Biodiversity eVects on ecosystem functioning in forests have only recently attracted increasing attention. The vast majority of studies in forests have focused on above-ground responses to diVerences in tree species diversity, while systematic analyses of the eVects of biodiversity on root systems are virtually non-existent. By investigating the Wne root systems in 12 temperate deciduous forest stands in Central Europe, we tested the hypotheses that (1) stand Wne root biomass increases with tree diversity, and (2) 'below-ground overyielding' of species-rich stands in terms of Wne root biomass is the consequence of spatial niche segregation of the roots of diVerent species. The selected stands represent a gradient in tree species diversity on similar bedrock from almost pure beech forests to mediumdiverse forests built by beech, ash, and lime, and highlydiverse stands dominated by beech, ash, lime, maple, and hornbeam. We investigated Wne root biomass and necromass at 24 proWles per stand and analyzed species diVerences in Wne root morphology by microscopic analysis. Fine root biomass ranged from 440 to 480 g m ¡2 in the species-poor to species-rich stands, with 63-77% being concentrated in the upper 20 cm of the soil. In contradiction to our two hypotheses, the diVerences in tree species diversity aVected neither stand Wne root biomass nor vertical root distribution patterns. Fine root morphology showed marked distinctions between species, but these root morphological diVerences did not lead to signiWcant diVerences in Wne root surface area or root tip number on a stand area basis. Moreover, diVerences in species composition of the stands did not alter Wne root morphology of the species. We conclude that 'below-ground overyielding' in terms of Wne root biomass does not occur in the species-rich stands, which is most likely caused by the absence of signiWcant spatial segregation of the root systems of these late-successional species.

Background Studying root biomass, root system distribution and belowground interactions is essential for understanding the composition of plant communities, the impact of global change, and terrestrial biogeochemistry. Most soil samples and minirhizotron pictures hold roots of more than one species or plant individual. The identification of taxa by their roots would allow species-specific questions to be posed; information about root affiliation to plant individuals could be used to determine intra-specific competition. Scope Researchers need to be able to discern plant taxa by roots as well as to quantify abundances in mixed root samples. However, roots show less distinctive features that permit identification than aboveground organs. This review discusses the primary use of available methods, outlining applications, shortcomings and future developments. Conclusion Methods are either non-destructive, e.g. visual examination of root morphological criteria in situ, or require excavated and excised root samples. Among the destructive methods are anatomical keys, chemotaxonomic approaches and molecular markers. While some methods allow for discerning the root systems of individual plants, others can distinguish roots on the functional group or plant taxa level; methods such as IR spectroscopy and qPCR allow for quantifying the root biomass proportion of species without manual sorting.

Differences in spatial rooting patterns among coexisting species have been recognized as an important mechanism for generating biodiversity effects on ecosystem functioning. However, it is not yet clear whether complementarity in root space exploration is a universal characteristic of multi-species woody communities. In a temperate broad-leaved forest with a mosaic of speciespoor and species-rich stands, we tested two hypotheses related to putative below-ground 'overyielding' in more diverse forests, (1) that species mixture results in a partial spatial segregation of the fine root systems of different species, and (2) that stand fine root biomass increases with tree species diversity. We investigated 12 stands either with one, three, or five dominant tree species (4 replicate stands each) under similar soil and climate conditions for stand fine root biomass and spatial root segregation in vertical and horizontal direction in the soil. Fine roots of different tree species were identified using a morphological key based on differences in colour, periderm surface structure, and branching patterns. In species-poor and species-rich stands, and in all tree species present, fine root density (biomass per soil volume) decreased exponentially with soil depth at very similar rates. Stand fine root biomass in the densely rooted upper soil (0-40 cm depth) was not significantly different between stands with 1, 3 or 5 dominant tree species. We conclude that 'below-ground overyielding' in terms of higher fine root biomasses in species-rich stands as compared to monospecific ones does not occur in these broad-leaved forests which most likely results from a missing complementarity in vertical rooting patterns of the present tree species.

In contrast to studies on aboveground processes, the effect of species diversity on belowground productivity and fine-root regrowth after disturbance is still poorly studied in forests. In 12 oldgrowth broad-leaved forest stands, we tested the hypotheses that (i) the productivity and recovery rate (regrowth per standing biomass) of the fineroot system (root diameter < 2 mm) increase with increasing tree species diversity, and that (ii) the seasonality of fine-root biomass and necromass is more pronounced in pure than in tree species-rich stands as a consequence of non-synchronous root biomass peaks of the different species. We investigated stands with 1, 3, and 5 dominant tree species growing under similar soil and climate conditions for changes in fine-root biomass and necromass during a 12-month period and estimated fine-root productivity with two independent approaches (ingrowth cores, sequential coring). According to the analysis of 360 ingrowth cores, fine-root growth into the root-free soil increased with tree species diversity from 72 g m -2 y -1 in the monospecific plots to 166 g m -2 y -1 in the 5-species plots, indicating an enhanced recovery rate of the root system after soil disturbance with increasing species diversity (0.26, 0.34, and 0.51 y -1 in 1-, 3-, and 5-species plots, respectively). Fine-root productivity as approximated by the sequential coring data also indicated a roughly threefold increase from the monospecific to the 5-species stand. We found no indication of a more pronounced seasonality of fine-root mass in species-poor as compared to species-rich stands. We conclude that species identification on the fine root level, as conducted here, may open new perspectives on tree species effects on root system dynamics. Our study produced first evidence in support of the hypothesis that the fine-root systems of more diverse forest stands are more productive and recover more rapidly after soil disturbance than that of species-poor forests.

Root discrimination of species is a pre-condition for studying belowground competition processes between crop and weed species. In this experiment, we tested Fourier transform mid-infrared (FT MIR)-attenuated total reflection (ATR) spectroscopy to discriminate roots of closely related crop and weed species grown in the greenhouse: maize/barnyard grass, barley/wild oat, wheat/blackgrass (Poaceae), and sugar beet/common lambsquarters (Chenopodiaceae). Fresh (moist) and dried root segments as well as ground roots were analyzed by FT MIR-ATR spectroscopy. Root absorption spectra showed species specific peak distribution and peak height. A clear separation according to species was not possible with fresh root segments. Dried root segments (including root basis, middle section, and root tip) of maize/barnyard grass and sugar beet/common lambsquarters formed completely separated species clusters. Wheat and blackgrass separated in species specific clusters when root tips were removed from cluster analysis. A clear separation of dried root segments according to species was not possible in the case of barley and wild oat. Cluster analyses of ground roots revealed a 100% separation of all tested crop and weed species combinations. Spectra grouped in Poaceae and Chenopodiaceae clusters. Within the Poaceae cluster, C3 and C4 species differed significantly in heterogeneity. Thus, root spectra reflected the degree of kinship. To quantify species proportion in root mixtures, a two- and a three-species model for species quantification in root mixtures of maize, barnyard grass, and wild oat was calculated. The models showed low standard errors of prediction (RMSEP) and high residual predictive deviation values in an external test set validation. Hence, FT MIR-ATR spectroscopy seems to be a promising tool for root research even between closely related plant species.

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The differentiation of roots of agricultural species is desired for a deeper understanding of the belowground root interaction which helps to understand the complex interaction in intercropping and crop-weed systems. The roots can be reliably differentiated via Fourier transform infrared spectroscopy with attenuated total reflection (FTIR-ATR). In two replicated greenhouse experiments, six pea cultivars, five oat cultivars as well as seven maize cultivars and five barnyard grass proveniences (n = 10 plants/cultivar or provenience) were grown under controlled conditions. One root of each plant was harvested and five different root segments of each root were separated, dried and measured with FTIR-ATR spectroscopy. The results showed that, firstly, the root spectra of single pea and single oat cultivars as well as single maize and single barnyard grass cultivars/proveniences separated species-specific in cluster analyses. In the majority of cases the species separation was correct, but in a few cases, the spectra of the root tips had to be omitted to ensure the precise separation between the species. Therefore, species differentiation is possible regardless of the cultivar or provenience. Consequently, all tested cultivars of pea and oat spectra were analyzed together and separated within a cluster analysis according to their affiliated species. The same result was found in a cluster analysis with maize and barnyard grass spectra. Secondly, a cluster analysis with all species (pea, oat, maize and barnyard grass) was performed. The species split up species-specific and formed a dicotyledonous pea cluster and a monocotyledonous cluster subdivided in oat, maize and barnyard grass subclusters. Thirdly, cultivar or provenience differentiations within one species were possible in one of the two replicated experiments. But these separations were less resilient.

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