Showing papers by "Christian Wirth published in 2014"

Designing forest biodiversity experiments : general considerations illustrated by a new large experiment in subtropical China

Abstract: Summary 1. Biodiversity–ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25� 8 9 25� 8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ‘ecoscape’ to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achiev

Long‐term study of root biomass in a biodiversity experiment reveals shifts in diversity effects over time

Abstract: Biodiversity experiments generally report a positive effect of plant biodiversity on aboveground biomass (overyielding), which typically increases with time. Various studies also found overyielding for belowground plant biomass, but this has never been measured over time. Also, potential underlying mechanisms have remained unclear. Differentiation in rooting patterns among plant species and plant functional groups has been proposed as a main driver of the observed biodiversity effect on belowground biomass, leading to more efficient belowground resource use with increasing diversity, but so far there is little evidence to support this. We analyzed standing root biomass and its distribution over the soil profile, along a 1–16 species richness gradient over eight years in the Jena Experiment in Germany, and compared belowground to aboveground overyielding. In our long-term dataset, total root biomass increased with increasing species richness but this effect was only apparent after four years. The increasingly positive relationship between species richness and root biomass, explaining 12% of overall variation and up to 28% in the last year of our study, was mainly due to decreasing root biomass at low diversity over time. Functional group composition strongly affected total standing root biomass, explaining 44% of variation, with grasses and legumes having strong overall positive and negative effects, respectively. Functional group richness or interactions between functional group presences did not strongly contribute to overyielding. We found no support for the hypothesis that vertical root differentiation increases with species richness, with functional group richness or composition. Other explanations, such as stronger negative plant–soil feedbacks in low-diverse plant communities on standing root biomass and vertical distribution should be considered.

Global relationship of wood and leaf litter decomposability: the role of functional traits within and across plant organs

Abstract: Aim Recent meta-analyses have revealed that plant traits and their phylogenetic history influence decay rates of dead wood and leaf litter, but it remains unknown if decay rates of wood and litter covary over a wide range of tree species and across ecosystems. We evaluated the relationships between species-specific wood and leaf litter decomposability, as well as between wood and leaf traits that control their respective decomposability. Location Global. Methods We compiled data on rates of wood and leaf litter decomposition for 324 and 635 tree species, respectively, and data on six functional traits for both organs. We used hierarchical Bayesian meta-analysis to estimate, for the first time, species-specific values for wood and leaf litter decomposability standardized to reference conditions (k*wood and k*leaf) across the globe. With these data, we evaluated the relationships: (1) between wood and leaf traits, (2) between each k* and the selected traits within and across organs, and (3) between wood and leaf k*. Results Across all species k*wood and k*leaf were positively correlated, phylogenetically clustered and correlated with plant functional traits within and across organs. k* of both organs was usually better described as a function of within- and cross-organ traits, than of within-organ traits alone. When analysed for angiosperms and gymnosperms separately, wood and leaf k* were no longer significantly correlated, but each k* was still significantly correlated to the functional traits. Main conclusions We demonstrate important relationships among wood and leaf litter decomposability as after-life effects of traits from the living plants. These functional traits influence the decomposability of senesced tissue which could potentially lead to alterations in the rates of biogeochemical cycling, depending on the phylogenetic structure of the species pool. These results provide crucial information for a better representation of decomposition rates in dynamic global vegetation models.

Predicting species’ range limits from functional traits for the tree flora of North America

Abstract: Using functional traits to explain species’ range limits is a promising approach in functional biogeography. It replaces the idiosyncrasy of species-specific climate ranges with a generic trait-based predictive framework. In addition, it has the potential to shed light on specific filter mechanisms creating large-scale vegetation patterns. However, its application to a continental flora, spanning large climate gradients, has been hampered by a lack of trait data. Here, we explore whether five key plant functional traits (seed mass, wood density, specific leaf area (SLA), maximum height, and longevity of a tree)—indicative of life history, mechanical, and physiological adaptations—explain the climate ranges of 250 North American tree species distributed from the boreal to the subtropics. Although the relationship between traits and the median climate across a species range is weak, quantile regressions revealed strong effects on range limits. Wood density and seed mass were strongly related to the lower but not upper temperature range limits of species. Maximum height affects the species range limits in both dry and humid climates, whereas SLA and longevity do not show clear relationships. These results allow the definition and delineation of climatic “no-go areas” for North American tree species based on key traits. As some of these key traits serve as important parameters in recent vegetation models, the implementation of trait-based climatic constraints has the potential to predict both range shifts and ecosystem consequences on a more functional basis. Moreover, for future trait-based vegetation models our results provide a benchmark for model evaluation.

Large-scale wind disturbances promote tree diversity in a Central Amazon Forest

Abstract: Canopy gaps created by wind-throw events, or blowdowns, create a complex mosaic of forest patches varying in disturbance intensity and recovery in the Central Amazon. Using field and remote sensing data, we investigated the short-term (four-year) effects of large (>2000 m(2)) blowdown gaps created during a single storm event in January 2005 near Manaus, Brazil, to study (i) how forest structure and composition vary with disturbance gradients and (ii) whether tree diversity is promoted by niche differentiation related to wind-throw events at the landscape scale. In the forest area affected by the blowdown, tree mortality ranged from 0 to 70%, and was highest on plateaus and slopes. Less impacted areas in the region affected by the blowdown had overlapping characteristics with a nearby unaffected forest in tree density (583 ± 46 trees ha(-1)) (mean ± 99% Confidence Interval) and basal area (26.7 ± 2.4 m(2) ha(-1)). Highly impacted areas had tree density and basal area as low as 120 trees ha(-1) and 14.9 m(2) ha(-1), respectively. In general, these structural measures correlated negatively with an index of tree mortality intensity derived from satellite imagery. Four years after the blowdown event, differences in size-distribution, fraction of resprouters, floristic composition and species diversity still correlated with disturbance measures such as tree mortality and gap size. Our results suggest that the gradients of wind disturbance intensity encompassed in large blowdown gaps (>2000 m(2)) promote tree diversity. Specialists for particular disturbance intensities existed along the entire gradient. The existence of species or genera taking an intermediate position between undisturbed and gap specialists led to a peak of rarefied richness and diversity at intermediate disturbance levels. A diverse set of species differing widely in requirements and recruitment strategies forms the initial post-disturbance cohort, thus lending a high resilience towards wind disturbances at the community level.

Disentangling the environmental-heterogeneity–species-diversity relationship along a gradient of human footprint

Abstract: Decades of study have attempted to define a generalized environmental-heterogeneity–biodiversity (EH–BD) relationship, with the traditional MacArthurian niche-based hypothesis remaining as the dominant reference point; i.e., increasing heterogeneity promotes biodiversity by increasing resource opportunities. However, studies have frequently reported negative or nonsignificant relationships. In a vast majority of them, environmental heterogeneity was defined along a gradient of increasing randomness, toward complete disorder. A new conceptual framework could help to reconcile the array of observed relationships. Using an extensive literature review, we test a conceptual framework proposing that the direction of environmental-heterogeneity–biodiversity relationships is contingent on the level of human footprint to which an ecosystem is subjected (the anthropocline). The results reveal that highly modified and seminatural ecosystems are characterized by a dominance of positive and negative EH–BD relationship...

Stand Structure and Recent Climate Change Constrain Stand Basal Area Change in European Forests: A Comparison Across Boreal, Temperate, and Mediterranean Biomes

Abstract: European forests have a prominent role in the global carbon cycle and an increase in carbon storage has been consistently reported during the twentieth century. Any further increase in forest carbon storage, however, could be hampered by increases in aridity and extreme climatic events. Here, we use forest inventory data to identify the relative importance of stand structure (stand basal area and mean d.b.h.), mean climate (water availability), and recent climate change (temperature and precipitation anomalies) on forest basal area change during the late twentieth century in three major European biomes. Using linear mixed-effects models we observed that stand structure, mean climate, and recent climatic change strongly interact to modulate basal area change. Although we observed a net increment in stand basal area during the late twentieth century, we found the highest basal area increments in forests with medium stand basal areas and small to medium-sized trees. Stand basal area increases correlated positively with water availability and were enhanced in warmer areas. Recent climatic warming caused an increase in stand basal area, but this increase was offset by water availability. Based on recent trends in basal area change, we conclude that the potential rate of aboveground carbon accumulation in European forests strongly depends on both stand structure and concomitant climate warming, adding weight to suggestions that European carbon stocks may saturate in the near future.

Of carrots and sticks

Abstract: Journals and funders increasingly require public archiving of the data that support publications. We argue that this mandate is necessary, but not sufficient: more incentives for data sharing are needed.

Plant diversity effects on the water balance of an experimental grassland

Abstract: In the literature, contrasting effects of plant species richness on the soil water balance are reported. Our objective was to assess the effects of plant species and functional richness and functional identity on soil water contents and water fluxes in the experimental grassland of the Jena Experiment. The Jena Experiment comprises 86 plots on which plant species richness (0, 1, 2, 4, 8, 16, and 60) and functional group composition (zero to four functional groups: legumes, grasses, tall herbs, and small herbs) were manipulated in a factorial design. We recorded meteorological data and soil water contents of the 0·0–0·3 and 0·3–0·7 m soil layers and calculated actual evapotranspiration (ETa), downward flux (DF), and capillary rise with a soil water balance model for the period 2003–2007. Missing water contents were estimated with a Bayesian hierarchical model. Species richness decreased water contents in subsoil during wet soil conditions. Presence of tall herbs increased soil water contents in topsoil during dry conditions and decreased soil water contents in subsoil during wet conditions. Presence of grasses generally decreased water contents in topsoil, particularly during dry phases; increased ETa and decreased DF from topsoil; and decreased ETa from subsoil. Presence of legumes, in contrast, decreased ETa and increased DF from topsoil and increased ETa from subsoil. Species richness probably resulted in complementary water use. Specific functional groups likely affected the water balance via specific root traits (e.g. shallow dense roots of grasses and deep taproots of tall herbs) or specific shading intensity caused by functional group effects on vegetation cover. Copyright © 2013 John Wiley & Sons, Ltd.

Maintenance of constant functional diversity during secondary succession of a subtropical forest in China

Abstract: Questions Does the importance of biotic interactions between tree species increase during secondary forest succession? Do functional trait values become increasingly divergent from early towards late successional stages and how is functional diversity affected by trait identity, species identity and species richness effects? Location Gutianshan National Nature Reserve, Zhejiang Province, southeast China. Methods Based on 26 leaf and wood traits for 120 woody species, we calculated functional diversity, using Rao's formula for quadratic entropy, trait dissimilarity, defined as half the mean trait-based distance of all species in the community, and functional evenness, defined as the degree to which functional diversity is maximized. We employed randomization techniques to disentangle the effects of trait identity, species identity and species richness on these three components of functional diversity. Results Against expectations, functional diversity did not show any successional trend because the communities compensated for a loss in trait dissimilarity by distributing the trait values more evenly among the resident species, thus increasing functional evenness. Randomization tests showed that functional diversity was not affected by trait identity, by species identity or by species richness, which indicates that functional diversity was neither determined by particular single traits or by single species with outstanding trait values. Conclusions The constant functional diversity suggests constant functionality in this subtropical forest, which might temporally maintain stable immigration conditions during the course of succession, and thus provides an explanation why these subtropical forests become more species-rich with time.

Biodiversity effects on nitrate concentrations in soil solution: a Bayesian model

Abstract: Ecosystems are faced with high rates of species loss which has consequences for their functions and services. To assess the effects of plant species diversity on the nitrogen (N) cycle, we developed a model for monthly mean nitrate (NO3-N) concentrations in soil solution in 0–30 cm mineral soil depth using plant species and functional group richness and functional composition as drivers and assessing the effects of conversion of arable land to grassland, spatially heterogeneous soil properties, and climate. We used monthly mean NO3-N concentrations from 62 plots of a grassland plant diversity experiment from 2003 to 2006. Plant species richness (1–60) and functional group composition (1–4 functional groups: legumes, grasses, non-leguminous tall herbs, non-leguminous small herbs) were manipulated in a factorial design. Plant community composition, time since conversion from arable land to grassland, soil texture, and climate data (precipitation, soil moisture, air and soil temperature) were used to develop one general Bayesian multiple regression model for the 62 plots to allow an in-depth evaluation using the experimental design. The model simulated NO3-N concentrations with an overall Bayesian coefficient of determination of 0.48. The temporal course of NO3-N concentrations was simulated differently well for the individual plots with a maximum plot-specific Nash–Sutcliffe Efficiency of 0.57. The model shows that NO3-N concentrations decrease with species richness, but this relation reverses if more than approx. 25 % of legume species are included in the mixture. Presence of legumes increases and presence of grasses decreases NO3-N concentrations compared to mixtures containing only small and tall herbs. Altogether, our model shows that there is a strong influence of plant community composition on NO3-N concentrations.

A novel photographic approach for monitoring the structural heterogeneity and diversity of grassland ecosystems

Abstract: Aims Studies that investigate the space-filling heterogeneity of biological structures in plant communities remain scarce. The main objective of this study was to evaluate the relationship between newly developed photographic measures of structural heterogeneity in digital images and plant species composition in the context of a long-term grassland experiment. Methods We tested a close-range photographic protocol using measures of structural heterogeneity in gray-tone images, namely mean infor mation gain (MIG) and spatial anisotropy, to assess differences in the compositional (species richness) and functional characteristics (plant height and flowering) of 78 managed grassland communities. We also implemented a random placement model of community assembly to explore the links between our measures of structural complexity and the geometric pattern of plant communities. Important Findings MIG and spatial anisotropy correlated with the growth and species richness of grassland communities. Simulations showed that structural heterogeneity in gray-tone digital images is a function of the size distribution and orientation pattern of plant modules. This easy, fast and non-destructive methodological approach could eventually serve to monitor the diversity and integrity of various ecosystems at different resolutions across space and time.

Tree species traits but not diversity mitigate stem breakage in a subtropical forest following a rare and extreme ice storm

Abstract: Future climates are likely to include extreme events, which in turn have great impacts on ecological systems. In this study, we investigated possible effects that could mitigate stem breakage caused by a rare and extreme ice storm in a Chinese subtropical forest across a gradient of forest diversity. We used Bayesian modeling to correct stem breakage for tree size and variance components analysis to quantify the influence of taxon, leaf and wood functional traits, and stand level properties on the probability of stem breakage. We show that the taxon explained four times more variance in individual stem breakage than did stand level properties; trees with higher specific leaf area (SLA) were less susceptible to breakage. However, a large part of the variation at the taxon scale remained unexplained, implying that unmeasured or undefined traits could be used to predict damage caused by ice storms. When aggregated at the plot level, functional diversity and wood density increased after the ice storm. We suggest that for the adaption of forest management to climate change, much can still be learned from looking at functional traits at the taxon level.

Readable workflows need simple data

Abstract: Sharing scientific analyses via workflows has the potential to improve the reproducibility of research results as they allow complex tasks to be split into smaller pieces and give a visual access to the flow of data between the components of an analysis. This is particularly useful for trans-disciplinary research fields such as biodiversity and ecosystem functioning (BEF), where complex syntheses integrate data over large temporal, spatial and taxonomic scales. However, depending on the data used and the complexity of the analysis, scientific workflows can grow very complex which makes them hard to understand and reuse. Here we argue that enabling simplicity starting from the beginning of the data life cycle adhering to good practices of data management can significantly reduce the overall complexity of scientific workflows. It can simplify the processes of data inclusion, cleaning, merging and imputation. To illustrate our points we chose a typical analysis in BEF research, the aggregation of carbon pools in a forest ecosystem. We propose indicators to measure the complexity of workflow components including the data sources. We illustrate that the complexity decreases exponentially during the course of the analysis and that simple text-based measures can help to identify bottlenecks in a workflow. Taken together we argue that focusing on the simplification of data sources and workflow components will improve and accelerate data and workflow reuse and improve the reproducibility of data-driven sciences