Species richness, but not phylogenetic diversity, influences community biomass production and temporal stability in a re-examination of 16 grassland biodiversity studies

* Hundreds of experiments have now manipulated species richness (SR) of various groups of organisms and examined how this aspect of biological diversity influences ecosystem functioning. Ecologists have recently expanded this field to look at whether phylogenetic diversity (PD) among species, often quantified as the sum of branch lengths on a molecular phylogeny leading to all species in a community, also predicts ecological function. Some have hypothesized that phylogenetic divergence should be a superior predictor of ecological function than SR because evolutionary relatedness represents the degree of ecological and functional differentiation among species. But studies to date have provided mixed support for this hypothesis. * Here, we reanalyse data from 16 experiments that have manipulated plant SR in grassland ecosystems and examined the impact on above-ground biomass production over multiple time points. Using a new molecular phylogeny of the plant species used in these experiments, we quantified how the PD of plants impacts average community biomass production as well as the stability of community biomass production through time. * Using four complementary analyses, we show that, after statistically controlling for variation in SR, PD (the sum of branches in a molecular phylogenetic tree connecting all species in a community) is neither related to mean community biomass nor to the temporal stability of biomass. These results run counter to past claims. However, after controlling for SR, PD was positively related to variation in community biomass over time due to an increase in the variances of individual species, but this relationship was not strong enough to influence community stability. * In contrast to the non-significant relationships between PD, biomass and stability, our analyses show that SR per se tends to increase the mean biomass production of plant communities, after controlling for PD. The relationship between SR and temporal variation in community biomass was either positive, non-significant or negative depending on which analysis was used. However, the increases in community biomass with SR, independently of PD, always led to increased stability. These results suggest that PD is no better as a predictor of ecosystem functioning than SR. * Synthesis. Our study on grasslands offers a cautionary tale when trying to relate PD to ecosystem functioning suggesting that there may be ecologically important trait and functional variation among species that is not explained by phylogenetic relatedness. Our results fail to support the hypothesis that the conservation of evolutionarily distinct species would be more effective than the conservation of SR as a way to maintain productive and stable communities under changing environmental conditions.

Counterbalancing effects of competition for resources and facilitation against grazing in alpine snowbed communities

Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species-specific and/or dependent on ontogeny, and whether the balance of positive and negative plant–plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species-specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants.

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity-ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m(-2) and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m(-2). One year after seed addition, local plant species richness had increased on average by six species m(-2) (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m(-2) (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump-shaped model appears to be the limiting outline of the natural diversity-productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.

Direct and indirect associations between plant species richness and productivity in grasslands: regional differences preclude simple generalization of productivity-biodiversity relationships

Plant species richness of permanent grasslands has often been found to be significantly associated with productivity. Concentrations of nutrients in biomass can give further insight into these productivity- plant species richness relationships, e.g. by reflecting land use or soil characteristics. However, the consistency of such relationships across different regions has rarely been taken into account, which might significantly compromise our potential for generalization. We recorded plant species richness and measured above-ground biomass and concentrations of nutrients in biomass in 295 grasslands in three regions in Germany that differ in soil and climatic conditions. Structural equation modelling revealed that nutrient concentrations were mostly indirectly associated with plant species richness via biomass production. However, negative associations between the concentrations of different nutrients and biomass and plant species richness differed considerably among regions. While in two regions, more than 40% of the variation in plant species richness could be attributed to variation in biomass, K, P, and to some degree also N concentrations, in the third region only 15% of the variation could be explained in this way. Generally, highest plant species richness was recorded in grasslands where N and P were co-limiting plant growth, in contrast to N or K (co-) limitation. But again, this pattern was not recorded in the third region. While for two regions land-use intensity and especially the application of fertilizers are suggested to be the main drivers causing the observed negative associations with productivity, in the third region the little variance accounted for, low species richness and weak relationships implied that former intensive grassland management, ongoing mineralization of peat and fluctuating water levels in fen grasslands have overruled effects of current land-use intensity and productivity. Finally, we conclude that regional replication is of major importance for studies seeking general insights into productivity-diversity relationships.

Tree species and diversity effects on soil water seepage in a tropical plantation

Plant diversity has been shown to influence the water cycle of forest ecosystems by differences in water consumption and the associated effects on groundwater recharge. However, the effects of biodiversity on soil water fluxes remain poorly understood for native tree species plantations in the tropics. Therefore, we estimated soil water fluxes and assessed the effects of tree species and diversity on these fluxes in an experimental native tree species plantation in Sardinilla (Panama). The study was conducted during the wet season 2008 on plots of monocultures and mixtures of three or six tree species. Rainfall and soil water content were measured and evapotranspiration was estimated with the Penman-Monteith equation. Soil water fluxes were estimated using a simple soil water budget model considering water input, output, and soil water and groundwater storage changes and in addition, were simulated using the physically based one-dimensional water flow model Hydrus-1D. In general, the Hydrus simulation did not reflect the observed pressure heads, in that modeled pressure heads were higher compared to measured ones. On the other hand, the results of the water balance equation (WBE) reproduced observed water use patterns well. In monocultures, the downward fluxes through the 200 cm-depth plane were highest below Hura crepitans (6.13 mm day−1) and lowest below Luehea seemannii (5.18 mm day−1). The average seepage rate in monocultures (±SE) was 5.66 ± 0.18 mm day−1, and therefore, significantly higher than below six-species mixtures (5.49 ± 0.04 mm day−1) according to overyielding analyses. The three-species mixtures had an average seepage rate of 5.63 ± 0.12 mm day−1 and their values did not differ significantly from the average values of the corresponding species in monocultures. Seepage rates were driven by the transpiration of the varying biomass among the plots (r = 0.61, p = 0.017). Thus, a mixture of trees with different growth rates resulted in moderate seepage rates compared to monocultures of either fast growing or slow growing tree species. Our results demonstrate that tree-species specific biomass production and tree diversity are important controls of seepage rates in the Sardinilla plantation during the wet season.

Topoclimatological case-study of Alpine pastures near the Albula Pass in the eastern Swiss Alps

Alpine grasslands are an important source of fodder for the cattle of Alpine farmers. Only during the short summer season can these pastures be used for grazing. With the anticipated climate change, it is likely that plant production – and thus the fodder basis for the cattle – will be influenced. Investigating the dependence of biomass production on topoclimatic factors will allow us to better understand how anticipated climate change may influence this traditional Alpine farming system. Because small-scale topoclimatological variations of the main meteorological variables: temperature, humidity, precipitation, shortwave incoming radiation and wind speed are not easily derived from available long-term climate stations in mountainous terrain, it was our goal to investigate the topoclimatic variations over the pastures belonging to the Alp Weissenstein research station north of the Albula Pass in the eastern Swiss Alps. We present a basic assessment of current topoclimatic conditions as a site characterization for ongoing ecological climate change studies. To be able to link short-term studies with long-term climate records, we related agrometeorological measurements with those of surrounding long-term sites run by MeteoSwiss, both on valley bottoms (Davos, Samedan), and on mountain tops (Weissfluhjoch, Piz Corvatsch). We found that the Davos climate station north of the study area is most closely correlated with the local climate of Alp Weissenstein, although a much closer site (Samedan) exists on the other side of the Albula Pass. Mountain top stations, however, did not provide a convincing approximation for the climate at Alp Weissenstein. Direct comparisons of near-surface measurements from a set of 11 small weather stations distributed over the domain where cattle and sheep are grazed indicate that nocturnal minimum air temperature and minimum vapor pressure deficit are mostly governed by the altitudinal gradient, whereas daily maxima – including also wind speed – are more strongly depending on vegetation cover and less on the altitude.

Holocene climate, fire and vegetation dynamics at the treeline in the Northwestern Swiss Alps

Treelines are expected to rise to higher elevations with climate warming; the rate and extent however are still largely unknown. Here we present the first multi-proxy palaeoecological study from the treeline in the Northwestern Swiss Alps that covers the entire Holocene. We reconstructed climate, fire and vegetation dynamics at Iffigsee, an alpine lake at 2,065 m a.s.l., by using seismic sedimentary surveys, loss on ignition, visible spectrum reflectance spectroscopy, pollen, spore, macrofossil and charcoal analyses. Afforestation with Larix decidua and tree Betula (probably B. pendula) started at ~9,800 cal. b.p., more than 1,000 years later than at similar elevations in the Central and Southern Alps, indicating cooler temperatures and/or a high seasonality. Highest biomass production and forest position of ~2,100–2,300 m a.s.l. are inferred during the Holocene Thermal Maximum from 7,000 to 5,000 cal. b.p. With the onset of pastoralism and transhumance at 6,800–6,500 cal. b.p., human impact became an important factor in the vegetation dynamics at Iffigsee. This early evidence of pastoralism is documented by the presence of grazing indicators (pollen, spores), as well as a wealth of archaeological finds at the nearby mountain pass of Schnidejoch. Human and fire impact during the Neolithic and Bronze Ages led to the establishment of pastures and facilitated the expansion of Picea abies and Alnus viridis. We expect that in mountain areas with land abandonment, the treeline will react quickly to future climate warming by shifting to higher elevations, causing drastic changes in species distribution and composition as well as severe biodiversity losses.

Grazing response patterns indicate isolation of semi-natural European grasslands

Identifying drivers of species diversity is a major challenge in understanding and predicting the dynamics of species-rich semi-natural grasslands. In particular in temperate grasslands changes in land use and its consequences, i.e. increasing fragmentation, the on-going loss of habitat and the declining importance of regional processes such as seed dispersal by livestock, are considered key drivers of the diversity loss witnessed within the last decades. It is a largely unresolved question to what degree current temperate grassland communities already reflect a decline of regional processes such as longer distance seed dispersal. Answering this question is challenging since it requires both a mechanistic approach to community dynamics and a sufficient data basis that allows identifying general patterns. Here, we present results of a local individual- and trait-based community model that was initialized with plant functional types (PFTs) derived from an extensive empirical data set of species-rich grasslands within the `Biodiversity Exploratories' in Germany. Driving model processes included above- and belowground competition, dynamic resource allocation to shoots and roots, clonal growth, grazing, and local seed dispersal. To test for the impact of regional processes we also simulated seed input from a regional species pool. Model output, with and without regional seed input, was compared with empirical community response patterns along a grazing gradient. Simulated response patterns of changes in PFT richness, Shannon diversity, and biomass production matched observed grazing response patterns surprisingly well if only local processes were considered. Already low levels of additional regional seed input led to stronger deviations from empirical community pattern. While these findings cannot rule out that regional processes other than those considered in the modeling study potentially play a role in shaping the local grassland communities, our comparison indicates that European grasslands are largely isolated, i.e. local mechanisms explain observed community patterns to a large extent.

The importance of biotic interactions and local adaptation for plant response to environmental changes: field evidence along an elevational gradient

Predicting the response of species to environmental changes is a great and on-going challenge for ecologists, and this requires a more in-depth understanding of the importance of biotic interactions and the population structuration in the landscape. Using a reciprocal transplantation experiment, we tested the response of five species to an elevational gradient. This was combined to a neighbour removal treatment to test the importance of local adaptation and biotic interactions. The trait studied was performance measured as survival and biomass. Species response varied along the elevational gradient, but with no consistent pattern. Performance of species was influenced by environmental conditions occurring locally at each site, as well as by positive or negative effects of the surrounding vegetation. Indeed, we observed a shift from competition for biomass to facilitation for survival as a response to the increase in environmental stress occurring in the different sites. Unlike previous studies pointing out an increase of stress along the elevation gradient, our results supported a stress gradient related to water availability, which was not strictly parallel to the elevational gradient. For three of our species, we observed a greater biomass production for the population coming from the site where the species was dominant (central population) compared to population sampled at the limit of the distribution (marginal population). Nevertheless, we did not observe any pattern of local adaptation that could indicate adaptation of populations to a particular habitat. Altogether, our results highlighted the great ability of plant species to cope with environmental changes, with no local adaptation and great variability in response to local conditions. Our study confirms the importance of taking into account biotic interactions and population structure occurring at local scale in the prediction of communities’ responses to global environmental changes.

Activated carbon may have undesired side effects for testing allelopathy in invasive plants

Activated carbon has become a widely used tool to investigate root-mediated allelopathy of plants, especially in plant invasion biology, because it adsorbs and thereby neutralizes root exudates. Allelopathy has been a controversially debated phenomenon for years, which revived in plant invasion biology as one possible reason for the success of invasive plants. Noxious plant exudates may harm other plants and provide an advantage to the allelopathic plant. However, root exudates are not always toxic, but may stimulate the microbial community and change nutrient availability in the rhizosphere. In a greenhouse experiment, we investigated the interacting effects of activated carbon, arbuscular mycorrhiza and plant competition between the invasive Senecio inaequidens and the native Artemisia vulgaris. Furthermore, we tested whether activated carbon showed any undesired effects by directly affecting mycorrhiza or soil chemistry. Contrary to the expectation, S. inaequidens was a weak competitor and we could not support the idea that allelopathy was involved in the competition. Activated carbon led to a considerable increase in the aboveground biomass production and reduced the infection with arbuscular mycorrhiza of both plant species. We expected that arbuscular mycorrhiza promotes plant growth by increasing nutrient availability, but we found the contrary when activated carbon was added. Chemical analyses of the substrate showed, that adding activated carbon resulted in a strong increase in plant available phosphate and in a decrease of the C(organic)/N(total) ration both of which suggest stimulated microbial activity. Thus, activated carbon not only reduced potential allelopathic effects, but substantially changed the chemistry of the substrate. These results show that activated carbon should be handled with great care in ecological experiments on allelopathy because of possible confounding effects on the soil community.

Higher plant diversity promotes higher diversity of fungal pathogens, while it decreases pathogen infection per plant

Fungal plant pathogens are common in natural communities where they affect plant physiology, plant survival, and biomass production. Conversely, pathogen transmission and infection may be regulated by plant community characteristics such as plant species diversity and functional composition that favor pathogen diversity through increases in host diversity while simultaneously reducing pathogen infection via increased variability in host density and spatial heterogeneity. Therefore, a comprehensive understanding of multi-host multi-pathogen interactions is of high significance in the context of biodiversity-ecosystem functioning. We investigated the relationship between plant diversity and aboveground obligate parasitic fungal pathogen (''pathogens'' hereafter) diversity and infection in grasslands of a long-term, large-scale, biodiversity experiment with varying plant species (1-60 species) and plant functional group diversity (1-4 groups). To estimate pathogen infection of the plant communities, we visually assessed pathogen-group presence (i.e., rusts, powdery mildews, downy mildews, smuts, and leaf-spot diseases) and overall infection levels (combining incidence and severity of each pathogen group) in 82 experimental plots on all aboveground organs of all plant species per plot during four surveys in 2006. Pathogen diversity, assessed as the cumulative number of pathogen groups on all plant species per plot, increased log-linearly with plant species diversity. However, pathogen incidence and severity, and hence overall infection, decreased with increasing plant species diversity. In addition, co-infection of plant individuals by two or more pathogen groups was less likely with increasing plant community diversity. We conclude that plant community diversity promotes pathogen-community diversity while at the same time reducing pathogen infection levels of plant individuals.

Little evidence for release from herbivores as a driver of plant invasiveness from a multi-species herbivore-removal experiment

Enemy release is frequently posed as a main driver of invasiveness of alien species. However, an experimental multi-species test examining performance and herbivory of invasive alien, non-invasive alien and native plant species in the presence and absence of natural enemies is lacking. In a common garden experiment in Switzerland, we manipulated exposure of seven alien invasive, eight alien non-invasive and fourteen native species from six taxonomic groups to natural enemies (invertebrate herbivores), by applying a pesticide treatment under two different nutrient levels. We assessed biomass production, herbivore damage and the major herbivore taxa on plants. Across all species, plants gained significantly greater biomass under pesticide treatment. However, invasive, non-invasive and native species did not differ in their biomass response to pesticide treatment at either nutrient level. The proportion of leaves damaged on invasive species was significantly lower compared to native species, but not when compared to non-invasive species. However, the difference was lost when plant size was accounted for. There were no differences between invasive, non-invasive and native species in herbivore abundance. Our study offers little support for invertebrate herbivore release as a driver of plant invasiveness, but suggests that future enemy release studies should account for differences in plant size among species.

Aluminum toxicity in a tropical montane forest ecosystem in southern Ecuador

Aluminum phytotoxicity frequently occurs in acid soils (pH < 5.5) and was therefore discussed to affect ecosystem functioning of tropical montane forests. The susceptibility to Al toxicity depends on the sensitivity of the plant species and the Al speciation in soil solution, which can vary highly depending e.g., on pH, ionic strength, and dissolved organic matter. An acidification of the ecosystem and periodic base metal deposition from Saharan dust may control plant available Al concentrations in the soil solutions of tropical montane rainforests in south Ecuador. The overall objective of my study was to assess a potential Al phytotoxicity in the tropical montane forests in south Ecuador. For this purpose, I exposed three native Al non-accumulating tree species (Cedrela odorata L., Heliocarpus americanus L., and Tabebuia chrysantha (Jacq.) G. Nicholson) to increased Al concentrations (0 – 2400 μM Al) in a hydroponic experiment, I established dose-response curves to estimate the sensitivity of the tree species to increased Al concentrations, and I investigated the mechanisms behind the observed effects induced by elevated Al concentrations. Furthermore, the response of Al concentrations and the speciation in soil solution to Ca amendment in the study area were determined. In a final step, I assessed all major Al fluxes, drivers of Al concentrations in ecosystem solutions, and indicators of Al toxicity in the tropical montane rainforest in Ecuador in order to test for indications of Al toxicity. In the hydroponic experiment, a 10 % reduction in aboveground biomass production occurred at 126 to 376 μM Al (EC10 values), probably attributable to decreased Mg concentrations in leaves and reduced potosynthesis. At 300 μM Al, increased root biomass production of T. chrysantha was observed. Phosphorus concentrations in roots of C. odorata and T. chrysantha were significantly highest in the treatment with 300 μM Al and correlated significantly with root biomass, being a likely reason for stimulated root biomass production. The degree of organic complexation of Al in the organic layer leachate, which is central to plant nutrition because of the high root density, and soil solution from the study area was very high (mean > 99 %). The resulting low free Al concentrations are not likely to affect plant growth, although the concentrations of potentially toxic Al3+ increased with soil depth due to higher total Al and lower dissolved organic matter concentrations in soil solutions. The Ca additions caused an increase of Al in the organic layer leachate, probably because Al3+ was exchanged against the added Ca2+ ions while pH remained constant. The free ion molar ratios of Ca2+:Al3+ (mean ratio ca. 400) were far above the threshold (≤ 1) for Al toxicity, because of a much higher degree of organo-complexation of Al than Ca. High Al fluxes in litterfall (8.8 – 14.2 kg ha−1 yr−1) indicate a high Al circulation through the ecosystem. The Al concentrations in the organic layer leachate were driven by the acidification of the ecosystem and increased significantly between 1999 and 2008. However, the Ca:Al molar ratios in organic layer leachate and all aboveground ecosystem solutions were above the threshold for Al toxicity. Except for two Al accumulating and one non-accumulating tree species, the Ca:Al molar ratios in tree leaves from the study area were above the Al toxicity threshold of 12.5. I conclude that toxic effects in the hydroponic experiment occurred at Al concentrations far above those in native organic layer leachate, shoot biomass production was likely inhibited by reduced Mg uptake, impairing photosynthesis, and the stimulation of root growth at low Al concentrations can be possibly attributed to improved P uptake. Dissolved organic matter in soil solutions detoxifies Al in acidic tropical forest soils and a wide distribution of Al accumulating tree species and high Al fluxes in the ecosystem do not necessarily imply a general Al phytotoxicity.

Spatial patterns and species performances in experimental plant communities

Amongst the various hypotheses that challenged to explain the coexistence of species with similar life histories, theoretical, and empirical studies suggest that spatial processes may slow down competitive exclusion and hence promote coexistence even in the absence of evident trade-offs and frequent disturbances. We investigated the effects of spatial pattern and density on the relative importance of intra- and interspecific competition in a field experiment. We hypothesized that weak competitors increased biomass and seed production within neighborhoods of conspecifics, while stronger competitors would show increased biomass and seed production within neighborhoods of heterospecifics. Seeds of four annual plant species (Capsella bursa-pastoris, Stachys annua, Stellaria media, Poa annua) were sown in two spatial patterns (aggregated vs. random) and at two densities (low vs. high) in three different species combinations (monocultures, three and four species mixtures). There was a hierarchy in biomass production among the four species and C. bursa-pastoris and S. media were among the weak competitors. Capsella and Stellaria showed increased biomass production and had more individuals in the aggregated compared to the random pattern, especially when both superior competitors (S. annua, P. annua) were present. For P. annua we observed considerable differences among species combinations and unexpected pattern effects. Our findings support the hypothesis that weak competitors increase their fitness when grown in the neighborhood of conspecifics, and suggested that for the weakest competitors the species identity is not important and all other species are best avoided through intraspecific aggregation. In addition, our data suggest that the importance of spatial pattern for the other competitors might not only depend on the position within the hierarchy but also on the identity of neighbor species, species characteristics, below ground interactions, and other nonspatial factors.