Nutrient Enrichment and Food Web Composition Affect Ecosystem Metabolism in an Experimental Seagrass Habitat

Background

Food web composition and resource levels can influence ecosystem properties such as productivity and elemental cycles. In particular, herbivores occupy a central place in food webs as the species richness and composition of this trophic level may simultaneously influence the transmission of resource and predator effects to higher and lower trophic levels, respectively. Yet, these interactions are poorly understood.

Methodology/Principal Findings

Using an experimental seagrass mesocosm system, we factorially manipulated water column nutrient concentrations, food chain length, and diversity of crustacean grazers to address two questions: (1) Does food web composition modulate the effects of nutrient enrichment on plant and grazer biomasses and stoichiometry? (2) Do ecosystem fluxes of dissolved oxygen and nutrients more closely reflect above-ground biomass and community structure or sediment processes? Nutrient enrichment and grazer presence generally had strong effects on biomass accumulation, stoichiometry, and ecosystem fluxes, whereas predator effects were weaker or absent. Nutrient enrichment had little effect on producer biomass or net ecosystem production but strongly increased seagrass nutrient content, ecosystem flux rates, and grazer secondary production, suggesting that enhanced production was efficiently transferred from producers to herbivores. Gross ecosystem production (oxygen evolution) correlated positively with above-ground plant biomass, whereas inorganic nutrient fluxes were unrelated to plant or grazer biomasses, suggesting dominance by sediment microbial processes. Finally, grazer richness significantly stabilized ecosystem processes, as predators decreased ecosystem production and respiration only in the zero- and one- species grazer treatments.

Conclusions/Significance

Overall, our results indicate that consumer presence and species composition strongly influence ecosystem responses to nutrient enrichment, and that increasing herbivore diversity can stabilize ecosystem flux rates in the face of perturbations.     

Effects of macrograzers and light on periphyton stoichiometry

Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.     

How enrichment,ecosystem size,and their effects on species richness co‐determine the stability of microcosm communities

Nutrient enrichment, ecosystem size, and richness each may directly affect the stability of both populations and communities. Alternatively, nutrient enrichment and ecosystem size each may directly affect richness, which in turn may affect stability. No previous studies, however, have tested empirically how these three factors interact and co‐determine stability. We manipulated nutrient input and ecosystem size in replicate microcosms containing a diverse bacterial flora, and a range of green algae and heterotrophic protozoa, and used these manipulations and the resulting variation in species richness to measure their combined effects on temporal stability of both populations and communities. Results showed that nutrient enrichment and ecosystem size controlled protist richness, and their effects on stability could be mediated by richness. In addition, both community‐level and population‐level stability increased with protist richness. Furthermore, mean species evenness and mean species richness was negatively related. Effects of statistical averaging, overyielding, and component population stability were identified as possible mechanisms involved explaini ng the stabilizing effects of richness on community stability. Their relative strength in influencing stability, however, is likely to change as mean evenness decreased with increasing richness. This decrease in evenness would tend to weaken the strength of the statistic averaging effect, but increase the strength of the other two mechanisms due to relatively lower population variability (component population stability) and higher mean biovolumes of dominant protists (overyielding).     

How biological diversity influences ecosystem function: a test with a tropical stream detritivore guild

We investigated the relationship between diversity and ecosystem function, which is controversial and has rarely been examined for consumer assemblages, for the process of leaf breakdown by the shredder guild in a tropical stream. We manipulated species richness, evenness and identity of four macroinvertebrate shredder species (three caddisflies and one mayfly) in microcosms and tested their effect on leaf breakdown rates measured as leaf mass loss per capita and per milligram of animal. Species richness, evenness and species identity all affected leaf breakdown rates. Breakdown rates tended to increase with higher richness, but only for the three caddisflies, probably through a release of intraspecific interference, although other mechanisms such as niche complementarity or facilitation cannot be discarded. Leaf breakdown by the caddisflies was reduced in the presence of the mayfly, possibly because of its mode of movement by swimming instead of crawling and its similarity to some predators that are common in leaf litter. Species identity was more important than species richness in determining leaf breakdown rates, indicating that some species within the shredder guild are not redundant, and suggesting important consequences of particular species loss for the functioning of the ecosystem.     

Influence of evenness on the litter-species richness-decomposition relationship in Mediterranean grasslands

Aims Human impacts on natural ecosystems induce changes in their functioning through alterations in species richness, composition and evenness of plant communities. Most litter diversity–decomposition processes studies have only manipulated species richness, ignoring the role of evenness. Here, results from a field litterbag experiment are presented to test whether changes in evenness of species distribution in litter mixtures affected the strength of the litter-species richness–decomposition relationship.Methods Ten herbaceous species abundant in Mediterranean grassland communities and representative of different genera and functional groups were used. Species richness was directly manipulated to produce litter mixtures of three and six plant species, as well as litter of each individual species used. Each level of species richness was replicated several times such that each repeat had a different species composition. Three- and six-species litter mixtures were also treated to vary in evenness (three levels). Decomposition rate was assessed by percentage dry weight loss over the 90 days of the experiment.Important findings Decomposition rate was positively related to the linear increase in litter-species richness and was affected by the composition of the litter-species mixture. Decomposition rates differed significantly between evenness treatments and moreover, the strength of the positive relationship between litter-species richness and decomposition rate decreased notably in the low-evenness treatment. The effects of evenness on decomposition rate, at different richness levels, were partially explained by the differences in the initial litter mixture's carbon-to-nitrogen ratio within them. This study reveals that short-term decomposition rate is positively affected by both components of Mediterranean grassland litter-species diversity.     

Invasion by mobile aquatic consumers enhances secondary production and increases top-down control of lower trophic levels

Increased biological diversity due to invasion by non-indigenous species (NIS) is a global phenomenon with potential effects on trophic interactions and ecosystem processes in the invaded habitat. We assessed the effects of resource availability and invasion of three non-indigenous invertebrate grazers (two crustaceans and a snail) on secondary production, relative dominance of NIS grazers and resource depletion in experimental freshwater mesocosms. The relative dominance of NIS grazers increased with increasing initial resource availability, although the effect was largest for one of the three species. The effect was due to the fact that all the included non-indigenous grazers were able to expand their populations quickly in response to resource addition. For the most dominating species, the increased grazer diversity due to invasion in turn resulted in higher production of grazer biomass and a more efficient depletion of the periphyton resource. The effect was largest at high initial resource availability, where NIS dominance was most pronounced. Our results show that an invasion-induced increase in species diversity can increase resource depletion and consequently production, but that the effect depends on identity of the introduced species. The results also suggest that properties of the recipient system, such as resource availability, can modulate ecosystem effects of NIS by affecting invader success and dominance.     

The independent and interactive effects of snail grazing and nutrient enrichment on structuring periphyton communities

We investigated the independent and interactive effects of nutrient enrichment and snail grazing on structuring periphyton communities in a northern temperate lake. Nutrient releasing substrates and grazer enclosures were used to simultaneously manipulate nutrient availability and herbivory. Periphyton was allowed 18 days to accrue before grazers (Elimia livescens = Goniobasis livescens) were introduced.Addition of nitrogen and phosphorus caused a significant increase in biovolume (p < 0.001), whereas grazing had no significant effect on biovolume but resulted in a shift in species composition. Four taxa were largely responsible for the increase in biovolume on the nutrient enriched substrates: Oedogonium sp, Stigeoclonium tenue, Navicula radiosa var. radiosa and Navicula radiosa var. tenella. By the 28th day, nutrient enrichment caused a shift from a community dominated by diatoms (Bacillariophyceae) to a community dominated by green algae (Chlorophyceae). Blue green algae (Myxophyceae) maintained an equal proportion in high and low-nutrient regimes.Grazing had a more pronounced effect on altering community composition on the nutrient enriched substrates than on the unenriched substrates. Grazing caused a decrease in diversity and an increase in dominance by green algae on the nutrient enriched substrates. The relative biovolume of green algae increased from 64% to 93% on grazed substrates, due to the significant increase in relative abundance of Stigeoclonium tenue. This taxon has both prostate basal cells and erect filamentous cells. The ratio of basal: filamentous cells increased from 4.7 to 5.2 with grazing, suggesting that the heretotrichous growth form of Stigeoclonium tenue is adapted to grazing by virtue of the basal cells which are able to adhere to the substratum and resist being grazed.     

Ecological stoichiometry of indirect grazer effects on periphyton nutrient content

Ecological stoichiometry has been successful in enhancing our understanding of trophic interactions between consumer and prey species. Consumer and prey dynamics have been shown to depend on the nutrient composition of the prey relative to the nutrient demand of the consumer. Since most experiments on this topic used a single consumer species, little is known about the validity of stoichiometric constraints on trophic interactions across consumers and ecosystems. We conducted a quantitative meta-analysis on grazer–periphyton experiments to test (1) if benthic grazers have consistent effects on the nutrient composition of their prey, and (2) whether these effects can be aligned to the nutrient stoichiometry of grazer and periphyton, other environmental factors, or experimental constraints. Grazers significantly lowered periphyton C:N and C:P ratios, indicating higher N- and P-content of grazed periphyton across studies. Grazer presence on average increased periphyton N:P ratios, but across studies the effect size did not differ significantly from zero. The sign and strength of grazer effects on periphyton nutrient ratios was strongly dependent on the nutrient content of grazers and their food, but also on grazer biomass, the amount of biomass removal and water column nutrients. Grazer with low P-content tended to reduce periphyton P-content, whereas grazers with high P-content increased periphyton P-content. This result suggests that low grazer P-content can be an indication of physiological P-limitation rather than a result of having relatively low and fixed P-requirements. At the across-system scale of this meta-analysis, predictions from stoichiometric theory are corroborated, but the plasticity of the consumer nutrient composition has to be acknowledged. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

广西马尾松林植物功能多样性与生产力的关系

探索植物多样性与生产力的关系可为森林经营与管理提供科学基础。本研究以广西4个地区的马尾松(Pinus massoniana)人工林群落为研究对象, 通过计算物种多样性、功能多样性和功能优势值, 运用相关分析、自动线性建模和方差划分等方法, 分析了多样性与生产力的关系。研究发现, 生产力与物种丰富度、Shannon指数、功能丰富度、功能均匀度极显著正相关(P < 0.01), 与物种均匀度、功能多样性、功能离散度、功能团个数、坡向显著正相关(P < 0.05), 与林龄极显著负相关(P < 0.01), 4个功能多样性参数和4个物种多样性参数两两之间皆为显著正相关; 未发现初始生物量制约生产力的提高; 方差划分最优模型中, 功能多样性参数效应、功能优势值效应和林龄效应分别解释生产力方差的56%、43%和33%, 功能多样性参数效应和功能优势值效应重叠部分高达27%; 生态位互补效应主要由功能丰富度和功能均匀度产生, 选择效应主要由生长型优势值产生; 生长型优势值为灌木的样地生产力较高, 次优种或过渡种对生态系统功能也有重要作用。以生产力为响应变量的自动线性建模最佳子集包括重要性由大到小的5个因素: 林龄、生长型优势值、功能丰富度、功能均匀度、功能团个数。建议维护森林功能多样性, 加强林下叶层植物保护, 用好功能重要的物种, 通过林下叶层的补偿性光合作用和生长竞争, 有效地提高生产力和生物多样性。     

Biodiversity effects on productivity and stability of marine macroalgal communities: the role of environmental context

The influence of biodiversity on ecosystem functioning has been the focus of much recent research, but the role of environmental context and the mechanisms by which it may influence diversity effects on production and stability remain poorly understood. We assembled marine macroalgal communities in two mesocosm experiments that varied nutrient supply, and at four field sites that differed naturally in environmental conditions. Concordant with theory, nutrient addition promoted positive species richness effects on algal growth in the first mesocosm experiment; however, it tended to weaken the positive diversity relationship found under ambient conditions in a second experiment the next year. In the field experiments, species richness increased algal biomass production at two of four sites. Together, these experiments indicate that diversity effects on algal biomass production are strongly influenced by environmental conditions that vary over space and time. In decomposing the net biodiversity effect into its component mechanisms, seven of the eight experimental settings showed positive complementarity effects (suggesting facilitation or complementary resource use) countered by negative selection effects (i.e. enhanced growth in mixture of otherwise slow growing species) to varying degrees. Under no conditions, including nutrient enrichment, did we find evidence of positive selection effects commonly thought to drive positive diversity effects. Species richness enhanced stability of algal community biomass across a range of environmental settings in our field experiments. Hence, while species richness can increase production, enhanced stability is also an important functional outcome of maintaining diverse marine macroalgal communities.     

Biodiversity influences ecosystem functioning in aquatic angiosperm communities

Knowledge of the connection between aquatic plant diversity and ecosystem processes is still limited. To examine how plant species diversity affects primary productivity, plant nutrient use, functional diversity of secondary producers and population/community stability, we manipulated submerged angiosperm species diversity in a field experiment lasting 15 weeks. Plant richness increased the shoot density for three of four species. Polyculture biomass production was enhanced by increasing richness, with positive complementarity and selection effects causing positive biodiversity effects. Species richness enhanced the community stability for biomass production and shoot density. Sediment ammonium availability decreased with plant diversity, suggesting improved nutrient usage with increasing plant richness. Interestingly, positive multitrophic effects of plant species richness on structural and functional diversity of macrobenthic secondary producers were recorded. The results suggest that mixed seagrass meadows play an important role for ecosystem functioning and thus contribute to the provision of goods and services in coastal areas.     

Consistent Effects of Biodiversity on Ecosystem Functioning Under Varying Density and Evenness

Biodiversity experiments typically vary only species richness and composition, yet the generality of their results relies on consistent effects of these factors even under varying starting conditions of density and evenness. We tested this assumption in a factorial species richness x density x evenness experiment using a pool of 60 common grassland species divided into four functional groups (grasses, legumes, tall herbs and short herbs). Richness varied from 1, 2, 4, 8 to 16 species, total planting density was 1,000 or 2,000 seeds/m², and species were sown in even or uneven proportions, where one functional group was made dominant. Aboveground plant biomass increased linearly with the logarithm of species richness in all density and evenness treatments during all three years of the experiment. This was due to a convergence of realized density and evenness within species richness levels, although functional groups which were initially made dominant retained their dominance. Between species richness levels, realized density increased, and realized evenness decreased with species richness. Thus, more individuals could coexist if they belonged to different species. Within species richness levels, higher biomass values were correlated with lower density, suggesting an underlying thinning process. However, communities with low realized evenness also had low biomass values; thus high biomass did not result from species dominance. So-called complementarity and selection effects were similar across density and evenness treatments, indicating that the mechanisms underpinning the biodiversity effects were not altered. Species richness was the dominant driver of aboveground biomass, irrespective of variations in total densities and species abundance distributions at the start of the experiment; rejecting the hypothesis that initial differences in species abundance distributions might lead to different “stable states” in community structure or biomass. Thus, results from previous biodiversity experiments that only manipulated species richness and composition should be quite robust and broadly generalizable.     

Species Diversity of Periphyton Communities in the Littoral of a Temperate Lake

Species diversity of littoral periphyton sampled from Elk Lake (B. C., Canada) was dependent on station location and period of substrate exposure. Site differences were not statistically related to physico-chemical variables. However, diversity was negatively correlated with age of the communities and standing crop, and appeared to be associated with biological interaction or competition for available substrate surface. Further investigation revealed that the Shannon-Weaver index, as commonly applied, was inappropriate for data summarization since it reflected only the changes in evenness of the most abundant species. Components of the diversity index, as illustrated by the periphyton, were discussed.     

Ecological responses of aquatic macrophytes and benthic macroinvertebrates to dams in the Henares River Basin (Central Spain)

The ecological responses of aquatic macrophytes and benthic macroinvertebrates to deep-release dams in three impounded rivers of the Henares River Basin (Central Spain) were studied, specially focusing on the effects of nutrient enrichment caused by deep releases on these two freshwater communities. Three sampling sites, one upstream and two downstream from the reservoir, were established in each impounded river. Sampling surveys to collect submersed macrophytes and benthic macroinvertebrates at each sampling site were carried out in spring–summer of 2009 and 2011. Water temperature tended to decrease downstream from dams, whereas nitrate and phosphate concentrations tended to increase. These abiotic changes, particularly the downstream nutrient enrichment, apparently affected the macrophyte and macroinvertebrate communities. In the case of submersed macrophytes, total coverage and taxa richness increased downstream from dams. In the case of benthic macroinvertebrates, total density and total biomass also increased downstream, but taxa richness tended to decrease. Scrapers appeared to be the macroinvertebrate feeding group most favored downstream from dams as a probable consequence of the positive effect of nutrient enrichment on periphyton and perilithon abundance. Nutrients would ultimately come from water runoff over agricultural lands and over semi-natural forests and pastures, being subsequently accumulated in the hypolimnion of reservoirs.     

Patterns of species diversity and soil nutrients along a chronosequence of vegetation recovery in Hainan Island, South China

Understanding the pattern of species diversity and soil factors can enhance our knowledge of the mechanism of vegetation recovery, however, there is still a gap in the knowledge of succession rate and trend for species diversity in relation to soil nutrients during the vegetation recovery process. Patterns of species diversity and soil nutrients during the tropical vegetation recovery as well as the correlation between species diversity and soil nutrients were explored in Hainan Island, located in southern China. Plots assigned as grassland stage (GS), shrub stage (SS), secondary forest stage (SFS), and primary forest stage (PFS) were established using a chronosequence approach. Results showed that species richness and evenness increased from GS to PFS. Species dominance/diversity curves were fitted using the lognormal distribution model (r ² ?=?0.891?C0.972). Species richness for the herb layer was maximal at SFS, whereas species richness for both the shrub layer and tree layer reached their maximum at PFS. Species turnover and soil total phosphorus decreased, whereas organic matter and total nitrogen increased from GS to PFS. Organic matter and total nitrogen were both positively correlated with species richness and total coverage, and total phosphorus was positively correlated with species turnover. The results clearly demonstrate that diversity asymptotically increases and positively correlates with increasing soil fertility, and the total phosphorus value is predicted to be an important soil factor that affects successional rate during tropical vegetation recovery processes.     

Species richness and evenness respond in a different manner to propagule density in developing prairie microcosm communities

Diversity has two basic components: richness, or number of species in a given area, and evenness, or how relative abundance or biomass is distributed among species. Previously, we found that richness and evenness can be negatively related across plant communities and that evenness can account for more variation in Shannon’s diversity index (H′) than richness, which suggests that relationships among diversity components can be complex. Non-positive relationships between evenness and richness could arise due to the effects of migration rate or local species interactions, and relationships could vary depending on how these two processes structure local communities. Here we test whether diversity components are equally or differentially affected over time by changes in seed density (and associated effects on established plant density and competition) in greenhouse communities during the very early stages of community establishment. In our greenhouse experiment, we seeded prairie microcosms filled with bare field soil at three densities with draws from a mix of 22 grass and forb species to test if increased competition intensity or seedling density would affect the relationships among diversity components during early community establishment. Increased seed density treatments caused diversity components to respond in a different manner and to have different relationships with time. Richness increased linearly with seed density early in the experiment when seedling emergence was high, but was unrelated to density later in the experiment. Evenness decreased log-linearly with seed densities on all sampling dates due to a greater dominance by Rudbeckia hirta with higher densities. Early in the experiment, diversity indices weakly reflected differences in richness, but later, after the competitive effects of Rudbeckia hirta became more intense, diversity indices more strongly reflected differences in evenness. This suggests that species evenness and diversity indices do not always positively covary with richness. Based on these results, we suggest that evenness and richness can be influenced by different processes, with richness being more influenced by the number of emerging seedlings and evenness more by species interactions like competition. These results suggest that both diversity components should be measured in plant diversity studies whenever it is possible.     

营养盐加富和鱼类添加对浮游植物群落演替和多样性的影响

浮游植物是水体生态系统敞水区最重要的初级生产者,其组成与多样性反映了群落的结构类型和存在状态。通过围隔实验,模拟水库春季发生的营养盐加富和鱼类放养的干扰,分析在这两种干扰下的浮游植物群落演替过程中优势种和稀有种的变化,并通过以丰度与生物量为变量的香农和辛普森多样性指数的计算,分析浮游植物群落演替过程中的多样性变化特征。结果表明,营养盐加富干扰下的浮游植物群落的优势种变化和演替更为明显,营养盐加富与鱼类添加对浮游植物群落多样性变化的影响符合中度干扰理论。在优势种优势度变化较大的浮游植物群落演替过程中,多样性指数与浮游植物生物量有较高的负相关性。在浮游植物群落演替过程中,香农和辛普森多样性指数的变化趋势基本一致,采用丰度与生物量为变量的两种多样性指数的计算结果对实验系统中浮游植物群落多样性的分析结果没有明显的影响。     

Species richness vs evenness: independent relationship and different responses to edaphic factors

The main components of species diversity are richness and evenness. Species richness has been the subject of biodiversity studies more often than species evenness. Some simulated models suggest that the relationship between these two components is equal and predictable in a stable environment. This study was conducted in a human-shaped and unstable environment in arable field boundaries. A total of 30 sampling plots were laid out at random in the boundaries. Vegetation at the sites was of meadow type. The relationship between species richness and evenness, and their responses to edaphic nutrient factors in the field boundaries, were examined by correlation analysis and an ordination method. Correlation analyses demonstrated that no consistent pattern was present in the relationship between species evenness and species richness in this human-shaped ecosystem. Species richness and evenness also had different responses to edaphic factors, with species richness being negatively correlated with phosphorus, and species evenness negatively correlated with the ratio of organic carbon to total nitrogen in soil. The results indicate that different and independent ecological processes determine species richness and evenness. The relationship between these two components may be site-specific, reflecting variation in resource utilization by plant species. The pattern thus reflects the spatial heterogeneity of disturbances or 'patchiness' of resources between sites in a semi-natural ecosystem.     

Diversity and ecological distribution of macrofungi in the Laojun Mountain region,southwestern China

Surveys of the macrofungi associated with eight different vegetation types in the Laojun Mountain region of southwestern China yielded approximately 520 species belonging to 175 genera. Species richness and diversity were highest in mixed conifer and broadleaf forests and deciduous broadleaf forests. In typical forests of temperate regions of the world, there are five dominant genera of ectomycorrhizal macrofungi. The distribution patterns of species in these genera for the different vegetation types indicate that they are able to associate with a wide variety of different trees. Analysis of data for common macrofungal species and taxonomic similarity of the communities present in the eight vegetation types suggest that the greater the differences in the plant species that comprise the vegetation, the less similar are the common macrofungal species associated with the most common host plants. These same data also show that some species of macrofungi occur only in one or two vegetation types. There were 156 species of edible fungi recorded from the different vegetation types, and these fungi appear to be abundant in the Laojun Mountain region. At different positions along the elevation gradient, positive correlations existed with respect to the relationship between species richness and diversity, and the general trend was for macrofungal species richness and diversity to decrease with increasing elevation, with the numbers of species recorded being significantly lower at the very highest elevation. The relative dominance of certain taxa in the assemblage of species present was found to increase with increasing elevation, and variation in the evenness component of diversity was slight. As such, the differences in dominance and evenness were also not significant (P > 0.05). Macrofungal species richness was slightly more diverse on shaded slopes than on more exposed (sunny) slopes, and the differences in species diversity, dominance and evenness were relatively minor. This suggests that slope aspect may only weakly influence the distribution of macrofungal species in the Laojun Mountain region.