Positive biodiversity–ecosystem functioning (BEF) relationships observed in experiments can be challenging to identify in natural communities. Freshwater animal communities are disproportionately harmed by global change that results in accelerated species loss. Understanding how animal-mediated ecosystems functions may change as a result of global change can help determine whether biodiversity or species-specific conservation will be effective at maintaining function. Unionid mussels represent half of imperiled species in freshwater ecosystems globally and perform important ecological functions such as water filtration and nutrient recycling. We explored BEF relationships for 22 naturally assembled mussel aggregations spanning three river basins. We used the Price equation to partition the contributions of species richness, composition, and context dependent interactions to two functions of interests: spatially-explicit standing-stock biomass (indirect proxy for function) and species-specific nitrogen (N) excretion rates (direct measure of N recycling). Random and non-random species loss each reduced biomass and N recycling. Many rare species with low contributions to biomass contributed to standing-stock biomass in all basins. Widespread species had variable function across sites, such that context dependent effects (CDEs) outweighed richness effects on standing-stock biomass in two basins, and were similar to richness effects in the third. Richness effects outweighed CDEs for N recycling. Thus, many species contributed a low proportion to overall N-recycling; a product we attribute to the high evenness and functional effect trait diversity associated with these communities. The loss of low-functioning species reduced the function of persisting species. This novel result using observational data adds evidence that positive species interactions, such as interspecific facilitation, may be a mechanism by which biodiversity enhances ecosystem functions. Our work stresses the importance of evaluating species-specific contributions to functions in diverse systems, such as nutrient cycling when maintaining specific animal-mediated functions is a management goal because indirect proxies may not completely characterize BEF relationships.
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Evenness effects mask richness effects on ecosystem functioning at macro‐scales in lakes
Biodiversity–ecosystem functioning (BEF) theory has largely focused on species richness, although studies have demonstrated that evenness may have stronger effects. While theory and numerous small‐scale studies support positive BEF relationships, regional studies have documented negative effects of evenness on ecosystem functioning. We analysed a lake dataset spanning the continental US to evaluate whether strong evenness effects are common at broad spatial scales and if BEF relationships are similar across diverse regions and trophic levels. At the continental scale, phytoplankton evenness explained more variance in phytoplankton and zooplankton resource use efficiency (RUE; ratio of biomass to resources) than richness. For individual regions, slopes of phytoplankton evenness–RUE relationships were consistently negative and positive for phytoplankton and zooplankton RUE, respectively, and most slopes did not significantly differ among regions. Findings suggest that negative evenness effects may be more common than previously documented and are not exceptions restricted to highly disturbed systems.
more » « less- NSF-PAR ID:
- 10457827
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Ecology Letters
- Volume:
- 22
- Issue:
- 12
- ISSN:
- 1461-023X
- Page Range / eLocation ID:
- p. 2120-2129
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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BACKGROUND The availability of nitrogen (N) to plants and microbes has a major influence on the structure and function of ecosystems. Because N is an essential component of plant proteins, low N availability constrains the growth of plants and herbivores. To increase N availability, humans apply large amounts of fertilizer to agricultural systems. Losses from these systems, combined with atmospheric deposition of fossil fuel combustion products, introduce copious quantities of reactive N into ecosystems. The negative consequences of these anthropogenic N inputs—such as ecosystem eutrophication and reductions in terrestrial and aquatic biodiversity—are well documented. Yet although N availability is increasing in many locations, reactive N inputs are not evenly distributed globally. Furthermore, experiments and theory also suggest that global change factors such as elevated atmospheric CO 2 , rising temperatures, and altered precipitation and disturbance regimes can reduce the availability of N to plants and microbes in many terrestrial ecosystems. This can occur through increases in biotic demand for N or reductions in its supply to organisms. Reductions in N availability can be observed via several metrics, including lowered nitrogen concentrations ([N]) and isotope ratios (δ 15 N) in plant tissue, reduced rates of N mineralization, and reduced terrestrial N export to aquatic systems. However, a comprehensive synthesis of N availability metrics, outside of experimental settings and capable of revealing large-scale trends, has not yet been carried out. ADVANCES A growing body of observations confirms that N availability is declining in many nonagricultural ecosystems worldwide. Studies have demonstrated declining wood δ 15 N in forests across the continental US, declining foliar [N] in European forests, declining foliar [N] and δ 15 N in North American grasslands, and declining [N] in pollen from the US and southern Canada. This evidence is consistent with observed global-scale declines in foliar δ 15 N and [N] since 1980. Long-term monitoring of soil-based N availability indicators in unmanipulated systems is rare. However, forest studies in the northeast US have demonstrated decades-long decreases in soil N cycling and N exports to air and water, even in the face of elevated atmospheric N deposition. Collectively, these studies suggest a sustained decline in N availability across a range of terrestrial ecosystems, dating at least as far back as the early 20th century. Elevated atmospheric CO 2 levels are likely a main driver of declines in N availability. Terrestrial plants are now uniformly exposed to ~50% more of this essential resource than they were just 150 years ago, and experimentally exposing plants to elevated CO 2 often reduces foliar [N] as well as plant-available soil N. In addition, globally-rising temperatures may raise soil N supply in some systems but may also increase N losses and lead to lower foliar [N]. Changes in other ecosystem drivers—such as local climate patterns, N deposition rates, and disturbance regimes—individually affect smaller areas but may have important cumulative effects on global N availability. OUTLOOK Given the importance of N to ecosystem functioning, a decline in available N is likely to have far-reaching consequences. Reduced N availability likely constrains the response of plants to elevated CO 2 and the ability of ecosystems to sequester carbon. Because herbivore growth and reproduction scale with protein intake, declining foliar [N] may be contributing to widely reported declines in insect populations and may be negatively affecting the growth of grazing livestock and herbivorous wild mammals. Spatial and temporal patterns in N availability are not yet fully understood, particularly outside of Europe and North America. Developments in remote sensing, accompanied by additional historical reconstructions of N availability from tree rings, herbarium specimens, and sediments, will show how N availability trajectories vary among ecosystems. Such assessment and monitoring efforts need to be complemented by further experimental and theoretical investigations into the causes of declining N availability, its implications for global carbon sequestration, and how its effects propagate through food webs. Responses will need to involve reducing N demand via lowering atmospheric CO 2 concentrations, and/or increasing N supply. Successfully mitigating and adapting to declining N availability will require a broader understanding that this phenomenon is occurring alongside the more widely recognized issue of anthropogenic eutrophication. Intercalibration of isotopic records from leaves, tree rings, and lake sediments suggests that N availability in many terrestrial ecosystems has steadily declined since the beginning of the industrial era. Reductions in N availability may affect many aspects of ecosystem functioning, including carbon sequestration and herbivore nutrition. Shaded areas indicate 80% prediction intervals; marker size is proportional to the number of measurements in each annual mean. Isotope data: (tree ring) K. K. McLauchlan et al. , Sci. Rep. 7 , 7856 (2017); (lake sediment) G. W. Holtgrieve et al. , Science 334 , 1545–1548 (2011); (foliar) J. M. Craine et al. , Nat. Ecol. Evol. 2 , 1735–1744 (2018)more » « less
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Abstract Although the positive effects of biodiversity on ecosystem functioning and stability have been extensively documented in the literature, previous studies have mostly explored the mechanisms of functioning and stability independently. It is unclear how biodiversity effects on functioning may covary with those on stability.
Here we developed an integrated framework to explore links between mechanisms underlying biodiversity effects on functioning and those on stability. Specifically, biodiversity effects on ecosystem functioning were partitioned into complementarity effects (
CE ) and selection effects (SE ), and those on stability were partitioned into species asynchrony and species stability. We investigated howCE andSE were linked to species asynchrony and stability and how their links might be mediated by species evenness, using a multi‐site grassland experiment.Our mixed‐effects models showed that a higher community productivity was mainly due to
CE and a higher community stability was mainly due to species asynchrony. Moreover,CE was positively related to species asynchrony, thus leading to a positive association between ecosystem productivity and stability.We used a structural equation model to illustrate how species evenness might mediate links between the various mechanisms. Communities with a higher evenness exhibited a higher
CE and species asynchrony, but a lowerSE and species stability. These evenness‐mediated associations enhanced the positive relationship betweenCE and species asynchrony, but blurred that betweenSE and species asynchrony.Synthesis . Our findings demonstrate mechanistic links between biodiversity effects on ecosystem functioning and stability. By doing so, our study contributes a novel framework for understanding ecological mechanisms of the functioning–stability relationship, which has important implications for developing management plans focused on strengthening synergies between ecosystem functioning and stability over the long term. -
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Abstract Plant performance is determined by the balance of intra‐ and interspecific neighbors within an individual's zone of influence. If individuals interact over smaller scales than the scales at which communities are measured, then altering neighborhood interactions may fundamentally affect community responses. These interactions can be altered by changing the number (species richness), abundances (species evenness), and positions (species pattern) of the resident plant species, and we aimed to test whether aggregating species at planting would alter effects of species richness and evenness on biomass production at a common scale of observation in grasslands. We varied plant species richness (2, 4, or 8 species and monocultures), evenness (0.64, 0.8, or 1.0), and pattern (planted randomly or aggregated in groups of four individuals) within 1 × 1 m plots established with transplants from a pool of 16 tallgrass prairie species and assessed plot‐scale biomass production and diversity over the first three growing seasons. As expected, more species‐rich plots produced more biomass by the end of the third growing season, an effect associated with a shift from selection to complementarity effects over time. Aggregating conspecifics at a 0.25‐m scale marginally reduced biomass production across all treatments and increased diversity in the most even plots, but did not alter biodiversity effects or richness–productivity relationships. Results support the hypothesis that fine‐scale species aggregation affects diversity by promoting species coexistence in this system. However, results indicate that inherent changes in species neighborhood relationships along grassland diversity gradients may only minimally affect community (meter) – scale responses among similarly designed biodiversity–ecosystem function studies. Given that species varied in their responses to local aggregation, it may be possible to use such species‐specific results to spatially design larger‐scale grassland communities to achieve desired diversity and productivity responses.
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Abstract A rich body of knowledge links biodiversity to ecosystem functioning (BEF), but it is primarily focused on small scales. We review the current theory and identify six expectations for scale dependence in the BEF relationship: (1) a nonlinear change in the slope of the BEF relationship with spatial scale; (2) a scale‐dependent relationship between ecosystem stability and spatial extent; (3) coexistence within and among sites will result in a positive BEF relationship at larger scales; (4) temporal autocorrelation in environmental variability affects species turnover and thus the change in BEF slope with scale; (5) connectivity in metacommunities generates nonlinear BEF and stability relationships by affecting population synchrony at local and regional scales; (6) spatial scaling in food web structure and diversity will generate scale dependence in ecosystem functioning. We suggest directions for synthesis that combine approaches in metaecosystem and metacommunity ecology and integrate cross‐scale feedbacks. Tests of this theory may combine remote sensing with a generation of networked experiments that assess effects at multiple scales. We also show how anthropogenic land cover change may alter the scaling of the BEF relationship. New research on the role of scale in BEF will guide policy linking the goals of managing biodiversity and ecosystems.
