Quantifying variation in forest disturbance,and its effects on aboveground biomass dynamics,across the eastern United States

The role of tree mortality in the global carbon balance is complicated by strong spatial and temporal heterogeneity that arises from the stochastic nature of carbon loss through disturbance. Characterizing spatio‐temporal variation in mortality (including disturbance) and its effects on forest and carbon dynamics is thus essential to understanding the current global forest carbon sink, and to predicting how it will change in future. We analyzed forest inventory data from the eastern United States to estimate plot‐level variation in mortality (relative to a long‐term background rate for individual trees) for nine distinct forest regions. Disturbances that produced at least a fourfold increase in tree mortality over an approximately 5 year interval were observed in 1–5% of plots in each forest region. The frequency of disturbance was lowest in the northeast, and increased southwards along the Atlantic and Gulf coasts as fire and hurricane disturbances became progressively more common. Across the central and northern parts of the region, natural disturbances appeared to reflect a diffuse combination of wind, insects, disease, and ice storms. By linking estimated covariation in tree growth and mortality over time with a data‐constrained forest dynamics model, we simulated the implications of stochastic variation in mortality for long‐term aboveground biomass changes across the eastern United States. A geographic gradient in disturbance frequency induced notable differences in biomass dynamics between the least‐ and most‐disturbed regions, with variation in mortality causing the latter to undergo considerably stronger fluctuations in aboveground stand biomass over time. Moreover, regional simulations showed that a given long‐term increase in mean mortality rates would support greater aboveground biomass when expressed through disturbance effects compared with background mortality, particularly for early‐successional species. The effects of increased tree mortality on carbon stocks and forest composition may thus depend partly on whether future mortality increases are chronic or episodic in nature.     

Fragmentation and pre-existing species turnover determine land-snail assemblages of tropical rain forest

Aim The main aims of the study were: (1) to investigate the effect of fragment age in relation to other patch‐ and landscape‐scale measures of forest fragmentation, and (2) to assess the relative importance of fragmentation, habitat degradation (i.e. degradation caused by selective logging and past shifting cultivation) and putative pre‐existing species turnover in structuring current land‐snail assemblages. Location South‐western Sri Lanka. Methods The land‐snail fauna was sampled using standardized belt transects. Fifty‐seven transects were sampled in 21 lowland rain forest fragments (c. 1–33,000 ha). The spatial arrangement of fragments in the study area was explicitly considered in an effort to take into account the non‐random nature of fragmentation and degradation and the possibility that current species composition may reflect patterns of species turnover that existed prior to fragmentation. The data set of 57 land‐snail species and 28 environmental and spatial variables was analysed using canonical correspondence analysis and partial canonical correspondence analysis. Results Fragment age, mean shape complexity (i.e. a landscape‐scale measure of shape complexity), altitude, and the spatial variables x (longitude), y (latitude) and y² explained significant variation in land‐snail species composition. None of the three nominal variables quantifying habitat degradation was significantly correlated with variation in species composition. The independent effects of fragment age and mean shape complexity were similar. The combined effect of the spatial variables alone was larger than the independent effects of fragment age, mean shape complexity or altitude, but was of the same order of magnitude. The total variation explained by the spatial variables was comparable to the total non‐spatial variation accounted for by fragment age, mean shape complexity and altitude. Main conclusions Fragment age was found to be one of only two key determinants (the other was shape complexity at the landscape scale) driving fragmentation‐related changes in community composition. The influence of pre‐fragmentation patterns of species turnover on assemblage structure can be stronger than the effects of fragmentation measures, such as age, and may override the effects of forest degradation. Thus, strong patterns of pre‐existing turnover may potentially confound interpretation of the effects of forest fragmentation and degradation.     

Survival and growth of seedlings of 12 Chilean rainforest trees in two light environments: Gas exchange and biomass distribution correlates

Abstract Mortality, growth, gas exchange and biomass distribution were measured in the seedlings of 12 Chilean temperate rainforest angiosperm trees in two contrasting artificial light environments (150 and 12 μmoles m^?2 s^?1), in order to explore life history diversity in this forest type, and examine the physiological and/or morphological traits associated with interspecific variation in seedling performance. Gas exchange traits were measured only in the high‐light treatment (150 μmoles m^?2 s^?1), owing to the very small size of leaves in the low‐light treatment. Relative growth rates (RGR) in high light were strongly correlated with photosynthetic capacity (P< 0.0001). Mortality rates in low light had a strong positive correlation with light compensation point (P = 0.007) and photosynthetic capacity (P = 0.004). Furthermore, high‐light RGR was strongly positively correlated with low‐light mortality (P = 0.001). Biomass distribution traits showed little relationship with mortality or growth within either light level, except for a marginally significant positive correlation (P = 0.04) between leaf area ratio and mortality in low light. In view of the large interspecific differences in final size, the weak relationships between biomass distribution parameters and plant performance could be partially attributable to ontogenetic drift in these traits. Among taxa with high mortality rates in low light, short‐lived species (e.g. Aristotelia chilensis) had lower light compensation points, had greater phenotypic plasticity and grew much faster in both light environments than did longer‐lived species (Nothofagus dombeyi, Weinmannia trichosperma, Eucryphia cordifolia). Results support the view that survival of first‐year seedlings in low light is not enhanced by morphological traits that maximize growth potential (e.g. high leaf area ratio), and that leaf‐level gas exchange traits have an important role as determinants of interspecific variation in seedling performance. However, the limited range of interspecific variation observed in light compensation points indicates that other traits apart from those that we measured (e.g. carbon storage) must also be involved in seedling shade tolerance differences. The weak relationship between longevity and shade tolerance level among our 12 species suggests that it may not be feasible to ordinate life histories of Chilean temperate rainforest trees on a single axis of trait variation.     

Fine‐scale forest variability and biodiversity in the boreal mixedwood forest

Local spatial variation in species distributions is driven by a mix of abiotic and biotic factors, and understanding such hierarchical variation is important for conservation of biodiversity across larger scales. We sought to understand how variation in species composition of understory vascular plants, spiders, and carabid beetles is associated with concomitant spatial variation in forest structure on a 1‐ha permanent plot in a never‐cut mixedwood forest in central Alberta (Canada). Using correlations among dendrograms produced by cluster analysis we associated data about mapped distribution of all living and dead stems > 1 cm diameter at breast height with distributions of the three focal taxa sampled from regular grids across the plot. Variation in each of these species assemblages were significantly associated with several forest structure variables at various spatial scales, but the scale of the associations varied among assemblages. Variation in species richness and abundance was explained mostly by changes in basal area of trees across the plot; however, other variables (e.g. snag density and tree density) were also important, depending on assemblage. We conclude that fine‐scale habitat variation is important in structuring spatial distribution of the species of the forest floor, even within a relatively homogeneous natural forest. Thus, assessments that ignore within‐stand heterogeneity and management that ignores its maintenance will have limited utility as conservation measures for these taxa, which are major elements of forest biodiversity.     

Disentangling environmental and spatial processes of community assembly in tropical forests from local to regional scales

Understanding patterns and mechanisms of variation in the compositional structure of communities across spatial scales is one of the fundamental challenges in ecology and biogeography. In this study, we evaluated the effects of spatial extent (i.e. size of study region) on: 1) whether community composition can be better explained by environmental (i.e. niche‐based) or spatial (e.g. dispersal‐based) processes ; and 2) how climate and soils contribute to the influence of environment on plant community composition. We surveyed community composition across a network of 398 forest plots spanning a ～4000 m elevational gradient in the Madidi region in northwestern Bolivia. Using redundancy analyses and hierarchical variation partitioning, we disentangled the effects of environmental and spatial predictors on species composition, further decomposing the environmental effect between its climatic and soil components. We repeated analyses for 200 sub‐regions ranging in spatial extent from ～250 to ～17 500 km². Our analyses show a high degree of idiosyncrasy in results that come from different sub‐regions. Despite this variability, we were able to identify various important patterns in the structure of tropical plant communities in our study system. First, even though sub‐regions varied in size by nearly two orders of magnitude, the total amount of explained variation in community composition was scale independent; at all spatial scales, environment and space accounted for about 25% of the differences in community composition among plots. Second, the measured environmental effect was higher than the spatial effect on average and in the vast majority of sub‐regions. This was true regardless of the spatial extent of analysis. Finally, we found that both climatic and soil variables accounted for significant fractions of variation, but climate was always more important than soils.     

Non‐trophic plant–animal interactions mediate positive density dependence among conspecific saplings

Trophic plant–animal interactions (e.g. browsing by ungulates, insect attack) are an important and well‐studied source of mortality in many tree populations. Non‐trophic tree–animal interactions (e.g. deer antler rubbing) also frequently lead to tree death, and thus have significant effects on forest ecosystem functioning, but they are much less well studied than trophic interactions are. As deer populations have increased in recent decades in the Northern Hemisphere, their impact on tree populations via browsing and antler rubbing will increase. The aim of the study was to illustrate the potential ability of non‐trophic plant–animal interactions to regulate the dynamics of a natural forest. Specifically, we wanted to determine whether and how density and distance‐dependent processes affect sapling mortality caused by an antler rubbing by red deer Cervus elaphus. We used a spatially explicit approach to examine density and distance‐dependent mortality effects in almost two thousand Picea abies saplings over 20 years, based on a fully mapped permanent 14.4 ha plot in a natural subalpine old‐growth spruce forest. Antler rubbing by deer was the main identified cause of sapling mortality, and it showed a strong spatial pattern: positive density dependence of survival among spruce saplings. Deer selectively killed spruce saplings that were isolated from conspecifics. In consequence, non‐trophic plant–deer interactions were a major driver of the spatial pattern of P. abies sapling survival. The other mortality causes (e.g. breaking, overturning) did not show density‐dependent patterns or their effects were much weaker. In the medium and long term, the density‐dependent pattern of sapling mortality due to antler rubbing can alter the tree stand structure. Our results highlight the ecological relevance of non‐trophic plant–animal interactions for forest ecosystem functioning.     

Can spatial variation in epiphyte diversity and community structure be predicted from sampling vascular epiphytes alone?

Aim Non‐vascular epiphytes have been largely ignored in studies examining the biotic and abiotic determinants of spatial variation in epiphyte diversity. Our aim was to test whether the spatial patterning of species richness, biomass and community composition across geographic regions, among trees within regions, and among branches within trees is consistent between the vascular and non‐vascular components of the temperate rain forest flora. Location Coastal lowland podocarp‐broadleaved forests on the west coast of the South Island of New Zealand. Methods We collected single samples (30 × 25 cm) from 96 epiphyte assemblages located on the inner branches of 40 northern rata (Metrosideros robusta) trees. For each sample, branch characteristics such as branch height, branch diameter, branch angle, branch aspect, and minimum and maximum epiphyte mat depth were recorded. The biomass for each individual epiphyte species was determined. Results Northern rata was host to a total of 157 species, comprising 32 vascular and 125 non‐vascular species, with liverworts representing 41% of all species. Within epiphyte mats, the average total organic biomass of 3.5 kg m^?2 of branch surface area consisted largely of non‐living biomass and roots. Vascular and non‐vascular epiphytes showed strikingly different spatial patterns in species richness, biomass and composition between sites, among trees within sites, and among branches within trees, which could not be explained by the branch structural characteristics we measured. The two plant groups had no significant association in community composition (r = 0.04, P = 0.08). However, the species richness of vascular plant seedlings was strongly linked to the presence/absence of lichens. Main conclusions Non‐vascular plants contributed substantially to the high species richness and biomass recorded in this study, which was comparable to that of some tropical rain forests. High variability in community composition among epiphyte mats, and very low correlation with any of the environmental factors measured possibly indicate high levels of stochasticity in seed or spore colonization, establishment success or community assembly among branches in these canopy communities. Although we found some evidence that vascular plant seedling establishment was linked to the presence of lichens and the biomass of non‐living components in the epiphyte mats, there was no correlation in the spatial patterning or determinants of species richness between non‐vascular and vascular plants. Consequently, variation in total epiphyte biodiversity could not be predicted from the measurement of vascular plant diversity alone, which highlights the crucial importance of sampling non‐vascular plants when undertaking epiphyte community studies.     

Defaunation impacts on seed survival and its effect on the biomass of future tropical forests

Large animal species, which provide important ecological functions such as dispersal of seeds or top–down control of seed predators, are very vulnerable in fragmented forests, being unable to survive in small fragments, and facing increasing hunting pressure. The loss of large animals affects two main ecological processes crucial for the tree reproductive cycle: seed dispersal of large seeds (e.g. provided by tapirs) and control of seed predator population (e.g. provided by large cats). The changes in both processes are expected to increase seed mortality since seeds are not dispersed away from conspecifics (causing increased pre‐dispersal mortality due to negative density dependent effects) and/or face increased predation after a dispersal event (post‐dispersal mortality). Although an extensive body of empirical knowledge exists on seed predation, the link between seed loss and adult tree community composition and structure is not well established, as well as the temporal scale seed changes affect adults. Using an individual‐based forest model (FORMIND), we evaluate the long‐term consequences of increased pre and post‐dispersal seed mortality on the future forest biomass retention of a Brazilian northeastern Atlantic forest. Our results show that forest biomass is significantly affected after 80–93% pre‐dispersal loss of large seeds, or post‐dispersal predation densities of 20–25 predators per parent tree. Large‐seeded tree species are at increased risk of local extinction causing up to 26.2% loss of forest biomass when both pre and post‐dispersal processes are combined. However, these changes can last up to 100 years after the occurrence of defaunation. In summary we conclude that large animal loss has the potential to reduce future forest biomass and tree species‐richness by impacting seed survival, and should be considered in the planning of biodiversity friendly landscapes as well as in calculations of the global carbon budget.     

Changes in miombo woodland structure under different land tenure and use systems in central Zambia

Aim Population pressure and communal land ownership are often perceived as serious threats to forest conservation in savanna woodlands of central and southern Africa. I aimed at testing the hypothesis that the rate of miombo woodland recovery after clearing and re‐growth structure are determined by land tenure and use. Location Miombo woodland under customary, leasehold, forest reserve and national park on ten permanent and temporary sites was studied in central Zambia. Two sites were in mature woodland and eight sites were in re‐growth miombo ranging in age from 1 to 30 years. Methods I enumerated and measured girth at breast height (1.3 m above ground) of trees/stems in sixty‐four 20 × 10 m plots in 1982, 1986 and 2000 at six sites and annually from 1990 to 2001 at four sites to determine stem density and status (live, dead or cut) and wood biomass. A total of 239 trees were cut, wood biomass measured and the data used to develop equations for estimating wood biomass on study plots. Distance between each study site and the nearest human settlement was estimated during each sampling period using aerial photographs, topographical maps and the global positioning system. Results Land tenure was responsible for significant differences in stem density, wood biomass and rate of biomass accumulation in re‐growth following clearing of mature miombo woodland. Although stem density was highest on customary land, wood biomass and accumulation rate were lowest. The highest biomass was on plots in forest reserves, with intermediate values for leasehold and national park. Fire was responsible for tree mortality at all the study sites and its impact was highest at a site in a national park. Sites close to human settlements had the highest density of cut stems but this activity did not significantly reduce wood biomass. Rate of woodland recovery was higher on sites cleared in the 1970s than on sites cleared in the 1990s, irrespective of age of re‐growth. The development of the first, second and third re‐growths following successive woodland clearing in 1972, 1981 and 1990, respectively, was not significantly different, except for stem density which was highest in the second re‐growth. Analysis of interactions between five land tenure and use factors (independent variables) and re‐growth structure revealed that 52% (P=0.0000) of the variation in stem density was because of re‐growth age and decade in which the woodland was cleared while distance to human settlements and age of re‐growth explained 42% (P=0.0000) of the variation in wood biomass. Individually, distance to human settlements explained 25% (P=0.0000) of the variation in wood biomass accumulation rate. Conclusion The results supported the hypothesis that rate of miombo woodland recovery and structure were influenced by land tenure and use. However, analysis of interactions between factors revealed that use related factors (i.e. decade in which woodland was cleared and distance to human settlements) and re‐growth related factors (age and type of re‐growth) were more important than land tenure per se in explaining variation in miombo recovery. The conclusion from these results is that regulation of land use is more important than change in land tenure to the proper management of miombo woodland.     

Role of environmental and spatial processes in structuring anuran communities across a tropical rain forest

This study aimed to evaluate if anuran species distributions in riparian and non‐riparian areas are influenced by environmental factors (i.e. niche) and/or by spatial factors (i.e. dispersal). The environmental variables analysed were altitude, distance from the stream and leaf litter depth. Spatial factors were represented by the eigenvectors extracted from geographical coordinates by eigenfunction analysis. The study was conducted in 24 km² of terra‐firme forest in Central Amazonia, Manaus – Amazonas, Brazil. Between November 2008 and May 2009, three samples were taken from 41 plots, 21 plots being placed at non‐riparian areas and another 20 placed in riparian areas. We submitted the assemblage dataset to a partial redundancy analysis to evaluate the contributions of environmental and spatial variables (selected with a forward selection procedure). In addition, we tested if communities differ from riparian and non‐riparian areas using a db‐MANOVA. Species richness and species composition differed between riparian and non‐riparian plots. Some species were restricted to riparian areas. Altitude was the only significant variable (P = 0.005) explaining 21% of the total variance. When analysing the data from all plots using the partial redundancy analysis, 27% of the variance was explained by spatial and environmental variables. The environmental variables explained exclusively 4% of the variance in assemblage composition, and 13% was explained by environmental variables that were also structured in space (i.e. the shared fraction), while 10% was explained exclusively by spatial variables. In conclusion, our results showed differences between the assemblages of riparian and non‐riparian areas which can be explained by the distribution of anuran species along environmental gradients altitude and distance to streams, with little evidence of dispersal limitation.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

Assembly mechanisms determining high species turnover in aquatic communities over regional and continental scales

Niche and neutral processes drive community assembly and metacommunity dynamics, but their relative importance might vary with the spatial scale. The contribution of niche processes is generally expected to increase with increasing spatial extent at a higher rate than that of neutral processes. However, the extent to what community composition is limited by dispersal (usually considered a neutral process) over increasing spatial scales might depend on the dispersal capacity of composing species. To investigate the mechanisms underlying the distribution and diversity of species known to have great powers of dispersal (hundreds of kilometres), we analysed the relative importance of niche processes and dispersal limitation in determining beta‐diversity patterns of aquatic plants and cladocerans over regional (up to 300 km) and continental (up to 3300 km) scales. Both taxonomic groups were surveyed in five different European regions and presented extremely high levels of beta‐diversity, both within and among regions. High beta‐diversity was primarily explained by species replacement (turnover) rather than differences in species richness (i.e. nestedness). Abiotic and biotic variables were the main drivers of community composition. Within some regions, small‐scale connectivity and the spatial configuration of sampled communities explained a significant, though smaller, fraction of compositional variation, particularly for aquatic plants. At continental scale (among regions), a significant fraction of compositional variation was explained by a combination of spatial effects (exclusive contribution of regions) and regionally‐structured environmental variables. Our results suggest that, although dispersal limitation might affect species composition in some regions, aquatic plant and cladoceran communities are not generally limited by dispersal at the regional scale (up to 300 km). Species sorting mediated by environmental variation might explain the high species turnover of aquatic plants and cladocerans at regional scale, while biogeographic processes enhanced by dispersal limitation among regions might determine the composition of regional biotas.     

Distinguishing the importance between habitat specialization and dispersal limitation on species turnover

Understanding what governs community assembly and the maintenance of biodiversity is a central issue in ecology, but has been a continuing debate. A key question is the relative importance of habitat specialization (niche assembly) and dispersal limitation (dispersal assembly). In the middle of the Loess Plateau, northwestern China, we examined how species turnover in Liaodong oak (Quercus wutaishanica) forests differed between observed and randomized assemblies, and how this difference was affected by habitat specialization and dispersal limitation using variation partitioning. Results showed that expected species turnover based on individual randomization was significantly lower than the observed value (P < 0.01). The turnover deviation significantly depended on the environmental and geographical distances (P < 0.05). Environmental and spatial variables significantly explained approximately 40% of the species composition variation at all the three layers (P < 0.05). However, their contributions varied among forest layers; the herb and shrub layers were dominated by environmental factors, whereas the canopy layer was dominated by spatial factors. Our results underscore the importance of synthetic models that integrate effects of both dispersal and niche assembly for understanding the community assembly. However, habitat specialization (niche assembly) may not always be the dominant process in community assembly, even under harsh environments. Community assembly may be in a trait‐dependent manner (e.g., forest layers in this study). Thus, taking more species traits into account would strengthen our confidence in the inferred assembly mechanisms.     

The suppression of Dendroctonus frontalis and subsequent wildfire have an impact on forest stand dynamics

Question: Interacting disturbance effects from Dendroctonus frontalis outbreaks and wildfire are thought to maintain Pinus spp. composition in the southeastern U. S. Our objective was to assess forest composition, structure, and succession following the interaction of two frequently occurring disturbance events in southern Pinus spp. forests: cut‐and‐leave suppression, a commonly used means for managing D. frontalis outbreaks, and wildfire. Location: Western Gulf Coastal Plain, Louisiana, USA. Method: Pinus taeda stands with cut‐and‐leave suppression and subsequent wildfire were compared to stands undisturbed by D. frontalis but with the same wildfire events twenty years after Pinus spp. mortality. The woody plant community was assessed in three different size classes and used to predict future forest types with the Forest Vegetation Simulator (50 years). Results: P. taeda is the most abundant (> 50%) species of saw‐ and poletimber‐sizes following cut‐and‐leave suppression with wildfire and in stands only with fire. Using canonical correspondence analysis, vegetation assemblages were primarily explained by slope position and elevation (7.6% variation explained). Fire intensity and stand age also accounted for variance in the ordination (4.4% and 3.1%, respectively). Dominant and co‐dominant P. taeda forest types were predicted by the model to be the most abundant forest types in each disturbance regime. In addition, new regeneration represents high hazard for future mortality from D. frontalis. Conclusion: Our study demonstrates that cut‐and‐leave suppression with additional wildfire disturbance maintains P. taeda composition, and does not alter forest composition differently from stands receiving only wildfire. As a result, predicted Pinus spp. basal area under both disturbances is great enough to facilitate future bark beetle disturbance.     

Prolonged tropical forest degradation due to compounding disturbances: Implications for CO2 and H2O fluxes

Drought, fire, and windstorms can interact to degrade tropical forests and the ecosystem services they provide, but how these forests recover after catastrophic disturbance events remains relatively unknown. Here, we analyze multi‐year measurements of vegetation dynamics and function (fluxes of CO₂ and H₂O) in forests recovering from 7 years of controlled burns, followed by wind disturbance. Located in southeast Amazonia, the experimental forest consists of three 50‐ha plots burned annually, triennially, or not at all from 2004 to 2010. During the subsequent 6‐year recovery period, postfire tree survivorship and biomass sharply declined, with aboveground C stocks decreasing by 70%–94% along forest edges (0–200 m into the forest) and 36%–40% in the forest interior. Vegetation regrowth in the forest understory triggered partial canopy closure (70%–80%) from 2010 to 2015. The composition and spatial distribution of grasses invading degraded forest evolved rapidly, likely because of the delayed mortality. Four years after the experimental fires ended (2014), the burned plots assimilated 36% less carbon than the Control, but net CO₂ exchange and evapotranspiration (ET) had fully recovered 7 years after the experimental fires ended (2017). Carbon uptake recovery occurred largely in response to increased light‐use efficiency and reduced postfire respiration, whereas increased water use associated with postfire growth of new recruits and remaining trees explained the recovery in ET. Although the effects of interacting disturbances (e.g., fires, forest fragmentation, and blowdown events) on mortality and biomass persist over many years, the rapid recovery of carbon and water fluxes can help stabilize local climate.     

Plant responses to climatic extremes: within‐species variation equals among‐species variation

Within‐species and among‐species differences in growth responses to a changing climate have been well documented, yet the relative magnitude of within‐species vs. among‐species variation has remained largely unexplored. This missing comparison impedes our ability to make general predictions of biodiversity change and to project future species distributions using models. We present a direct comparison of among‐ versus within‐species variation in response to three of the main stresses anticipated with climate change: drought, warming, and frost. Two earlier experiments had experimentally induced (i) summer drought and (ii) spring frost for four common European grass species and their ecotypes from across Europe. To supplement existing data, a third experiment was carried out, to compare variation among species from different functional groups to within‐species variation. Here, we simulated (iii) winter warming plus frost for four grasses, two nonleguminous, and two leguminous forbs, in addition to eleven European ecotypes of the widespread grass Arrhenatherum elatius. For each experiment, we measured: (i) C/N ratio and biomass, (ii) chlorophyll content and biomass, and (iii) plant greenness, root ¹⁵N uptake, and live and dead tissue mass. Using coefficients of variation (CVs) for each experiment and response parameter, a total of 156 within‐ vs. among‐species comparisons were conducted, comparing within‐species variation in each of four species with among‐species variation for each seed origin (five countries). Of the six significant differences, within‐species CVs were higher than among‐species CVs in four cases. Partitioning of variance within each treatment in two of the three experiments showed that within‐species variability (ecotypes) could explain an additional 9% of response variation after accounting for the among‐species variation. Our observation that within‐species variation was generally as high as among‐species variation emphasizes the importance of including both within‐ and among‐species variability in ecological theory (e.g., the insurance hypothesis) and for practical applications (e.g., biodiversity conservation).     

On the significance of belowground overyielding in temperate mixed forests: separating species identity and species diversity effects

Complementary soil exploration by the root systems of coexisting tree species has been hypothesised to result in a higher root biomass of mixed forests than of monocultures but the existing evidence for a belowground diversity effect in forests is scarce and not conclusive. In a species‐rich temperate broad‐leaved forest, we analysed the fine root biomass (roots ≤ 2 mm) and necromass in 100 plots differing in tree species diversity (one to three species) and species composition (all possible combinations of five species of the genera Acer, Carpinus, Fagus, Fraxinus and Tilia) which allowed us to separate possible species diversity and species identity effects on fine root biomass. We found no evidence of a positive diversity effect on standing fine root biomass and thus of overyielding in terms of root biomass. Root necromass decreased with increasing species diversity at marginal significance. Various lines of evidence indicate significant species identity effects on fine root biomass (10–20% higher fine root biomass in plots with presence of maple and beech than in plots with hornbeam; 100% higher fine root biomass in monospecific beech and ash plots than in hornbeam plots; differences significant). Ash fine roots tended to be over‐represented in the 2‐ and 3‐species mixed plots compared to monospecific ash plots pointing at apparent belowground competitive superiority of Fraxinus in this mixed forest. Our results indicate that belowground overyielding and spatial complementarity of root systems may be the exception rather than the rule in temperate mixed forests.     

Landscape host abundance and configuration regulate periodic outbreak behavior in spruce budworm Choristoneura fumiferana

Landscape‐level forest management has long been hypothesized to affect forest insect outbreak dynamics, but empirical evidence remains elusive. We hypothesized that the combination of increased hardwood relative to host tree species, prevalence of younger forests, and fragmentation of those forests due to forest harvesting legacies would reduce outbreak intensity, increase outbreak frequency, and decrease spatial synchrony in spruce budworm Choristoneura fumiferana outbreaks. We investigated these hypotheses using tree ring samples collected across 51 sites pooled into 16 subareas distributed across a large ecoregion spanning the international border between Ontario (Canada), and Minnesota (USA). This ecoregion contains contrasting land management zones with clear differences in forest landscape structure (i.e. forest composition and spatial configuration) while minimizing the confounding influence of climate. Cluster analyses of the 76‐yr time‐series generally grouped by subareas found within the same land management zone. Spatial nonparametric covariance analysis indicated that the highest and lowest degree of spatial synchrony of spruce budworm outbreaks were found within unmanaged wilderness and lands managed at fine spatial scales in Minnesota, respectively. Using multivariate analysis, we also found that forest composition, configuration, and climate together accounted for a total of 40% of the variance in outbreak chronologies, with a high level of shared variance between composition and configuration (13%) and between composition and climate (9%). At the scale of our study, climate on its own did not explain any of the spatial variation in outbreaks. Outbreaks were of higher frequency, lower intensity, and less spatially synchronized in more fragmented, younger forests with a lower proportion of host species, with opposing outbreak characteristics observed in regions characterised by older forests with more concentrated host species. Our study is the first quantitative evaluation of the long‐standing ‘silvicultural hypothesis’ of spruce budworm management specifically conducted at a spatio‐temporal scale for which it was intended.     

Climate‐ and successional‐related changes in functional composition of European forests are strongly driven by tree mortality

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late‐successional short‐statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm‐like strategies at low water availability in forests formerly dominated by broad‐leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm‐like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.