Tree regeneration responds more to shade casting by the overstorey and competition in the understorey than to abundance per se

Manipulating the overstorey is the key tool for forest managers to steer natural regeneration. Opening up the canopy does not only create favourable light conditions for tree seedling growth, but also for (competitive) understorey species. Therefore, a thorough understanding of how changes in the abundance of the overstorey and understorey influence tree regeneration is needed to attain successful regeneration. To this end, we used the regional Flemish Forest Inventories, which contain vegetation plots that were surveyed at two times and include large variation in species composition and abundance of both overstorey and understorey layers. These plots were classified into poor and rich forest types, which differ in overstorey and understorey species composition and soil fertility. For each forest type, we first investigated the effect of overstorey abundance and shade-casting ability on the understorey herbaceous vegetation cover and its competitive nature. Then, we modelled how both these strata influence the presence-absence as well as the cover of tree regeneration, using the zero-inflated beta distribution. Our results show that the understorey cover and its competitiveness mainly increase when the abundance and shade-casting ability of the overstorey is reduced. The shade-casting ability of the overstorey and competitiveness of the understorey were more important in determining tree regeneration, especially probability of presence, than the abundance of these layers per se. This was consistent for both forest types, although directions and magnitudes of the effects differed. In predictions mimicking several thinning scenarios we found that in the poor forests, reducing overstorey abundance could lead to an increase in seedling cover, whereas in rich forests, the opposite is true and seedling cover will potentially be reduced. Finally, in a single-species analysis focusing on Quercus, we found a trade-off between sufficiently reducing overstorey abundance, while at the same retaining parent trees as potential seed sources. These findings can be used to guide forest management decisions in order to attain successful forest regeneration in temperate forests

Interactive effects of past land use and recent forest management on the understorey community in temperate oak forests in South Sweden

Questions: Past agricultural land use and forest management have shaped and influenced the understorey composition in European forests for centuries. We investigated whether understorey vegetation assemblages are affected by (a) legacies from a historical infield/outland agricultural system (i.e., a system with nutrient-enriched vs nutrient-depleted areas), (b) recent management intensity (i.e., thinning/felling activities), and (c) the interaction of recent management and potential legacies. Location: Oak forests in Skane, south Sweden. Methods: We use three vegetation surveys (1983, 1993/94 and 2014) and notes on management and land-use history, available for 62 permanent 500 m(2) plots. We conducted linear mixed effect modelling to detect both main and interactive effects of past land use and recent management on understorey diversity measures and vegetation indicator values for light and fertility. We combined nonmetric multidimensional scaling with permutational multivariate analysis of variance and indicator species analysis to detect compositional differences caused by past land use and/or recent management. Results: Understorey diversity was mainly affected by management activities, but the former infield/outland agricultural system was an important determinant of understorey composition. Understorey composition of former infields reflected higher nutrient availability and lower light availability compared to former outland. Past land use and recent management had interactive effects on light-related understorey variables: for the less intensively managed plots, the outland plots contained more light-demanding species than the infield plots, while for the more intensively managed plots, the light-demanding signature of the understorey was similar for infield and outland plots. Conclusions: Different intensities of past land use as well as recent forest management influenced the composition of the forest understorey, and interactions were present. Therefore, careful consideration of both the long-term land-use history and the more recent disturbances due to forest management are necessary when making future predictions of understorey composition and diversity

Individualistic responses of forest herb traits to environmental change

Intraspecific trait variation (ITV; i.e. variability in mean and/or distribution of plant attribute values within species) can occur in response to multiple drivers. Environmental change and land-use legacies could directly alter trait values within species but could also affect them indirectly through changes in vegetation cover. Increasing variability in environmental conditions could lead to more ITV, but responses might differ among species. Disentangling these drivers on ITV is necessary to accurately predict plant community responses to global change. We planted herb communities into forest soils with and without a recent history of agriculture. Soils were collected across temperate European regions, while the 15 selected herb species had different colonizing abilities and affinities to forest habitat. These mesocosms (384) were exposed to two-level full-factorial treatments of warming, nitrogen addition and illumination. We measured plant height and specific leaf area (SLA). For the majority of species, mean plant height increased as vegetation cover increased in response to light addition, warming and agricultural legacy. The coefficient of variation (CV) for height was larger in fast-colonizing species. Mean SLA for vernal species increased with warming, while light addition generally decreased mean SLA for shade-tolerant species. Interactions between treatments were not important predictors. Environmental change treatments influenced ITV, either via increasing vegetation cover or by affecting trait values directly. Species' ITV was individualistic, i.e. species responded to different single resource and condition manipulations that benefited their growth in the short term. These individual responses could be important for altered community organization after a prolonged period

Direct and understorey-mediated indirect effects of human-induced environmental changes on litter decomposition in temperate forest

Human-induced environmental changes in temperature, light availability due to forest canopy management, nitrogen deposition, and land-use legacies can alter ecosystem processes such as litter decomposition. These influences can be both direct and indirect via altering the performance of understorey vegetation. To identify the direct and indirect effects of environmental changes on litter decomposition, we performed an experiment with standardised green and rooibos teas. The experiment was conducted in a temperate mixed deciduous forest, and treatments (temperature, light, and nitrogen) were applied to mesocosms filled with ancient and post-agricultural forest soil. Both green tea and rooibos teas were more rapidly decomposed in oligotrophic soil than in eutrophic soil. The direct effects of the treatments on litter decomposition varied among the two litter types, incubation times, and soil fertility groups. Warming and agricultural legacy had a negative direct effect on the decomposition of the green tea in the high soil fertility treatment during the early decomposition stage. In contrast, agricultural legacy had a positive direct effect on the decomposition of rooibos tea. Soil enriched with nitrogen had a negative direct effect on the decomposition of green tea in mesotrophic soil in the early decomposition stage and on rooibos tea in later stage. The indirect effects of the treatments were consistently negative, as treatments (especially the temperature and light treatments in the early decomposition stage) had a positive effect on plant cover, which negatively affected litter decomposition. Our results indicate that warming, increased nitrogen deposition, and land use legacy can directly stimulate the decomposition of labile litter on more fertile soils. Furthermore, warming and increased light had stronger positive direct effects on understorey herbaceous cover, which leads to slower decomposition rates, especially in more fertile soils. Therefore, the indirect effects of environmental changes related to the understorey layer on litter decomposition can be more important than their direct effects, thus should not be overlooked

Light and warming drive forest understorey community development in different environments

Plant community composition and functional traits respond to chronic drivers such as climate change and nitrogen (N) deposition. In contrast, pulse disturbances from ecosystem management can additionally change resources and conditions. Community responses to combined environmental changes may further depend on land‐use legacies. Disentangling the relative importance of these global change drivers is necessary to improve predictions of future plant communities. We performed a multifactor global change experiment to disentangle drivers of herbaceous plant community trajectories in a temperate deciduous forest. Communities of five species, assembled from a pool of 15 forest herb species with varying ecological strategies, were grown in 384 mesocosms on soils from ancient forest (forested at least since 1850) and postagricultural forest (forested since 1950) collected across Europe. Mesocosms were exposed to two‐level full‐factorial treatments of warming, light addition (representing changing forest management) and N enrichment. We measured plant height, specific leaf area (SLA) and species cover over the course of three growing seasons. Increasing light availability followed by warming reordered the species towards a taller herb community, with limited effects of N enrichment or the forest land‐use history. Two‐way interactions between treatments and incorporating intraspecific trait variation (ITV) did not yield additional inference on community height change. Contrastingly, community SLA differed when considering ITV along with species reordering, which highlights ITV’s importance for understanding leaf morphology responses to nutrient enrichment in dark conditions. Contrary to our expectations, we found limited evidence of land‐use legacies affecting community responses to environmental changes, perhaps because dispersal limitation was removed in the experimental design. These findings can improve predictions of community functional trait responses to global changes by acknowledging ITV, and subtle changes in light availability. Adaptive forest management to impending global change could benefit the restoration and conservation of understorey plant communities by reducing the light availability

Understorey removal effects on tree regeneration in temperate forests: a meta‐analysis

1. The unwanted development of dense understorey vegetation composed of resource‐acquisitive, tall plant species competing strongly with tree regeneration can pose formidable problems for managers attempting to regenerate temperate forests. Despite many studies on the effects of understorey removal, no comprehensive review has summarised and quantified its effects on subsequent life stages of tree regeneration in temperate forests. 2. We synthesised data from 32 experimental studies from temperate forest regions. We used meta‐analytic techniques to find general patterns in terms of the characteristics of the understorey, overstorey and characteristics of the regenerating tree species, which are most responsible for possible positive understorey removal effects on early life stages of tree regeneration, i.e. emergence, survival and growth. 3. Both seedling survival and growth increased in response to understorey removal; emergence did not show a clear pattern. Seedlings growing free from competition mainly increased their biomass growth (total and above‐ground), whereas diameter and height growth responded less. These positive effects were largest when removing denser understorey vegetation and under more open overstorey conditions. Multiple management options influenced the regeneration responses to understorey removal. For instance, growth of older, planted seedlings responded less to removal, whereas protection against large browsers increased growth responses. 4. Tree species with differing strategies responded differently to understorey removal. Growth and survival responses of early‐successional species responded more strongly to understorey removal than mid‐ or late‐successional tree species. 5. Synthesis and applications. Our study showed that understorey removal can have strong positive effects on tree regeneration across temperate forest contexts. The magnitude of these effects depended on overstorey and understorey conditions, but also on the type of tree species that is regenerated. Our results can support forest managers in their decision‐making and help assess under what conditions understorey removal will be most justified

The functional role of temperate forest understorey vegetation in a changing world

Temperate forests cover 16% of the global forest area. Within these forests, the understorey is an important biodiversity reservoir that can influence ecosystem processes and functions in multiple ways. However, we still lack a thorough understanding of the relative importance of the understorey for temperate forest functioning. As a result, understoreys are often ignored during assessments of forest functioning and changes thereof under global change. We here compiled studies that quantify the relative importance of the understorey for temperate forest functioning, focussing on litter production, nutrient cycling, evapotranspiration, tree regeneration, pollination and pathogen dynamics. We describe the mechanisms driving understorey functioning and develop a conceptual framework synthesizing possible effects of multiple global change drivers on understorey-mediated forest ecosystem functioning. Our review illustrates that the understorey's contribution to temperate forest functioning is significant but varies depending on the ecosystem function and the environmental context, and more importantly, the characteristics of the overstorey. To predict changes in understorey functioning and its relative importance for temperate forest functioning under global change, we argue that a simultaneous investigation of both overstorey and understorey functional responses to global change will be crucial. Our review shows that such studies are still very scarce, only available for a limited set of ecosystem functions and limited to quantification, providing little data to forecast functional responses to global change

Responses of competitive understorey species to spatial environmental gradients inaccurately explain temporal changes

Understorey plant communities play a key role in the functioning of forest ecosystems. Under favourable environmental conditions, competitive understorey species may develop high abundances and influence important ecosystem processes such as tree regeneration. Thus, understanding and predicting the response of competitive understorey species as a function of changing environmental conditions is important for forest managers. In the absence of sufficient temporal data to quantify actual vegetation changes, space-for-time (SFT) substitution is often used, i.e. studies that use environmental gradients across space to infer vegetation responses to environmental change over time. Here we assess the validity of such SFT approaches and analysed 36 resurvey studies from ancient forests with low levels of recent disturbances across temperate Europe to assess how six competitive understorey plant species respond to gradients of overstorey cover, soil conditions, atmospheric N deposition and climatic conditions over space and time. The combination of historical and contemporary surveys allows (i) to test if observed contemporary patterns across space are consistent at the time of the historical survey, and, crucially, (ii) to assess whether changes in abundance over time given recorded environmental change match expectations from patterns recorded along environmental gradients in space. We found consistent spatial relationships at the two periods: local variation in soil variables and overstorey cover were the best predictors of individual species’ cover while interregional variation in coarse-scale variables, i.e. N deposition and climate, was less important. However, we found that our SFT approach could not accurately explain the large variation in abundance changes over time. We thus recommend to be cautious when using SFT substitution to infer species responses to temporal changes.</p

Combining biodiversity resurveys across regions to advance global change research

More and more ecologists have started to resurvey communities sampled in earlier decades to determine long-term shifts in community composition and infer the likely drivers of the ecological changes observed. However, to assess the relative importance of and interactions among multiple drivers, joint analyses of resurvey data from many regions spanning large environmental gradients are needed. In this article, we illustrate how combining resurvey data from multiple regions can increase the likelihood of driver orthogonality within the design and show that repeatedly surveying across multiple regions provides higher representativeness and comprehensiveness, allowing us to answer more completely a broader range of questions. We provide general guidelines to aid the implementation of multiregion resurvey databases. In so doing, we aim to encourage resurvey database development across other community types and biomes to advance global environmental change research