disturbances could have an important influence on future forest dynamics and warrant attention.

Influence of Pleistocene climatic oscillations on the phylogeography and demographic history of endemic vulnerable trees (section Magnolia) of the Tropical Montane Cloud Forest in Mexico.

The Tropical Montane Cloud Forest (TMCF) is a highly dynamic ecosystem that has undergone frequent spatial changes in response to the interglacial-glacial cycles of the Pleistocene. These climatic fluctuations between cold and warm cycles have led to species range shifts and contractions-expansions, resulting in complex patterns of genetic structure and lineage divergence in forest tree species. In this study, we sequenced four regions of the chloroplast DNA (trnT-trnL, trnK5-matk, rpl32-trnL, trnS-trnG) for 20 populations and 96 individuals to evaluate the phylogeography, historical demography, and paleodistributions of vulnerable endemic TMCF trees in Mexico: Magnolia pedrazae (north-region), M. schiedeana (central-region), and M. schiedeana population Oaxaca (south-region). Our data recovered 49 haplotypes that showed a significant phylogeographic structure in three regions: north, central, and south. Bayesian Phylogeographic and Ecological Clustering (BPEC) analysis also supported the divergence in three lineages and highlighted the role of environmental factors (temperature and precipitation) in genetic differentiation. Our historical demography analyses revealed demographic expansions predating the Last Interglacial (LIG, ~125,000 years ago), while Approximate Bayesian Computation (ABC) simulations equally supported two contrasting demographic scenarios. The BPEC and haplotype network analyses suggested that ancestral haplotypes were geographically found in central Veracruz. Our paleodistributions modeling showed evidence of range shifts and expansions-contractions from the LIG to the present, which suggested the complex evolutionary dynamics associated to the climatic oscillations of the Pleistocene. Habitat management of remnant forest fragments where large and genetically diverse populations occur in the three TMCF regions analyzed would be key for the conservation of these magnolia populations.

Genome assemblies for two Neotropical trees: Jacaranda copaia and Handroanthus guayacan.

The lack of genomic resources for tropical canopy trees is impeding several research avenues in tropical forest biology. We present genome assemblies for two Neotropical hardwood species, Jacaranda copaia and Handroanthus (formerly Tabebuia) guayacan, that are model systems for research on tropical tree demography and flowering phenology. For each species, we combined Illumina short-read data with in vitro proximity-ligation (Chicago) libraries to generate an assembly. For Jacaranda copaia, we obtained 104X physical coverage and produced an assembly with N50/N90 scaffold lengths of 1.020/0.277 Mbp. For H. guayacan, we obtained 129X coverage and produced an assembly with N50/N90 scaffold lengths of 0.795/0.165 Mbp. J. copaia and H. guayacan assemblies contained 95.8% and 87.9% of benchmarking orthologs, although they constituted only 77.1% and 66.7% of the estimated genome sizes of 799 and 512 Mbp, respectively. These differences were potentially due to high repetitive sequence content (>59.31% and 45.59%) and high heterozygosity (0.5% and 0.8%) in each species. Finally, we compared each new assembly to a previously sequenced genome for Handroanthus impetiginosus using whole-genome alignment. This analysis indicated extensive gene duplication in H. impetiginosus since its divergence from H. guayacan.

Permafrost thaw in boreal peatlands is rapidly altering forest community composition

Abstract Boreal peatlands are frequently underlain by permafrost, which is thawing rapidly. A common ecological response to thaw is the conversion of raised forested plateaus to treeless wetlands, but unexplained spatial variation in responses, combined with a lack of stand‐level data, make it difficult to predict future trajectories of boreal forest composition and structure. We sought to characterize patterns and identify drivers of forest structure, composition, mortality and recruitment in a boreal peatland experiencing permafrost thaw. To do this, we established a large (10 ha) permanent forest plot (completed in 2014), located in the Northwest Territories, Canada, that includes 40,584 mapped and measured trees. In 2018, we conducted a comprehensive mortality and recruitment recensus. We also measured frost table depth, soil moisture, soil humification and organic layer thickness within the plot between 2012 and 2018, and used habitat association tests to link these variables to forest characteristics and dynamics. Forest composition and structure varied markedly throughout the plot and were strongly governed by patterns in permafrost presence and organic layer thickness. Overall, there was a net loss of trees from the plot at a rate of 0.7% year−1. Mortality of black spruce, the dominant tree species, was more than double that of recruitment and was strongly associated with permafrost thaw. In contrast, recruitment of larch was over four times greater than mortality, and occurred primarily in low‐lying, permafrost‐free wetlands with mineral soil near the surface. Synthesis. The trends in tree demography and underlying drivers suggest that spruce‐dominated permafrost plateaus will be converted into larch‐dominated wetlands as permafrost thaw progresses in boreal peatlands, particularly in areas where mineral soil is near the surface. In the longer term, thaw could increase the hydrologic connectivity of the landscape, resulting in widespread drainage and re‐vegetation by spruce, but we did not find evidence that this is occurring yet. Given the increasing rates of permafrost thaw, and positive feedbacks between thaw and forest change, we predict that larch abundance will continue to increase in boreal peatlands over the coming decades, leading to shifts in ecosystem function, wildlife habitat, albedo and snow dynamics.

Climate‐induced variation in the demography of 14 tree species is not sufficient to explain their distribution in eastern North America

AbstractAimDynamic range models are proposed to investigate species distributions and to project range shifts under climate change. They are based upon the Hutchinsonian niche theory, specifying that the occurrence of a species in an environmental space should be limited to positions where the intrinsic growth rate is positive. Evaluating population growth rate is, however, difficult for physiologically structured populations, such as forest stands, owing to size‐induced individual variation in performance. Therefore, we still have a limited understanding of which aspect of tree demography contributes the most to their geographical range limit. We develop an index of demographic performance for size‐structured populations and study its variation across a climatic gradient. We then investigate the relationship between the demographic performance index and species distribution.LocationNorth America (57–124° W, 26–52° N).Time period1963–2010.Major taxa studiedFourteen tree species.MethodsWe represent forest dynamics with a size‐structured population model and neighbourhood competition with the perfect plasticity approximation. We then derive the lifetime reproduction per individual, , in the absence of density dependence. Using forest inventory data, we assess how tree demography for each species varies with climate. We test the model by comparing and the probability of occurrence within species ranges.ResultsWe find that both growth and mortality rates vary across species distributions, but climate explains little of the observed variation. Individual size and neighbourhood competition are the primary explanatory variables of tree demography. Finally, we find that relates weakly to the probability of occurrence, with no systematic decline in population growth rates towards the range limits.Main conclusionsSpatial and size‐induced variation in tree growth and mortality do not explain range limits and are insufficient to enable an understanding of tree dynamics. We propose that phenomena perceived mostly at the metapopulation scale should also be considered.

Nitrogen‐fixing trees have no net effect on forest growth in the coterminous United States

Abstract Nitrogen (N)‐fixing trees fulfil a unique and important biogeochemical role in forests through their ability to convert atmospheric N2 gas to plant‐available N. Due to their high N fixation rates, it is often assumed that N‐fixing trees facilitate neighbouring trees and enhance forest growth. This assumption is supported by some local studies but contradicted by others, leaving the overall effect of N‐fixing trees on forest growth unresolved. Here we use the US Forest Service's Forest Inventory and Analysis database to evaluate the effects of N‐fixing trees on plot‐scale basal area change and individual‐scale neighbouring tree demography across the coterminous US. First we discuss the average trends. At the plot and individual scales, N‐fixing trees do not affect the relative growth rates of neighbouring trees, but they facilitate recruitment and inhibit survival rates, suggesting that they are drivers of tree turnover in the coterminous US. At the plot scale, N‐fixing trees facilitate the basal area change of non‐fixing neighbours. In addition to the average trends, there is wide variation in the effect of N‐fixing trees on forest growth, ranging from strong facilitation to strong inhibition. This variation does not show a clear geographical pattern, though it does vary with certain local factors. N‐fixing trees facilitate forest growth when they are likely to be less competitive: under high N deposition and high soil moisture or when neighbouring trees occupy different niches (e.g. high foliar C:N trees and non‐fixing trees). Synthesis. N‐fixing trees have highly variable effects on forest growth and neighbour demographics across the coterminous US. This suggests that the effect of N‐fixing trees on forest development and carbon storage depends on local factors, which may help reconcile the conflicting results found in previous localized studies on the effect of N‐fixing trees on forest growth.

Competition and demography rather than dispersal limitation slow down upward shifts of trees’ upper elevation limits in the Alps

Abstract Species range limits are expected to be dramatically altered under future climate change and many species are predicted to shift their distribution upslope to track their suitable conditions (i.e. based on their niche). However, there might be large discrepancies between the speed of the upward shift of the climatic niche and the actual migration velocity of the species, especially in long‐lived organisms such as trees. In fact, most studies did not find any significant upward shift of the distributional limits of temperate forest trees over the last decades. It therefore beckons the questions why trees are moving upslope much slower than their bioclimatic envelope and what are the implications for ecosystem functioning. Here, we compared the simulations of the upslope displacement of the bioclimatic envelope of 16 tree species inhabiting temperate mountain forests under ongoing and future climate change obtained by correlative species distribution models (SDMs) to those from a dynamic forest model accounting for dispersal, competition and demography. We then partitioned the discrepancy in upslope migration velocity between the SDMs and the dynamic forest model into different components by manipulating dispersal limitation, interspecific competition and demography. Tree species in the dynamic forest model migrated only slowly upslope in contrast to the SDMs. Most of the difference in migration velocity can directly be attributed to tree's demography (long life cycle), followed by effects of competition and only a marginal contribution of dispersal limitation. Additionally, lower elevation species (‘non‐treeline’) shifted slower upslope than high elevation species (‘treeline’) indicating a strong effect of interspecific competition at their leading edge. Synthesis. Forests have a high inertia to climate change because of their longevity and ability to acclimatize to high climatic fluctuations. Lower elevation tree species (deciduous) only slowly establish in stands at higher elevation where coniferous species dominate and likely profit from facilitation by disturbance events. Therefore, forest ecosystems seem to persist, even if climate becomes unfavourable, until they approach a tipping point at which an extreme event (e.g. drought, storm or insect attack) leads to a large dieback and resource change enabling new suitable species to spread and establish.

Tree Encroachment Varies by Plant Community in a Large Boreal Peatland Complex in the Boreal-Temperate Ecotone of Northeastern USA

Large peatland complexes at the boreal-temperate ecotone are essential habitats for boreal species at their southern range limits where they are threatened by tree encroachment accelerated by climate change and nitrogen deposition. To inform vascular plant and biodiversity conservation, we studied tree encroachment patterns in a large (> 400 ha) boreal peatland complex in the northeastern United States across vegetation types and environmental gradients. We characterized vascular plant composition, environmental drivers and tree demography on 50 plots (each 25 m2). We used non-metric multidimensional scaling (NMS) to identify two main drivers of vascular plant composition in the herbaceous layer—pH and tree canopy openness—that described three broad plant community types (open bog, forested bog, and fen). Tree demography suggested that woody encroachment (i.e., tree seedling recruitment) varied across these community types; open bog was colonized by Picea mariana seedlings, while forested bog and fen (dominated by evergreen conifers, Picea mariana and Thuja occidentalis, respectively) were colonized by deciduous tree species (Acer rubrum and Betula alleghaniensis). Our findings provide early warning signs of vegetation change in boreal peatlands near their southern range limits caused by the encroachment of temperate tree species into forested peatlands and expanding tree cover in open bogs.

Drought mitigation by thinning: Benefits from the stem to the stand along 15 years of experimental rainfall exclusion in a holm oak coppice

In northern Mediterranean forests, increasing drought stress due to the on-going climate change is combined with stand ageing due to the lack of management. Management by thinning may alleviate drought stress by reducing competition, but its application is challenging in coppices of resprouting species where its long-term consequences for tree demography and stand dynamics are difficult to evaluate. In this study, we investigate the long-term (15 years) demographic responses of holm oak (Quercus ilex L.) to a combination of thinning from below (−30% basal area) and experimental rainfall exclusion (−27% precipitation). Stem growth, survival and resistance to an extreme drought event were positively linked to both stem size and local competition release after thinning. Thinning improvement of growth and survival were thus due to both a selection of the biggest, most vigorous, trees and to a release of competition for water. Rainfall exclusion, on the other hand, led to a shift of the tree size-mortality relationship, which resulted in the death of bigger trees, in a faster loss of stool density and in a slower evolution of the stand basal area compared to the control. Thinning was beneficial by cancelling the rainfall exclusion effects on growth and mortality, and by doubling the stand basal area increment compared to unthinned control. The initial loss of stools due to thinning was compensated by a lower mortality, suggesting that thinning do not reduce further the amount of unique genotypes on the long-term. Positive thinning effects on stem growth decreased over time but remained significant 15 years after thinning, while resprouting dynamics strongly decreased with time. These results indicate that moderate thinning from below is a relevant strategy to increase stem vitality and stand production in old coppices, particularly in a context of a chronic rise in drought stress and more frequent extreme drought episodes.

Soil-borne pathogens as determinants of regeneration patterns at community level in Mediterranean forests.

Emergent diseases are an increasing problem in forests worldwide. Exotic pathogens are now threatening forests where pathogens have not traditionally been considered to be major ecological drivers of tree demography, such as water-limited Mediterranean forests. However, how pathogens might limit regeneration in invaded forests is largely unknown. Here we used fungicide to analyse the impact of soil-borne oomycete pathogens on seedling establishment at community level in Mediterranean forests invaded by the exotic oomycete Phytophthora cinnamomi. Fungicide effects were modelled as a function of the tree neighbourhood composition, the seed mass of the target species, and the abiotic environment. Fungicide application had positive effects on seedling performance that varied in magnitude and spatial structure among coexisting species. Seed mass predicted fungicide effects on seedling emergence, but not on survival or growth. Positive fungicide effects were modulated by levels of abiotic resources, mainly water, increasing with soil moisture. Our results support a novel role for soil-borne oomycete pathogens as one more axis of the regeneration niche of woody species in water-limited forests. Given the increasing numbers of exotic oomycete pathogens worldwide, more research is needed to understand the role of this relevant microbial group as a factor shaping seedling establishment.

Simulating functional diversity of European natural forests along climatic gradients

AbstractAimWe analyse how functional diversity (FD) varies across European natural forests to understand the effects of environmental and competitive filtering on plant trait distribution.LocationForest ecosystems in Europe from 11°W to 36°E and 29.5°N to 62°N.TaxonPinaceae, Fagaceae and Betulaceae, Oleaceae, Tiliaceae, Aceraceae, Leguminosae (unspecific).MethodsWe adopted the existing Dynamic Global Vegetation Model Lund‐Potsdam‐Jena managed Land of flexible individual traits (LPJmL‐FIT) for Europe by eliminating both bioclimatic limits of plant functional types (PFTs) and replacing prescribed values of functional traits for PFTs with emergent values under influence of environmental filtering and competition. We quantified functional richness (FR), functional divergence (FDv) and functional evenness (FE) in representative selected sites and at Pan‐European scale resulting from simulated functional and structural trait combinations of individual trees. While FR quantifies the amount of occupied trait space, FDv and FE describe the distribution and abundance of trait combinations, respectively, in a multidimensional trait space.ResultsLund‐Potsdam‐Jena managed Land of flexible individual traits reproduces spatial PFTs and local trait distributions and agrees well with observed productivity, biomass and tree height of European natural forests. The observed site‐specific trait distributions and spatial gradients of traits of the leaf‐ and stem‐resource economics spectra coincide with environmental filtering and the competition for light and water in environments with strong abiotic stress. Where deciduous and needle‐leaved trees co‐occur, for example, in boreal and mountainous forests, the potential niche space is wide (high FR), and extreme ends in the niche space are occupied (high FDv). We find high FDv in Mediterranean forests where drought increasingly limits tree growth, thus niche differentiation becomes more important. FDv decreases in temperate forests where a cold climate increasingly limits growth efficiency of broad‐leaved summer green trees, thus reducing the importance of competitive exclusion. Highest FE was simulated in wet Atlantic and southern Europe which indicated relatively even niche occupation and thus high resource‐use efficiency.Main ConclusionsWe find FD resulting from both environmental and competitive filtering. Pan‐European FR, FDv and FE demonstrate the influence of climate gradients and intra‐ and inter‐PFT competition. The indices underline a generally high FD of natural forests in Europe. Co‐existence of functionally diverse trees across PFTs emerges from alternative (life‐history) strategies, disturbance and tree demography.

Late Miocene origin and recent population collapse of the Malagasy savanna olive tree (Noronhia lowryi)

Abstract Debates regarding the origin of tropical savannas have attempted to disentangle the role of human, biotic and abiotic factors. Understanding the origins of savanna remains essential to identifying processes that gave rise to habitat mosaics, particularly those found in the Central Plateau of Madagascar. Documenting the evolutionary history and demography of native trees occurring in open habitats may reveal footprints left by past and recent environmental changes. We conducted a population genetic analysis of an endangered Malagasy shrub (Noronhia lowryi, Oleaceae) of the Central Plateau. Seventy-seven individuals were sampled from three sites and genotyped at 14 nuclear and 24 chloroplast microsatellites. We found a highly contrasting nuclear and plastid genetic structure, suggesting that pollen-mediated gene flow allows panmixia, while seed-based dispersal may rarely exceed tens of metres. From a phylogeny based on full plastomes, we dated the surprisingly old crown age of maternal lineages back to ~6.2 Mya, perhaps co-occurring with the global expansion of savanna. In contrast, recent demographic history inferred from nuclear data shows a bottleneck signature ~350 generations ago, probably reflecting an environmental shift during the Late Pleistocene or the Holocene. Ancient in situ adaptation and recent demographic collapse of an endangered woody plant highlight the unique value and vulnerability of the Malagasy savannas.

Long-term forest dynamics in response to climate change in northern mixed forests in Japan: A 38-year individual-based approach

Climate change is expected to cause a directional change in northern mixed forests from sub-boreal conifer-dominated to cool-temperate broad-leaved trees dominated forests due to species-specific responses to the changing climate regime. However, few studies have examined both the long-term effects of climate (temperature, precipitation and wind disturbance) and non-climate (stand structure and topography) factors.In this study, we analyzed forest census data from 15 forest plots (17.5 ha, 8460 trees) in two primeval reserve areas over a 38-year period (1979–2016) using individual-based or grid-based state-space models to evaluate the effects of climate and non-climate factors that affect the demographic parameters (growth rate, mortality, and recruitment rate) of the three coniferous and seven deciduous broad-leaved species.Two climate factors during the growth period differentially affected the growth rate of trees: high temperature and high precipitation had negative effects on coniferous species and positive effects on broad-leaved species, respectively. The change of stand basal area and its interaction between precipitation in non-growth period affected the growth rate of some species. Furthermore, the mortality of large conifers was strongly affected by a typhoon in 2004. The dominance of coniferous species has declined especially in plots with west-facing slopes and from snowfall that has largely diminished since 1979.In northern Japan, the temperature and precipitation of the growth period are increasing, and snowfall is decreasing. Our results indicated that this climate change reduced the growth rates of coniferous species and increased those of broad-leaved species. Moreover, wind disturbances will increase, which will accelerate the change in species composition by removing dominant coniferous trees. This study is the first to show these directional changes in the species composition of northern mixed forests from conifer-dominated to broad-leaved tree-dominated based on the long-term tree demography.