Leaf Traits, Neighbors, and Abiotic Factors: Ways That Context Can Mediate the Impact of Invasive Species on Nitrogen Cycling

Species invasions are more prevalent than ever before. While the addition of a species can dramatically change critical ecosystem processes, factors that mediate the direction and magnitude of those impacts have received less attention. A better understanding of the factors that mediate invasion impacts on ecosystem functioning is needed in order to target which exotic species will be most harmful and which systems are most vulnerable. The role of invasion on nitrogen (N) cycling is particularly important since N cycling controls ecosystem services that provision human health, e.g. nutrient retention and water quality.

We conducted a meta-analysis and in-depth studies focused on the invasive grass species, Microstegium vimineum, to better understand how (i) plant characteristics, (ii) invader abundance and neighbor identity, and (iii) environmental conditions mediate the impacts of invasion on N pools and fluxes. The results of our global meta-analysis support the concept that invasive species and reference community traits such as leaf %N and leaf C:N are useful for understanding invasion impacts on soil N cycling, but that trait dissimilarities between invaded and reference communities are most informative. Regarding the in-depth studies of Microstegium, we did not find evidence to suggest that invasion increases net nitrification as other studies have shown. Instead, we found that an interaction between its abundance and the neighboring plant identify were important for determining soil nitrate concentrations and net nitrification rates in the greenhouse. In field, we found that variability in environmental conditions mediated the impact of Microstegium invasion on soil N pools and fluxes, primarily net ammonification, between sites through direct, indirect, and interactive pathways. Notably, we detected a scenario in which forest openness has a negative direct effect and indirect positive effect on ammonification in sites with high soil moisture and organic matter. Collectively, our findings suggest that dissimilarity in plant community traits, neighbor identity, and environmental conditions can be important drivers of invasion impacts on ecosystem N cycling and should be considered when evaluating the ecosystem impacts of invasive species across heterogeneous landscapes.

Collection of aboveground community and species-specific plant biomass from the Jena Experiment (time series since 2002).

This data set comprises time series of aboveground community plant biomass (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) and species-specific biomass from the sown species of several experiments at the field site of a large grassland biodiversity experiment (the Jena Experiment; see further details below). Aboveground community biomass was normally harvested twice a year just prior to mowing (during peak standing biomass twice a year, generally in May and August; in 2002 only once in September) on all experimental plots in the Jena Experiment. This was done by clipping the vegetation at 3 cm above ground in up to four rectangles of 0.2 x 0.5 m per large plot. The location of these rectangles was assigned by random selection of new coordinates every year within the core area of the plots. The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: individual species for the sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. Sown plant community biomass was calculated as the sum of the biomass of the individual sown species. The data for individual samples and the mean over samples for the biomass measures on the community level are given. Overall, analyses of the community biomass data have identified species richness as well as functional group composition as important drivers of a positive biodiversity-productivity relationship. The following series of datasets are contained in this collection: 1. Plant biomass form the Main Experiment: In the Main Experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). 2. Plant biomass from the Dominance Experiment: In the Dominance Experiment, 206 grassland plots of 3.5 x 3.5 m were established from a pool of 9 species that can be dominant in semi-natural grassland communities of the study region. In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 3, 4, 6, and 9 species). 3. Plant biomass from the monoculture plots: In the monoculture plots the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species like the other experiments in May 2002. All plots were maintained by bi-annual weeding and mowing.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2003)

This data set contains aboveground plant biomass in 2003 (Sown plant community, Weed plant community, and Dead plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2003 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. excluding an outer edge of 0.5 m). The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), and detached dead plant material (i.e., dead plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2010)

This data set contains aboveground plant biomass in 2010 (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. One of the replicate plots per species was given up after the vegetation period of 2007 for all but the nine species belonging also to the so called dominance experiment in Jena. These nine species are: Alopecurus pratensis, Anthriscus sylvestris, Arrhenatherum elatius, Dactylis glomerata, Geranium pratense, Poa trivialis, Phleum pratense, Trifolium repens and Trifolium pratense.In 2010 plot size was reduced to 1 x 1 m. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2010 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 1 rectangle of 0.2 x 0.5 m per plot. The location of this rectangle was in the center of the plot area. The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2005)

This data set contains aboveground plant biomass in 2005 (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2005 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. excluding an outer edge of 0.5 m). The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2006)

This data set contains aboveground plant biomass in 2006 (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2006 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. excluding an outer edge of 0.5 m). The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2007)

This data set contains aboveground plant biomass in 2007 (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2007 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. excluding an outer edge of 0.5 m). The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2008)

This data set contains aboveground plant biomass in 2008 (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. One of the replicate plots per species was given up after the vegetation period of 2007 for all but the nine species belonging also to the so called dominance experiment in Jena. These nine species are: Alopecurus pratensis, Anthriscus sylvestris, Arrhenatherum elatius, Dactylis glomerata, Geranium pratense, Poa trivialis, Phleum pratense, Trifolium repens and Trifolium pratense.In 2008 plot size was reduced to 2.5 x 2.5 m. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2008 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. excluding an outer edge of 0.5 m). The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2009)

This data set contains aboveground plant biomass in 2009 (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. One of the replicate plots per species was given up after the vegetation period of 2007 for all but the nine species belonging also to the so called dominance experiment in Jena. These nine species are: Alopecurus pratensis, Anthriscus sylvestris, Arrhenatherum elatius, Dactylis glomerata, Geranium pratense, Poa trivialis, Phleum pratense, Trifolium repens and Trifolium pratense.In 2008 plot size was reduced to 2.5 x 2.5 m. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2009 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was in the center of the plot area. The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2002)

This data set contains aboveground plant biomass in 2002 (Sown plant community; measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2002 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. excluding an outer edge of 0.5 m). The positions of the rectangles within plots were identical for all plots. From the harvested biomass only the separated biomass of the sown plant species was kept. All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Aboveground plant biomass from the Jena Experiment (Monocultures, year 2004)

This data set contains aboveground plant biomass in 2004 (Sown plant community, Weed plant community, Dead plant material, and Unidentified plant material; all measured in biomass as dry weight) of the monoculture plots of a large grassland biodiversity experiment (the Jena Experiment). In the monoculture plots the biomass of the sown plant community contains only a single species per plot and this species is a different one for each plot. Which species has been sown in which plot is stated in the plot information table for monocultures (see further details below). The monoculture plots of 3.5 x 3.5 m were established for all of the 60 plant species of the Jena Experiment species pool with two replicates per species. These 60 species comprising the species pool of the Jena Experiment belong to four functional groups (grasses, legumes, tall and small herbs). Plots were sown in May 2002 and are since maintained by bi-annual weeding and mowing. Aboveground plant biomass was harvested twice in 2004 just prior to mowing (during peak standing biomass in early June and in late August) on all experimental plots of the monocultures. This was done by clipping the vegetation at 3 cm above ground in 2 rectangles of 0.2 x 0.5 m per plot. The location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. excluding an outer edge of 0.5 m). The positions of the rectangles within plots were identical for all plots. The harvested biomass was sorted into categories: sown plant species, weed plant species (species not sown at the particular plot), detached dead plant material (i.e., dead plant material in the data file), and remaining plant material that could not be assigned to any category (i.e., unidentified plant material in the data file). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data for individual subsamples (i.e. rectangles) and the mean over samples for all biomass measures are given.

Irrigation and drainage canals role for plant diversity and nature conservation

Aims: With this research, we wanted to investigate and promote the conservation of biodiversity in the network of drainage canals of the Po Valley Study area: The canal network of Bologna plain, long more than 1150 km (Po Valley, North Italy) Methods: In Chapter II we analyzed the geographical patterns that characterize our transects, the land use of their upstream basins, the water quality at the closure points of their river basins. In Chapter III we described the plant communities with some ecological information and we also tested the effect of the canal size on the plant communities. In Chapter IV we described the relation beetween some functional traits of the plant species sampled and some environmental parameters Results: A total of 272 species were sampled in 118 transects. The plant communities of the drainage canals have been found to have a significant influence: the geographical pattern "proximity to protected areas", the class of land use "agrozootechnical settlements", and some water parameters. The analysis of the parameter "canal depth" indicated a significant distinction between small and large canals based on plant communities. The functional composition of the plant communities was affected by the bank aspect, the inclusion/exclusion from the protected areas and the upstream basin land uses. Moreover, the functional groups of species responded differently to environmental drivers, water quality gradients and were influenced by a combination of environmental stresses Conclusions: This research confirms the key role of the canals network in sustaining the plant richness in oversimplified landscapes. Considering the fragility of the floodplains and the global warming that is taking place, it is necessary to rethink the role of irrigation canals and their plant communities in the near future. This work reinforces the belief that long-term sampling plans and greater knowledge about canal management practices are needed

Leaf Manganese Accumulation And Phosphorus-acquisition Efficiency.

Plants that deploy a phosphorus (P)-mobilising strategy based on the release of carboxylates tend to have high leaf manganese concentrations ([Mn]). This occurs because the carboxylates mobilise not only soil inorganic and organic P, but also a range of micronutrients, including Mn. Concentrations of most other micronutrients increase to a small extent, but Mn accumulates to significant levels, even when plants grow in soil with low concentrations of exchangeable Mn availability. Here, we propose that leaf [Mn] can be used to select for genotypes that are more efficient at acquiring P when soil P availability is low. Likewise, leaf [Mn] can be used to screen for belowground functional traits related to nutrient-acquisition strategies among species in low-P habitats.

Functional biogeography of oceanic islands and the scaling of functional diversity in the Azores

Analyses of species-diversity patterns of remote islands have been crucial to the development of biogeographic theory, yet little is known about corresponding patterns in functional traits on islands and how, for example, they may be affected by the introduction of exotic species. We collated trait data for spiders and beetles and used a functional diversity index (FRic) to test for nonrandomness in the contribution of endemic, other native (also combined as indigenous), and exotic species to functional-trait space across the nine islands of the Azores. In general, for both taxa and for each distributional category, functional diversity increases with species richness, which, in turn scales with island area. Null simulations support the hypothesis that each distributional group contributes to functional diversity in proportion to their species richness. Exotic spiders have added novel trait space to a greater degree than have exotic beetles, likely indicating greater impact of the reduction of immigration filters and/or differential historical losses of indigenous species. Analyses of species occurring in native-forest remnants provide limited indications of the operation of habitat filtering of exotics for three islands, but only for beetles. Although the general linear (not saturating) pattern of trait-space increase with richness of exotics suggests an ongoing process of functional enrichment and accommodation, further work is urgently needed to determine how estimates of extinction debt of indigenous species should be adjusted in the light of these findings.

Species distribution models reveal apparent competitive and facilitative effects of a dominant species on the distribution of tundra plants

Abiotic factors are considered strong drivers of species distribution and assemblages. Yet these spatial patterns are also influenced by biotic interactions. Accounting for competitors or facilitators may improve both the fit and the predictive power of species distribution models (SDMs). We investigated the influence of a dominant species, Empetrum nigrum ssp. hermaphroditum, on the distribution of 34 subordinate species in the tundra of northern Norway. We related SDM parameters of those subordinate species to their functional traits and their co-occurrence patterns with E. hermaphroditum across three spatial scales. By combining both approaches, we sought to understand whether these species may be limited by competitive interactions and/or benefit from habitat conditions created by the dominant species. The model fit and predictive power increased for most species when the frequency of occurrence of E. hermaphroditum was included in the SDMs as a predictor. The largest increase was found for species that 1) co-occur most of the time with E. hermaphroditum, both at large (i.e. 750 m) and small spatial scale (i.e. 2 m) or co-occur with E. hermaphroditum at large scale but not at small scale and 2) have particularly low or high leaf dry matter content (LDMC). Species that do not co-occur with E. hermaphroditum at the smallest scale are generally palatable herbaceous species with low LDMC, thus showing a weak ability to tolerate resource depletion that is directly or indirectly induced by E. hermaphroditum. Species with high LDMC, showing a better aptitude to face resource depletion and grazing, are often found in the proximity of E. hermaphroditum. Our results are consistent with previous findings that both competition and facilitation structure plant distribution and assemblages in the Arctic tundra. The functional and co-occurrence approaches used were complementary and provided a deeper understanding of the observed patterns by refinement of the pool of potential direct and indirect ecological effects of E. hermaphroditum on the distribution of subordinate species. Our correlative study would benefit being complemented by experimental approaches.