[show abstract][hide abstract] ABSTRACT: Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally-replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance varied most among regions (subcontinents), whereas cover was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds mono-dominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions. This article is protected by copyright. All rights reserved.
Global Change Biology 08/2013; · 6.91 Impact Factor
[show abstract][hide abstract] ABSTRACT: Large-scale exploitation of higher trophic levels by humans, together with global-scale nutrient enrichment, highlights the need to explore interactions between predator loss and resource availability. The hypothesis of exploitation ecosystems suggests that top–down and bottom–up control alternate between trophic levels, resulting in a positive relationship between primary production and the abundance of every second trophic level. Specifically, in food webs with three effective trophic levels, primary producers and predators should increase with primary production, while in food webs with two trophic levels, only herbivores should increase. We provided short-term experimental support for these model predictions in a natural benthic community with three effective trophic levels, where the number of algal recruits, but not the biomass of gastropod grazers, increased with algal production. In contrast, when the food web was reduced to two trophic levels by removing larger predators, the number of algal recruits was unchanged while gastropod grazer biomass increased with algal production. Predator removal only affected the consumer-controlled early life-stages of algae, indicating that both the number of trophic levels and the life-stage development of the producer trophic level determine the propagation of trophic cascades in benthic systems. Our results support the hypothesis that predators interact with resource availability to determine food-web structure.
Ecological Research 08/2013; 27(1). · 1.55 Impact Factor
[show abstract][hide abstract] ABSTRACT: The information on temperature-mediated changes in biodiversity in local assemblages is scarce and mainly addresses the change in species richness. However, warming may have more consistent effects on species turnover than on the number of species. Moreover, very few studies extended the analysis of changes in biodiversity and species composition to questions of associated ecosystem functions such as primary production. Here, we synthesize 4 case studies employing microalgal microcosms within the Aquashift priority program to ask (1) do warming-related shifts in species richness correspond to changes in the rate of biomass production, (2) do similar relationships prevail for evenness, and (3) do warming-related shifts in species turnover stabilize or destabilize biomass production? Two of the four cases are previously unpublished, and for a third case, the link between diversity and functional consequences of temperature was not analyzed before. We found accelerated loss of species with warming in all cases. Biomass production was lower with lower species richness in most cases but increased with lower evenness. Most importantly, the relation between functional and compositional stability was different between cases: More rapid extinction resulted in more variable biomass in 2 cases conducted with a limited species pool, indicating that compositional destabilization relates to functional variability. By contrast, the only experiment with a large species pool (30 species) allowed previously rare species to become dominant in the community and showed more stable biomass at high turnover, indicating that compensatory dynamics (turnover) can promote functional stability. These 4 independent experiments highlight the need to consider both compositional and functional consequences of altered temperature regimes.
[show abstract][hide abstract] ABSTRACT: The linkages between biological communities and ecosystem function remain poorly understood along gradients of human-induced stressors. We examined how resource provisioning (nutrient recycling), mediated by native freshwater mussels, influences the structure and function of benthic communities by combining observational data and a field experiment. We compared the following: (1) elemental and community composition (algal pigments and macroinvertebates) on live mussel shells and on nearby rocks across a gradient of catchment agriculture and (2) experimental colonisation of benthic communities on live vs. sham shells controlling for initial community composition and colonisation duration. We show that in near pristine systems, nutrient heterogeneity mediated by mussels relates to greater biodiversity of communities, which supports the notion that resource heterogeneity can foster biological diversity. However, with increased nutrients from the catchment, the relevance of mussel-provisioned nutrients was nearly eliminated. While species can persist in disturbed systems, their functional relevance may be diminished or lost.
[show abstract][hide abstract] ABSTRACT: In their seminal paper, Goldman et al. suggested that phytoplankton close to maximum growth rate attains a
restricted optimal N: P ratio close to the Redfield ratio of molar N: P 5 16. Recently, the presence of such a
global attractor for optimal phytoplankton stoichiometry has been questioned in models and empirical analyses.
As the chemical composition of phytoplankton is of major importance for our understanding of global elemental cycles and biogeochemical transformations, we assembled 55 data sets of phytoplankton growth rate and biomass N: P ratios in a meta-analysis testing (1) whether phytoplankton N: P converges at high growth rates, (2) whether N: P ratios scale with growth rate, and (3) whether the optimal N: P ratios achieved at highest growth rates reflect organism traits or environmental conditions. Across systems and species, phytoplankton N: P decreased with increasing growth rate and at the same time showed decreasing variance, i.e., fast-growing phytoplankton is more P rich and has a more confined elemental composition. Optimal N: P increased with increasing N: P of available nutrients, i.e., with increasing P limitation. Other differences were rare, except cyanobacteria showed higher optimal N: P than diatoms. Understanding the role of phytoplankton in biogeochemical transformation requires modeling approaches that are stoichiometrically flexible to reflect the dynamics of growth and nutrient supply in primary producers.
Limnology and oceanography 05/2013; 58:2076-2088. · 3.41 Impact Factor
[show abstract][hide abstract] ABSTRACT: The nutrition of animal consumers is an important regulator of ecological processes due to its effects on their physiology, life-history and behaviour. Understanding the ecological effects of poor nutrition depends on correctly diagnosing the nature and strength of nutritional limitation. Despite the need to assess nutritional limitation, current approaches to delineating nutritional constraints can be non-specific and imprecise. Here, we consider the need and potential to develop new complementary approaches to the study of nutritional constraints on animal consumers by studying and using a suite of established and emerging biochemical and molecular responses. These nutritional indicators include gene expression, transcript regulators, protein profiling and activity, and gross biochemical and elemental composition. The potential applications of nutritional indicators to ecological studies are highlighted to demonstrate the value that this approach would have to future studies in community and ecosystem ecology.
[show abstract][hide abstract] ABSTRACT: Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade-off possibilities. The competition-defence hypothesis posits a trade-off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth-defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth-defence trade-off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life-history groups and within the majority of individual sites. Thus, a growth-defence trade-off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.
[show abstract][hide abstract] ABSTRACT: In the course of the biodiversity-ecosystem functioning debate, the issue of multifunctionality of species communities has recently become a major focus. Elemental stoichiometry is related to a variety of processes reflecting multiple plant responses to the biotic and abiotic environment. It can thus be expected that the diversity of a plant assemblage alters community level plant tissue chemistry. We explored elemental stoichiometry in aboveground plant tissue (ratios of carbon, nitrogen, phosphorus, and potassium) and its relationship to plant diversity in a 5-year study in a large grassland biodiversity experiment (Jena Experiment). Species richness and functional group richness affected community stoichiometry, especially by increasing C:P and N:P ratios. The primacy of either species or functional group richness effects depended on the sequence of testing these terms, indicating that both aspects of richness were congruent and complementary to expected strong effects of legume presence and grass presence on plant chemical composition. Legumes and grasses had antagonistic effects on C:N (-27.7% in the presence of legumes, +32.7% in the presence of grasses). In addition to diversity effects on mean ratios, higher species richness consistently decreased the variance of chemical composition for all elemental ratios. The diversity effects on plant stoichiometry has several non-exclusive explanations: The reduction in variance can reflect a statistical averaging effect of species with different chemical composition or a optimization of nutrient uptake at high diversity, leading to converging ratios at high diversity. The shifts in mean ratios potentially reflect higher allocation to stem tissue as plants grew taller at higher richness. By showing a first link between plant diversity and stoichiometry in a multiyear experiment, our results indicate that losing plant species from grassland ecosystems will lead to less reliable chemical composition of forage for herbivorous consumers and belowground litter input.
PLoS ONE 01/2013; 8(3):e58179. · 3.73 Impact Factor
[show abstract][hide abstract] ABSTRACT: ABSTRACT: Two decades of biodiversity-ecosystem functioning research have not yet led to consensus view on how local dominance and biodiversity affect the stability of ecosystems in the presence of disturbances. As natural systems are usually affected by multiple stressors, we tested the diversity−stability hypothesis on phytoplankton communities subjected to combinations of 2 stressors (pH reduction and simulated grazing). Monocultures revealed uncorrelated species’ sensitivities to the 2 stressors and the presence of a best-performing species (Scenedesmus obliquus). We thus selected 2 experimental species combinations, either (A) including or (B) excluding S. obliquus, and inoculated at 3 diversity levels (2, 4, 8 species). Both stressors induced significant biovolume loss and increased temporal biovolume variation, but the effect on final biomass (Btot) differed between combinations and with richness. The fast recovery of S. obliquus precluded long-term disturbance effects on Btot in Combination A. Correspondingly, these communities showed slower recovery rates and lower Btot with more species present, since the strong performance of S. obliquus was diluted by more species being present. More species-rich assemblages in Combination B lost more biovolume after stressor application, whereas the recovery rates showed a more complex pattern, depending on stressor sequence and species richness. Our study suggests that the stability of communities in the face of multiple disturbances cannot be predicted from a general diversity−stability relationship alone, but depends on the correlation between species sensitivities to different stressors and on the population dynamics of a well-performing species when dominant.
[show abstract][hide abstract] ABSTRACT: The dependence of competitive interactions on abiotic conditions is attracting increasing interest in the face of globally
rising temperatures and altered biogeochemical cycles of major nutrients. In a microcosm experiment involving a natural inoculum
of benthic microalgae, temperature and nutrient supply ratios were manipulated in order to test three main hypotheses: (1)
temperature and nutrient supply ratios determine species composition and diversity of the assemblage, (2) the identity of
the dominating species depends on nutrient supply and temperature, and (3) higher temperature leads to faster competitive
exclusion and thus more rapid decline in species richness. Over a period of 7weeks, algal biomass reached an equilibrium
carrying capacity, with was higher at colder temperatures and intermediate N:P supply ratios (N:P=16). Initial growth rate
increased with temperature and under high P-supply. Species richness in the stationary phase of the experiment decreased with
increasing temperature, reflecting a higher extinction rate in the warmer treatments, which were also characterized by higher
dominance of single species. Thus, increasing temperature both altered the identity of the dominating species and accelerated
competitive displacement. This experiment thus indicates that warming might influence outcome and temporal dynamics in species
interactions, and thereby eventually local diversity.
KeywordsExtinction–Species interactions–Warming–Species richness–Dominance–Microphytobenthos
Ecological Research 04/2012; 26(3):533-539. · 1.55 Impact Factor
[show abstract][hide abstract] ABSTRACT: The evidence for species diversity effects on ecosystem functions is mainly based on studies not explicitly addressing local
or regional processes regulating coexistence or the importance of community structure in terms of species evenness. In experimental
communities of marine benthic microalgae, we altered the successional stages and thus the strength of local species interactions
by manipulating rates of dispersal and disturbance. The treatments altered realized species richness, evenness and community
biomass. For species richness, dispersal mattered only at high disturbance rates; when opening new space, dispersal led to
maximized richness at intermediate dispersal rates. Evenness, in contrast, decreased with dispersal at low or no disturbance,
i.e. at late successional stages. Community biomass showed a non-linear hump-shaped response to increasing dispersal at all
disturbance levels. We found a positive correlation between richness and biomass at early succession, and a strong negative
correlation between evenness and biomass at late succession. In early succession both community biomass and richness depend
directly on dispersal from the regional pool, whereas the late successional pattern shows that if interactions allow the most
productive species to become dominant, diverting resources from this species (i.e. higher evenness) reduces production. Our
study emphasizes the difference in biodiversity–function relationships over time, as different mechanisms contribute to the
regulation of richness and evenness in early and late successional stages.
KeywordsBiodiversity ecosystem functioning-Richness-Evenness-Dispersal-Disturbance-Successional stage
[show abstract][hide abstract] ABSTRACT: Pan et al. claim that our results actually support a strong linear positive relationship between productivity and richness, whereas Fridley et al. contend that the data support a strong humped relationship. These responses illustrate how preoccupation with bivariate patterns distracts from a deeper understanding of the multivariate mechanisms that control these important ecosystem properties.
[show abstract][hide abstract] ABSTRACT: Food webs may be affected by evolutionary processes, and effective evolutionary time ultimately affects the probability of species evolving to fill the niche space. Thus, ecosystem history may set important evolutionary constraints on community composition and food web structure. Food chain length (FCL) has long been recognized as a fundamental ecosystem attribute. We examined historical effects on FCL in large lakes spanning >6 orders of magnitude in age. We found that food chains in the world's ancient lakes (n = 8) were significantly shorter than in recently formed lakes (n = 10) and reservoirs (n = 3), despite the fact that ancient lakes harbored much higher species richness, including many endemic species. One potential factor leading to shorter FCL in ancient lakes is an increasing diversity of trophic omnivores and herbivores. Speciation could simply broaden the number of species within a trophic group, particularly at lower trophic levels and could also lead to a greater degree of trophic omnivory. Our results highlight a counter-intuitive and poorly-understood role of evolutionary history in shaping key food web properties such as FCL.
PLoS ONE 01/2012; 7(6):e37856. · 3.73 Impact Factor
[show abstract][hide abstract] ABSTRACT: Biological diversity comprises both species richness, i.e., the number of species in a community, and evenness, measuring how similar species are in their abundances. The relationship between species richness and evenness (RRE) across communities remains, however, a controversial issue in ecology because no consistent pattern has been reported. We conducted a systematic meta-review of RRE in aquatic ecosystems along regional to continental gradients and across trophic groups, differing in body size by 13 orders of magnitude. Hypotheses that RRE responded to latitudinal and scale variability across trophic groups were tested by regression analyses. Significant correlations of species richness and evenness only existed in 71 out of 229 datasets. Among the RRE, 89 were negative and 140 were positive. RRE did not vary with latitude but showed a positive response to scale. In a meta-analysis with ecosystem type as a single explaining variable, RRE did not vary among ecosystem types, i.e. between marine and freshwater. Finally, autotrophs had more positive RRE than heterotrophs. The weak RRE in many aquatic datasets suggests that richness and evenness often reflect independent components of biodiversity, highlighting that richness alone may be an incomplete surrogate for biodiversity. Our results further elucidate that RRE is driven by organismal and environmental properties, both of which must be considered to gain a deeper understanding of large-scale patterns of biodiversity.
[show abstract][hide abstract] ABSTRACT: More diverse communities have been shown to have higher and more temporally stable ecosystem functioning than less diverse ones, suggesting they should also have a consistently higher level of functioning over time. Diverse communities could maintain consistently high function because the species driving function change over time (functional turnover) or because they are more likely to contain key species with temporally stable functioning. Across 7 y in a large biodiversity experiment, we show that more diverse plant communities had consistently higher productivity, that is, a higher level of functioning over time. We identify the mechanism for this as turnover in the species driving biomass production; this was substantial, and species that were rare in some years became dominant and drove function in other years. Such high turnover allowed functionally more diverse communities to maintain high biomass over time and was associated with higher levels of complementarity effects in these communities. In contrast, turnover in communities composed of functionally similar species did not promote high biomass production over time. Thus, turnover in species promotes consistently high ecosystem function when it sustains functionally complementary interactions between species. Our results strongly reinforce the argument for conservation of high biodiversity.
Proceedings of the National Academy of Sciences 09/2011; 108(41):17034-9. · 9.74 Impact Factor
[show abstract][hide abstract] ABSTRACT: For more than 30 years, the relationship between net primary productivity and species richness has generated intense debate in ecology about the processes regulating local diversity. The original view, which is still widely accepted, holds that the relationship is hump-shaped, with richness first rising and then declining with increasing productivity. Although recent meta-analyses questioned the generality of hump-shaped patterns, these syntheses have been criticized for failing to account for methodological differences among studies. We addressed such concerns by conducting standardized sampling in 48 herbaceous-dominated plant communities on five continents. We found no clear relationship between productivity and fine-scale (meters(-2)) richness within sites, within regions, or across the globe. Ecologists should focus on fresh, mechanistic approaches to understanding the multivariate links between productivity and richness.
[show abstract][hide abstract] ABSTRACT: Synergistic interactions between multiple limiting resources are common, highlighting the importance of co-limitation as a constraint on primary production. Our concept of resource limitation has shifted over the past two decades from an earlier paradigm of single-resource limitation towards concepts of co-limitation by multiple resources, which are predicted by various theories. Herein, we summarise multiple-resource limitation responses in plant communities using a dataset of 641 studies that applied factorial addition of nitrogen (N) and phosphorus (P) in freshwater, marine and terrestrial systems. We found that more than half of the studies displayed some type of synergistic response to N and P addition. We found support for strict definitions of co-limitation in 28% of the studies: i.e. community biomass responded to only combined N and P addition, or to both N and P when added separately. Our results highlight the importance of interactions between N and P in regulating primary producer community biomass and point to the need for future studies that address the multiple mechanisms that could lead to different types of co-limitation.
[show abstract][hide abstract] ABSTRACT: Recent theory suggests that both biodiversity and productivity are constrained by resource supply rates and ratios and that resource stoichiometry is the key to understanding the relationship between biodiversity and productivity. We experimentally tested this theory using pelagic metacommunities. We amended existing predictions by explicitly considering evenness as an aspect of biodiversity and including control of algal biomass by consumption in addition to competition. The metacommunities received a different phosphorus (P) supply and the three patches within each metacommunity differed in their nitrogen (N) supply, which created different N∶P ratios (2, 16, and 128). All patches were inoculated with a phytoplankton assemblage consisting of five species, and half of the metacommunities received two ciliate species as consumers. At the level of the entire metacommunity, algal biomass increased with increasing P supply, whereas species richness and evenness decreased with increasing P supply. Without consumers, resource use efficiency (RUE; realized biomass per unit of P) increased with increasing richness and evenness. Consumer presence reduced overall biomass and richness and precluded a correlation between RUE and biodiversity. At the patch level, local evenness correlated with higher RUE at both imbalanced N∶P ratios (2 and 128) but not at a balanced N∶P ratio. In conclusion, overall P supply constrained realized biomass and altered diversity, whereas resource stoichiometry shaped the relationship between biodiversity and RUE.
The American Naturalist 08/2011; 178(2):171-81. · 4.55 Impact Factor
[show abstract][hide abstract] ABSTRACT: 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.
[show abstract][hide abstract] ABSTRACT: Presence of habitat-forming macroalgae is supposed to mitigate effects of altered resources on benthic microalgae and their consumers. In a field experiment in a microtidal area of the Western Baltic Sea, we tested the interactive effects of nutrient enrichment, artificial shading, and habitat complexity on microalgal biomass and diversity as well as invertebrate abundance and richness. Habitat complexity comprised three levels, the presence of macroalgal canopy of Fucus vesiculosus, the presence of macroalgal propagules, and the absence of both (=control). Microalgal biomass (and richness) was significantly reduced by canopy presence (−88%, compared to control) and shading (−42%), with the highest biomass in the absence of both canopy and macroalgal propagules at ambient light. Within the microalgal assemblage, higher biomass was related to lower evenness (higher dominance). Density of two main invertebrate groups (snails and amphipods) strongly increased with canopy presence (on average from 53 to 154 individuals m−2 stone area for snails, and from 234 to 1203 individuals m−2 for amphipods) and so did invertebrate richness (from 4.3 to 10.3). Additionally, snail density doubled with increasing light availability. The snail responses to light and canopy were independent, the former relating to higher availability of microalgal prey, the latter to more structure. Microalgal taxon richness and biomass decreased with increasing invertebrate richness and with density of snails and amphipods. Our experiment thus showed that the presence of habitat-forming macroalgae alters biomass and diversity across trophic levels in benthic coastal communities as well as their response to resource manipulations.