Grazer diversity affects resistance to multiple stressors in an experimental seagrass ecosystem

When multiple stressors act simultaneously, their effects on ecosystems become more difficult to predict. In the face of multiple stressors, diverse ecosystems may be more stable if species respond differently to stressors or if functionally similar species can compensate for stressor effects on focal species. Many habitats around the globe are threatened by multiple stressors, including highly productive seagrass habitats. For example, in Chesapeake Bay, USA, regional climate change predictions suggest that elevated temperature and freshwater inputs are likely to be increasingly important stressors. Using seagrass mesocosms as a model system, we tested whether species richness of crustacean grazers buffers ecosystem properties against the impacts of elevated temperature and freshwater pulse stressors in a fully factorial experiment. Grazer species responded to pulsed salinity changes differently; abundance of Elasmopus levis responded negatively to freshwater pulses, whereas abundance of Gammarus mucronatus and Erichsonella attenuata responded positively or neutrally. Consistent with the hypothesis that biodiversity provides resistance stability, biomass of epiphytic algae that form the base of the food web was less affected by stressors in species‐rich grazer treatments than in single‐species grazer treatments. Stochastic (among‐replicate) variation of sessile invertebrate biomass within treatments was also reduced in more diverse grazer treatments. Therefore, grazer species richness tended to increase the resistance stability of both major components of the seagrass fouling community, algae and invertebrates, in the face of environmental stressors. Finally, in our model system, multi‐stressor impacts suggested a pattern of antagonism contrary to previous assumptions of synergistic stressor effects. Overall, our results confirm that invertebrate grazer species are functionally diverse in their response to environmental stressors, but are largely functionally redundant in their grazing effects leading to greater resistance stability of certain ecosystem properties in diverse grazer assemblages even when influenced by multiple environmental stressors.     

Stressor richness intensifies productivity loss but mitigates biodiversity loss

Ecosystems are subject to a multitude of anthropogenic environmental changes. Experimental research in the field of multiple stressors has typically involved varying the number of stressors, here termed stressor richness, but without controlling for total stressor intensity. Mistaking stressor intensity effects for stressor richness effects can misinform management decisions when there is a trade‐off between mitigating these two factors. We incorporate multiple stressors into three community models and show that, at a fixed total stressor intensity, increasing stressor richness aggravates joint stressor effects on ecosystem functioning, but reduces effects on species persistence and composition. In addition, stressor richness weakens the positive selection and negative complementarity effects on ecosystem function. We identify the among‐species variation of stressor effects on traits as a key determinant of the resulting community‐level stressor effects. Taken together, our results unravel the mechanisms linking multiple environmental changes to biodiversity and ecosystem function.     

Trophic Dependence of Ecosystem Resistance and Species Compensation in Experimentally Acidified Lake 302S (Canada)

Ecosystem resistance to the impacts of diverse human insults depends on the replacement of sensitive species by ones more tolerant of the stressor. Here we present evidence from a whole-lake acidification experiment (Lake 302S, Experimental Lakes Area, Canada) that resistance and species compensation decline with increasing trophic level. Diverse and fast-growing algal and rotifer assemblages with high dispersal potentials showed significant compensatory species dynamics, resulting in the maintenance of total biomass despite 30%–80% declines in species richness. Canonical correspondence analysis showed that significant compensatory algal and rotifer dynamics were best explained by differential species tolerances of acidified chemical conditions coupled with release from resource limitation and predation. However, less diverse cladoceran, copepod, and fish assemblages showed significant declines in total biomass and weak species compensation with loss of species during acidification. In comparison, algal and zooplankton species dynamics remained relatively synchronized in a nearby unperturbed reference lake (Lake 239) during the experiment. As a result, Lake 302S showed limited ecosystem resistance to anthropogenic acidification. Therefore, we hypothesize that lost species will increase the susceptibility of acidified lakes to the adverse impacts of other environmental stressors (for example, climate warming, stratospheric ozone depletion, invasive species). Consequently, the ecosystem stability of boreal lakes is expected to decline as global change proceeds. Received 2 January 2001; accepted 12 July 2002.     

Impact of multiple stressors on juvenile fish in estuaries of the northeast Pacific

A key step in identifying global change impacts on species and ecosystems is to quantify effects of multiple stressors. To date, the science of global change has been dominated by regional field studies, experimental manipulation, meta‐analyses, conceptual models, reviews, and studies focusing on a single stressor or species over broad spatial and temporal scales. Here, we provide one of the first studies for coastal systems examining multiple stressor effects across broad scales, focused on the nursery function of 20 estuaries spanning 1,600 km of coastline, 25 years of monitoring, and seven fish and invertebrate species along the northeast Pacific coast. We hypothesized those species most estuarine dependent and negatively impacted by human activities would have lower presence and abundances in estuaries with greater anthropogenic land cover, pollution, and water flow stress. We found significant negative relationships between juveniles of two of seven species (Chinook salmon and English sole) and estuarine stressors. Chinook salmon were less likely to occur and were less abundant in estuaries with greater pollution stress. They were also less abundant in estuaries with greater flow stress, although this relationship was marginally insignificant. English sole were less abundant in estuaries with greater land cover stress. Together, we provide new empirical evidence that effects of stressors on two fish species culminate in detectable trends along the northeast Pacific coast, elevating the need for protection from pollution, land cover, and flow stressors to their habitats. Lack of response among the other five species could be related to differing resistance to specific stressors, type and precision of the stressor metrics, and limitations in catch data across estuaries and habitats. Acquiring improved measurements of impacts to species will guide future management actions, and help predict how estuarine nursery functions can be optimized given anthropogenic stressors and climate change scenarios.     

Decadal‐scale regional changes in Canadian freshwater zooplankton: the likely consequence of complex interactions among multiple anthropogenic stressors

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Dominant species maintain ecosystem function with non-random species loss

Loss of species caused by widespread stressors, such as drought and fragmentation, is likely to be non‐random depending on species abundance in the community. We experimentally reduced the number of rare and uncommon plant species while independently reducing only the abundance of dominant grass species in intact, native grassland. This allowed us to simulate a non‐random pattern of species loss, based on species abundances, from communities shaped by natural ecological interactions and characterized by uneven species abundance distributions. Over two growing seasons, total above‐ground net primary productivity (ANPP) declined with reductions in abundance of the dominant species but was unaffected by a threefold decline in richness of less common species. In contrast, productivity of the remaining rare and uncommon species decreased with declining richness, in part due to loss of complementary interactions among these species. However, increased production of the dominant grasses offset the negative effects of species loss. We conclude that the dominant species, as controllers of ecosystem function, can provide short‐term resistance to reductions in ecosystem function when species loss is nonrandom. However, the concurrent loss of complementary interactions among rare and uncommon species, the most diverse component of communities, may contribute to additional species loss and portends erosion of ecosystem function in the long term.     

Impacts of sedimentation and nitrogen enrichment on wetland plant community development

Many factors influence which plant species are found in a particular wetland. The species pool is composed of the species present in the seed bank and species able to disperse into the wetland, and many abiotic and biotic factors interact to influence a species performance and abundance in the plant community. Anthropogenic activities produce specific stressors on wetland systems that alter these abiotic and biotic interactions, potentially altering species composition. We simulated three common wetland hydrogeomorphic (HGM) subclasses in a greenhouse to examine the effects of two stressors-sedimentation and nitrogen (N) enrichment-on the performance of 8 species grown in artificial communities. Species establishment, height, biomass, and foliar N and P concentrations were measured to explore species responses to stressors and competition, as well as the potential impacts of changes in species composition on ecosystem processes. Species were affected differently by sedimentation and N enrichment, and there were differences in overall community sensitivity to stressors between wetland subclasses. Sedimentation generally reduced seedling establishment, while N enrichment produced variable effects on height and biomass. Interspecific competition had little effect on establishment but significantly reduced most species biomass. Sedimentation generally lowered community biomass, diversity, and richness, while enrichment increased community biomass. Establishment, biomass, and foliar nutrient concentrations significantly differed between many species, suggesting that shifts in species composition may impact ecosystem processes such as nutrient cycling and carbon storage. Phalaris arundinacea, an aggressive clonal graminoid, universally dominated all wetland subclasses. This dominance across a range of environmental conditions (sedimentation, fertility, and hydrology) has important implications for both restoration and predicting the impacts of human activities on species composition. Our results suggest that, in regions where P. arundinacea is common, restoration projects that establish communities from seeds and human activities that cause vegetation removal are likely to become dominated by P. arundinacea.     

Multiple anthropogenic stressors cause ecological surprises in boreal lakes

The number of combinations of anthropogenic stressors affecting global change is increasing; however, few studies have empirically tested for their interactive effects on ecosystems. Most importantly, interactions among ecological stressors generate nonadditive effects that cannot be easily predicted based on single‐stressor studies. Here, we corroborate findings from an in situ mesocosm experiment with evidence from a whole‐ecosystem manipulation to demonstrate for the first time that interactions between climate and acidification determine their cumulative impact on the food‐web structure of coldwater lakes. Interactions among warming, drought, and acidification, rather than the sum of their individual effects, best explained significant changes in planktonic consumer and producer biomass over a 23‐year period. Further, these stressors interactively exerted significant synergistic and antagonistic effects on consumers and producers, respectively. The observed prevalence of long‐ and short‐term ecological surprises involving the cumulative impacts of multiple anthropogenic stressors highlights the high degree of uncertainty surrounding current forecasts of the consequences of global change.     

Biodiversity as a solution to mitigate climate change impacts on the functioning of forest ecosystems

Forest ecosystems are critical to mitigating greenhouse gas emissions through carbon sequestration. However, climate change has affected forest ecosystem functioning in both negative and positive ways, and has led to shifts in species/functional diversity and losses in plant species diversity which may impair the positive effects of diversity on ecosystem functioning. Biodiversity may mitigate climate change impacts on (I) biodiversity itself, as more‐diverse systems could be more resilient to climate change impacts, and (II) ecosystem functioning through the positive relationship between diversity and ecosystem functioning. By surveying the literature, we examined how climate change has affected forest ecosystem functioning and plant diversity. Based on the biodiversity effects on ecosystem functioning (B→EF), we specifically address the potential for biodiversity to mitigate climate change impacts on forest ecosystem functioning. For this purpose, we formulate a concept whereby biodiversity may reduce the negative impacts or enhance the positive impacts of climate change on ecosystem functioning. Further B→EF studies on climate change in natural forests are encouraged to elucidate how biodiversity might influence ecosystem functioning. This may be achieved through the detailed scrutiny of large spatial/long temporal scale data sets, such as long‐term forest inventories. Forest management strategies based on B→EF have strong potential for augmenting the effectiveness of the roles of forests in the mitigation of climate change impacts on ecosystem functioning.     

Multiple stressors and multifunctionality: limited effects on an illuminated benthic system

The bulk of experiments that study stressor effects on ecosystem functioning consider only individual functions one at a time, and such narrow focus may well bias our understanding of the overall impact on ecosystem functioning. We used data from six published experiments in which marine illuminated sediment systems were exposed to nutrient enrichment, toxicants, sedimentation and warming, either alone or in combination. Measured functions were primary production, community respiration, inorganic nitrogen and phosphorus fluxes, and autotrophic biomass. We calculated two indices of multifunctionality that simultaneously considered all six functions: (i) a weighted average level of the functions and (ii) the number of functions that simultaneously exceed a critical threshold level. Stressors affected individual functions both positively and negatively, but multifunctionality was generally unaffected by both single and joint stressors. The filtering capacity of coastal illuminated sediment systems thus appears resilient to exposure to moderate levels of multiple stressors, most probably due to the robustness of the benthic microalgal community. We recommend using a multifunctionality approach in future studies on cumulative stressor effects on ecosystem functioning, particularly when considering functions related to ecosystem services.     

Effects of three stressors on vegetation in an oligohaline marsh

1. When multiple stressors have interactive effects they can lead to important changes in ecosystem function. We examined how three stressors affected the plant community in an oligohaline marsh in southeastern Louisiana, U.S.A. These stressors included herbivory (mostly by the introduced rodent Myocastor coypus ), disturbance (herbicide application) and nutrient enrichment (three levels of N–P–K fertilizer). Sampling was conducted six times over 4 years.
2. Recovery after disturbance was slow, such that after 26 months biomass in disturbed plots was 36% that of controls. Slow recovery appeared to be due to herbivory, as exclusion of herbivores for 14 months led to much more biomass compared to non-excluded plots. Exclusion did not, however, aid recovery of species richness; this recovery required 51 months in total.
3. Nutrient enrichment increased biomass by 41% and decreased species richness by c. 20% in later sampling periods. Decreased species richness was due primarily to a reduced ability of dominant species to co-exist (as determined with Hill's diversity number N1). Nutrient enrichment did not interact with the other treatments.
4. Disturbance favoured two grasses ( Echinochloa crus-galli and Leptochloa fascicularis ), while lack of disturbance favoured two herbs ( Sagittaria lancifolia and Polygonum punctatum ) and two vines ( Ipomoea sagittata and Cuscuta pentagona ). Nutrient enrichment positively affected abundance of two species ( C. pentagona and L. fascicularis ). Herbivory did not affect species composition.
5. The effect of one stressor (experimental disturbance) on plant biomass depended on the strength of another stressor (herbivory). Nutrient enrichment was also important in affecting the plant community, but only as a single stressor. All effects changed over time, and it was clear that to understand properly the effects of multiple stressors, long-term, manipulative field experiments are necessary.     

Patterns of benthic algae and cyanobacteria along twin‐stressor gradients of nutrients and fine sediment: a stream mesocosm experiment

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Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a synthetic macroalgal assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of S. muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of S. muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming, producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.     

Evaluating methods to quantify anthropogenic stressors on wild animals

Humans have a variety of direct and indirect impacts on wildlife and a number of methods have been proposed to identify and quantify anthropogenic stressors that negatively impact wildlife. The ideal method would ultimately help predict the presence, absence, or population viability of animals living with a particular stressor. We critically review seven methods that have been used, or are potentially useful, to identify anthropogenic stressors on animals. We rank them from fitness indicators to disturbance indicators: breeding success, mate choice, fluctuating asymmetry, flight initiation distance, immunocompetence, glucocorticoids, and cardiac response. We describe each method's ease of use, precision in quantifying the stressor, accuracy in predicting the presence, absence, or population viability of a species experiencing a given stressor, and the repeatability of the results across populations and species. From this analysis, we conclude that there is no single optimal method to quantify anthropogenic stressors; method selection will depend on precise goals and fiscal constraints.     

Evaluating stress in rocky shore and shallow reef habitats using the macrofauna of kelp holdfasts

Kelp often dominates hard rocky substrata in the coastal watersof temperate, boreal and subantarctic regions of the world's oceans. The habitat providedby kelp holdfasts supports high species richness and high abundance of a range of invertebratetaxa. Studies conducted on the effects of stress on these communities have indicated that theyare sensitive to stressors and that the response is often predictable. With the growing body ofliterature on natural variation in holdfast community structure, studies of the impacts of stressare becoming more readily interpretable making the macrofauna of this habitat a potentiallyimportant tool for evaluating anthropogenic effects on hard substrata in coastal environments. Limitations of the use of holdfast fauna in studies of stress include the fact that brownseaweeds may disappear from polluted areas of reef, and where they persist, holdfaststructure may be altered directly by the stressor. An additional problem is that manipulative,experimental studies are difficult. Artificial holdfasts may provide a means by which thedemonstrated sensitivity of the holdfast community can be combined with a manipulative approach to testspecific hypotheses related to effects of stress in shallow coastal environments.     

The effects of elevated temperature and dissolved ρCO2 on a marine foundation species

Understanding how climate change and other environmental stressors will affect species is a fundamental concern of modern ecology. Indeed, numerous studies have documented how climate stressors affect species distributions and population persistence. However, relatively few studies have investigated how multiple climate stressors might affect species. In this study, we investigate the impacts of how two climate change factors affect an important foundation species. Specifically, we tested how ocean acidification from dissolution of CO₂ and increased sea surface temperatures affect multiple characteristics of juvenile eastern oysters (Crassostrea virginica). We found strong impacts of each stressor, but no interaction between the two. Simulated warming to mimic heat stressed summers reduced oyster growth, survival, and filtration rates. Additionally, we found that CO₂‐induced acidification reduced strength of oyster shells, which could potentially facilitate crab predation. As past studies have detected few impacts of these stressors on adult oysters, these results indicate that early life stages of calcareous marine organisms may be more susceptible to effects of ocean acidification and global warming. Overall, these data show that predicted changes in temperature and CO₂ can differentially influence direct effects on individual species, which could have important implications for the nature of their trophic interactions.     

Land use impacts on stream community composition and concordance along a natural stress gradient

Most ecosystems are subjected to multiple stressors derived from natural and anthropogenic sources and community responses to human disturbance in naturally stressful habitats may differ from those in more benign habitats. We examined the influence of a natural (geology-driven acidity) vs. human-induced stress (land drainage) and their interaction on the composition and concordance of stream diatom, bryophyte and invertebrate communities. To account for differing drainage impacts in circumneutral (sedimentation) and naturally acid (reduced pH and increased metal concentrations) streams we investigated concordance in three groups of streams: reference (circumneutral and naturally acidic reference), circumneutral (reference and drained) and naturally acidic (reference and drained) streams. We expected diatoms to respond more strongly to anthropogenic acidification and more weakly to sedimentation compared to bryophytes and invertebrates. We expected overall strong concordance among the three taxonomic groups, but especially so in reference streams. All three organism groups had distinct species composition in naturally acidic vs. circumneutral streams. Concordance between communities was overall strong, especially so in the reference streams. All groups responded to drainage disturbance in both types of streams. Invertebrates were slightly less responsive to increased acidification in the naturally acidic streams but were more affected by sedimentation in the circumneutral streams than were the other two groups. The natural stressor affected communities more than the anthropogenic stressors. Naturally stressed communities were affected by an anthropogenic stressor as much as those in more benign habitats, although the additional stressor was similar to the initial stress (further reduction of stream pH). Naturally acid streams may need special concern in bioassessment because models based on circumneutral reference sites will likely produce biased predictions for these streams.     

Can biological traits of stream invertebrates help disentangle the effects of multiple stressors in an agricultural catchment?

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Shifts of community composition and population density substantially affect ecosystem function despite invariant richness

There has been considerable focus on the impacts of environmental change on ecosystem function arising from changes in species richness. However, environmental change may affect ecosystem function without affecting richness, most notably by affecting population densities and community composition. Using a theoretical model, we find that, despite invariant richness, (1) small environmental effects may already lead to a collapse of function; (2) competitive strength may be a less important determinant of ecosystem function change than the selectivity of the environmental change driver and (3) effects on ecosystem function increase when effects on composition are larger. We also present a complementary statistical analysis of 13 data sets of phytoplankton and periphyton communities exposed to chemical stressors and show that effects on primary production under invariant richness ranged from ?75% to +10%. We conclude that environmental protection goals relying on measures of richness could underestimate ecological impacts of environmental change.     

Ecosystem engineering effects on species diversity across ecosystems: a meta‐analysis

Ecosystem engineering is increasingly recognized as a relevant ecological driver of diversity and community composition. Although engineering impacts on the biota can vary from negative to positive, and from trivial to enormous, patterns and causes of variation in the magnitude of engineering effects across ecosystems and engineer types remain largely unknown. To elucidate the above patterns, we conducted a meta‐analysis of 122 studies which explored effects of animal ecosystem engineers on species richness of other organisms in the community. The analysis revealed that the overall effect of ecosystem engineers on diversity is positive and corresponds to a 25% increase in species richness, indicating that ecosystem engineering is a facilitative process globally. Engineering effects were stronger in the tropics than at higher latitudes, likely because new or modified habitats provided by engineers in the tropics may help minimize competition and predation pressures on resident species. Within aquatic environments, engineering impacts were stronger in marine ecosystems (rocky shores) than in streams. In terrestrial ecosystems, engineers displayed stronger positive effects in arid environments (e.g. deserts). Ecosystem engineers that create new habitats or microhabitats had stronger effects than those that modify habitats or cause bioturbation. Invertebrate engineers and those with lower engineering persistence (<1 year) affected species richness more than vertebrate engineers which persisted for >1 year. Invertebrate species richness was particularly responsive to engineering impacts. This study is the first attempt to build an integrative framework of engineering effects on species diversity; it highlights the importance of considering latitude, habitat, engineering functional group, taxon and persistence of their effects in future theoretical and empirical studies.