Marine meso-herbivore consumption scales faster with temperature than seaweed primary production, supplementary material

Respiration of ectotherms is predicted to increase faster with rising environmental temperature than photosynthesis of primary producers because of the differential temperature dependent kinetics of the key enzymes involved. Accordingly, if biological processes at higher levels of complexity are constrained by underlying metabolic functions food consumption by heterotrophs should increase more rapidly with rising temperature than photo-autoptrophic primary production. We compared rates of photosynthesis and growth of the benthic seaweed Fucus vesiculosus with respiration and consumption of the isopod Idotea baltica to achieve a mechanistic understanding why warming strengthens marine plant-herbivore interactions. In laboratory experiments thallus pieces of the seaweed and individuals of the grazer were exposed to constant temperatures at a range from 10 to 20°C. Photosynthesis of F. vesiculosus did not vary with temperature indicating efficient thermal acclimation whereas growth of the algae clearly increased with temperature. Respiration and food consumption of I. baltica also increased with temperature. Grazer consumption scaled about 2.5 times faster with temperature than seaweed production. The resulting mismatch between algal production and herbivore consumption may result in a net loss of algal tissue at elevated temperatures. Our study provides an explanation for faster decomposition of seaweeds at elevated temperatures despite the positive effects of high temperatures on algal growth.

(Table A1) Annual biomass of plants and animals recorded on Bylot Island between 1993-2009

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator--rey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.

Initial biomass and biomass change of tundra and forest plants in three herbivore treatments in Norway and Sweden

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana, has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (> 0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (< 0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.

Aboveground community and species-specific plant biomass from the Jena Experiment (Main Experiment, year 2004)

This data set contains aboveground community 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 the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). 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). Plots were maintained by bi-annual weeding and mowing. Aboveground community biomass was harvested twice in 2004 just prior to mowing (during peak standing biomass in late May and in late August) on all experimental plots of the main experiment. This was done by clipping the vegetation at 3 cm above ground in four rectangles of 0.2 x 0.5 m per large 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. the central 10 x 15 m). 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.

(Table 1) Species richness and diversity of tundra plants in herbivore exclosure and control plots near Barrow, Alaska

To determine the role lemmings play in structuring plant communities and their contribution to the 'greening of the Arctic', we measured plant cover and biomass in 50 + year old lemming exclosures and control plots in the coastal tundra near Barrow, Alaska. The response of plant functional types to herbivore exclusion varied among land cover types. In general, the abundance of lichens and bryophytes increased with the exclusion of lemmings, whereas graminoids decreased, although the magnitude of these responses varied among land cover types. These results suggest that sustained lemming activity promotes a higher biomass of vascular plant functional types than would be expected without their presence and highlights the importance of considering herbivory when interpreting patterns of greening in the Arctic. In light of the rapid environmental change ongoing in the Arctic and the potential regional to global implications of this change, further exploration regarding the long-term influence of arvicoline rodents on ecosystem function (e.g. carbon and energy balance) should be considered a research priority.

Mean cover and frequencies of vascular plant species during three decades around Abisko Scientific Research Station

Plant species distributions are expected to shift and diversity is expected to decline as a result of global climate change, particularly in the Arctic where climate warming is amplified. We have recorded the changes in richness and abundance of vascular plants at Abisko, sub-Arctic Sweden, by re-sampling five studies consisting of seven datasets; one in the mountain birch forest and six at open sites. The oldest study was initiated in 1977-1979 and the latest in 1992. Total species number increased at all sites except for the birch forest site where richness decreased. We found no general pattern in how composition of vascular plants has changed over time. Three species, Calamagrostis lapponica, Carex vaginata and Salix reticulata, showed an overall increase in cover/frequency, while two Equisetum taxa decreased. Instead, we showed that the magnitude and direction of changes in species richness and composition differ among sites.

Aboveground community and species-specific plant biomass from the Jena Experiment (Main Experiment, year 2007)

This data set contains aboveground community 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 the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). 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). Plots were maintained by bi-annual weeding and mowing. Aboveground community 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 main experiment. This was done by clipping the vegetation at 3 cm above ground in four (May) or three (August) rectangles of 0.2 x 0.5 m per large 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. the central 10 x 15 m). 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.

Aboveground community and species-specific plant biomass from the Jena Experiment (Main Experiment, year 2006)

This data set contains aboveground community 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 the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). 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). Plots were maintained by bi-annual weeding and mowing. Aboveground community 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 main experiment. This was done by clipping the vegetation at 3 cm above ground in four rectangles of 0.2 x 0.5 m per large 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. the central 10 x 15 m). 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.

Collection of data on plant height along the plant diversity gradient in the Jena Experiment (Main Experiment, time series since 2002)

This data set contains a time series of plant height measurements (vegetative and reproductive) from the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In addition, data on species specific plant heights for the main experiment are available from 2002. 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). Plots were maintained by bi-annual weeding and mowing. 1. Plant height was recorded, generally, twice a year just before biomass harvest (during peak standing biomass in late May and in late August). Methodologies of measuring height have varied somewhat over the years. In earlier year the streched plant height was measured, while in later years the standing height without streching the plant was measured. Vegetative height was measured either as the height of the highest leaf or as the length of the main axis of non-flowering plants. Regenerating height was measured either as the height of the highest flower on a plant or as the height of the main axis of flowering. Sampled plants were either randomly selected in the core area of plots or along transects in defined distances. For details refer to the description of individual years. Starting in 2006, also the plots of the management experiment, that altered mowing frequency and fertilized subplots (see further details in the general description of the Jena Experiment) were sampled. 2. Species specific plant height was recorded two times in 2002: in late July (vegetative height) and just before biomass harvest during peak standing biomass in late August (vegetative and regenerative height). For each plot and each sown species in the species pool, 3 plant individuals (if present) from the central area of the plots were randomly selected and used to measure vegetative height (non-flowering indviduals) and regenerative height (flowering individuals) as stretched height. Provided are the means over the three measuremnts per plant species per plot.

Plant height along the plant diversity gradient in the Jena Experiment (Main Experiment, year 2002)

This data set contains measurements of plant height: vegetative height (non-flowering indviduals) and regenerative height (flowering individuals) in 2002 from the Main Experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). 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). Plots were maintained by bi-annual weeding and mowing. In 2002, plant height was recorded twice a year: in late June and just before biomass harvest during peak standing biomass in late August. For 3 target plant individuals (if present) per sown species from the central area of the plots, vegetative height (non-flowering indviduals) and regenerative height (flowering individuals) were measured as stretched height. Provided are the indivdiual measurements and the mean over the measured plants per plot (in June) and the mean over the measured plants per plot (in August).

Aboveground community and species-specific plant biomass from the Jena Experiment (Main Experiment, year 2003)

This data set contains aboveground community 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 the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). 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). Plots were maintained by bi-annual weeding and mowing. Aboveground community biomass was harvested twice in 2003 just prior to mowing (during peak standing biomass in late May and in late August) on all experimental plots of the main experiment. This was done by clipping the vegetation at 3 cm above ground in four rectangles of 0.2 x 0.5 m per large 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. the central 10 x 15 m). 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.

Aboveground community and species-specific plant biomass from the Jena Experiment (Dominance Experiment, year 2006)

This data set contains 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 the dominance experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the dominance experiment, 206 grassland plots of 3.5 x 3.5 m were established from a pool of 9 plant 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). Plots were maintained by bi-annual weeding and mowing. Aboveground community biomass was harvested twice in May and August 2006 on all experimental plots of the dominance experiment. This was done by clipping the vegetation at 3 cm above ground in two rectangles of 0.2 x 0.5 m per experimental plot. The location of these rectangles was assigned by random selection of coordinates within the central area of the plots (excluding an outer edge of 50cm). 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, and remaining plant material that could not be assigned to any category. 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 mean of both samples per plot and the individual measurements are provided in the data file. Overall, analyses of the community biomass data have identified species richness and the presence of particular species as an important driver of a positive biodiversity-productivity relationship.

Species-specific plant height along the plant diversity gradient in the Jena Experiment (Main Experiment, year 2002)

This data set contains measurements of species-specific plant height: vegetative height (non-flowering indviduals) and regenerative height (flowering individuals) measured for all sown species separetly in 2002. Data was recorded in the Main Experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). 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). Plots were maintained by bi-annual weeding and mowing. In 2002, plant height was recorded two times: in late July (vegetative height) and just before biomass harvest during peak standing biomass in late August (vegetative and regenerative height). For each plot and each sown species in the species pool, 3 plant individuals (if present) from the central area of the plots were randomly selected and used to measure vegetative height (non-flowering indviduals) and regenerative height (flowering individuals) as stretched height. Provided are the means over the three measuremnts per plant species per plot.

Aboveground community and species-specific plant biomass from the Jena Experiment (Dominance Experiment, year 2003)

This data set contains aboveground community plant biomass (Sown plant community, Weed plant community, and Dead plant material; all measured in biomass as dry weight) and species-specific biomass from the sown species of the dominance experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the dominance experiment, 206 grassland plots of 3.5 x 3.5 m were established from a pool of 9 plant 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). Plots were maintained by bi-annual weeding and mowing. Aboveground community biomass was harvested twice in May and August 2003 on all experimental plots of the dominance experiment. This was done by clipping the vegetation at 3 cm above ground in two rectangles of 0.2 x 0.5 m per experimental plot. The location of these rectangles was assigned by random selection of coordinates within the central area of the plots (excluding an outer edge of 50cm). 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, and remaining plant material that could not be assigned to any category. 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 mean of both samples per plot and the individual measurements are provided in the data file. Overall, analyses of the community biomass data have identified species richness and the presence of particular species as an important driver of a positive biodiversity-productivity relationship.