P.W.F.M. Hommel

Microclimate moderates plant responses to macroclimate warming

Significance Around the globe, climate warming is increasing the dominance of warm-adapted species—a process described as “thermophilization.” However, thermophilization often lags behind warming of the climate itself, with some recent studies showing no response at all. Using a unique database of more than 1,400 resurveyed vegetation plots in forests across Europe and North America, we document significant thermophilization of understory vegetation. However, the response to macroclimate warming was attenuated in forests whose canopies have become denser. This microclimatic effect likely reflects cooler forest-floor temperatures via increased shading during the growing season in denser forests. Because standing stocks of trees have increased in many temperate forests in recent decades, microclimate may commonly buffer understory plant responses to macroclimate warming. Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., “thermophilization” of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that “climatic lags” may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12–67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass—e.g., for bioenergy—may open forest canopies and accelerate thermophilization of temperate forest biodiversity.

Effects of tree species composition on within-forest distribution of understorey species

Abstract Question: Do tree species, with different litter qualities, affect the within-forest distribution of forest understorey species on intermediate to base-rich soils? Since habitat loss and fragmentation have caused ancient forest species to decline, those species are the main focus of this study. Location: Three ancient forests, along a soil gradient from acidification-sensitive to base-rich, were studied: Limbrichterbosch and Savelsbos in The Netherlands and Holtkrat in Denmark. Methods: Canopy and soil surveys along transects generated data for Redundancy Analysis on tree - humus relationships. We analysed the distribution of forest plant species with Canonical Correspondence Analysis. The explanatory factors were soil characteristics (pH, organic matter, loam content and thickness of the humus layers), external crown projection, groundwater and canopy data. We further analysed the relationship between forest species and humus characteristics with Spearman correlations. Results: Tree species have a significant impact on humus characteristics through the nature of their litter. Humus characteristics significantly explain the distribution of forest understorey species. The pH of the first 25 cm mineral soil and the thickness of the F- (fermentation) layer are the primary factors affecting the distribution of ancient forest species. Conclusion: This study indicates that the species composition of the forest canopy affects the distribution of forest understorey species. Ancient forest species are more abundant and frequent underneath trees with base-rich litter. On acidification-sensitive soils these relationships were stronger than on more base-rich, loamy soils. Abbreviations: Ah = Soil horizon consisting of mineral soil with a high organic matter content; CEC = Cation exchange capacity; F = Fermenting litter (layer); Ah = Humus-rich mineral soil (layer); L = Litter (layer); RDA = Redundancy analysis.

A model‐based approach to studying changes in compositional heterogeneity

1. Non-random species loss and gain in local communities change the compositional heterogeneity between communities over time, which is traditionally quantified with dissimilarity-based approaches. Yet, dissimilarities summarize the multivariate species data into a univariate index and obscure the species-level patterns of change, which are central to understand the causes and consequences of the community changes. 2. Here, we propose a model-based approach that looks for species-level effects of time period and construct a multiple-site metric as a sum across species to test the consistency of the individual species responses. Species fall into different response types, showing how they influence the changes in community heterogeneity. 3. In a comparison with other multiple-sitemetrics, we illustrate the properties of our method and the differences and similarities with other approaches. For instance, ourmetric estimates the total variation in a community data set based on species-level contributions, not the compositional dissimilarities between particular sites. Similar to some other approaches, we can distinguish between heterogeneity derived from turnover or richness differences. 4. Our approach was applied to a set of 23 forest understorey resurvey studies spread across Europe. We show the species gains and lossesmay as well decrease or increase levels of community heterogeneity. Although species occurrences and communities have not changed in a consistent way along continental-scale environmental gradients such as climatic conditions, several species shifted in a similar way across the different data sets. 5. Testing the significance of shifts in species prevalence over time to infer corresponding changes in the compositional heterogeneity among sites provides a very intuitive tool for community resurvey studies. The main strengths of our framework are the explicit consideration of the relative roles of species gains and losses and the straightforward generalization to different sets of hypotheses related to community changes. Key-words: biodiversity, community composition, biotic homogenization, binomial deviance, dissimilarity, beta diversity,multivariate analysis,meta-analysis, forest understorey