Interactive community responses to disturbance in streams: disturbance history moderates the influence of disturbance types

Both disturbance history and disturbance type act to structure communities through selecting for particular species traits but they may also interact. For example, flooding selects for species with flood‐resistant traits in streams, but those traits could make communities susceptible to other disturbances and so could cause shifts in community composition due to anthropogenic climate change. To better understand the interactive influences of disturbance history and type on community composition, we investigated the response of macroinvertebrate communities to disturbance using in‐stream channels. Using a split‐plot design, individual channels in five ‘stable’ streams and five ‘frequently disturbed’ streams (disturbance history) were subject to different disturbance type treatments (flooding, drying and a control). Disturbance type independently drove effects on species diversity, but all other effects of disturbance type depended on disturbance history. In particular, the interaction of disturbance type and history determined overall community response. Both disturbance types tested produced similar community responses in frequently disturbed streams, including changes in community composition and alterations to the abundance of less mobile taxa, but low‐flow had a significantly greater effect in stable streams. Macroinvertebrate drift was greatest in the rock‐rolling treatments and significantly less in the low‐flow treatment for both disturbance histories. Therefore, disturbance history moderated the effects of disturbance type and determined the mechanism of community response by determining how well species were adapted to disturbance. This outcome suggests that previous disturbances strongly influence how vulnerable communities are to changes in disturbance, and so should be considered when predicting how changes in disturbance regimes will affect future community composition.     

A trait‐based framework for stream algal communities

The use of trait‐based approaches to detect effects of land use and climate change on terrestrial plant and aquatic phytoplankton communities is increasing, but such a framework is still needed for benthic stream algae. Here we present a conceptual framework of morphological, physiological, behavioural and life‐history traits relating to resource acquisition and resistance to disturbance. We tested this approach by assessing the relationships between multiple anthropogenic stressors and algal traits at 43 stream sites. Our “natural experiment” was conducted along gradients of agricultural land‐use intensity (0–95% of the catchment in high‐producing pasture) and hydrological alteration (0–92% streamflow reduction resulting from water abstraction for irrigation) as well as related physicochemical variables (total nitrogen concentration and deposited fine sediment). Strategic choice of study sites meant that agricultural intensity and hydrological alteration were uncorrelated. We studied the relationships of seven traits (with 23 trait categories) to our environmental predictor variables using general linear models and an information‐theoretic model‐selection approach. Life form, nitrogen fixation and spore formation were key traits that showed the strongest relationships with environmental stressors. Overall, FI (farming intensity) exerted stronger effects on algal communities than hydrological alteration. The large‐bodied, non‐attached, filamentous algae that dominated under high farming intensities have limited dispersal abilities but may cope with unfavourable conditions through the formation of spores. Antagonistic interactions between FI and flow reduction were observed for some trait variables, whereas no interactions occurred for nitrogen concentration and fine sediment. Our conceptual framework was well supported by tests of ten specific hypotheses predicting effects of resource supply and disturbance on algal traits. Our study also shows that investigating a fairly comprehensive set of traits can help shed light on the drivers of algal community composition in situations where multiple stressors are operating. Further, to understand non‐linear and non‐additive effects of such drivers, communities need to be studied along multiple gradients of natural variation or anthropogenic stressors.     

Anthropogenic disturbance and streams: land use and land‐use change affect stream ecosystems via multiple pathways

1. Ecosystems are strongly influenced by land use practices. However, identifying the mechanisms behind these influences is complicated by the many potential pathways (often indirect) between land use and ecosystems and by the long‐lasting effects of past land use. To support ecosystem restoration and conservation efforts, we need to better understand these indirect and lasting effects. 2. We constructed structural equation models (SEM) to evaluate the direct and indirect effects of contemporary (2002) land use (agriculture and development) and change in land use from 1952 to 2002 on present‐day streams (n = 190) in Maryland, U.S.A. Additional variables examined included site location, system size, altitude, per cent sand in soils, riparian condition, habitat quality, stream water NO₃‐N and benthic macroinvertebrate and fish measures of stream condition. Our first SEM (2002 Land Use) included the proportions of contemporary agriculture and development in catchments in the model. The second SEM (Land Use Change) included five measures of land use change (proportion agricultural in both times, developed in both times, agricultural in 1952 and developed in 2002, forested in 1952 and developed in 2002 and agricultural in 1952 and forested in 2002). 3. The data set fit both SEMs well. The 2002 Land Use model explained 71% of variation in NO₃‐N and 55%, 42% and 38% of variation in riffle quality, macroinvertebrate condition and fish condition, respectively. The Land Use Change model explained similar amounts of variation in NO₃‐N (R² = 0.72), riffle quality (R² = 0.57) and macroinvertebrate condition (R² = 0.44) but slightly more variation in fish condition (R² = 0.43). 4. Both models identified pathways through which landscape variables affect stream responses, including negative direct effects of latitude on macroinvertebrate and fish conditions and positive direct and indirect effects of altitude on NO₃‐N, riffle quality and macroinvertebrate and fish conditions. The 2002 Land Use model showed contemporary development and agriculture had positive total effects on NO₃‐N (both through direct pathways); contemporary development had negative effects on macroinvertebrate condition. The Land Use Change model showed that contemporary developed land that was forested in 1952 had no effects on NO₃‐N; current developed land that was developed or agricultural in 1952 showed positive effects on NO₃‐N. Forests that were agricultural in 1952 had negative effects on NO₃‐N, suggesting reduced NO₃‐N export with reforestation. The Land Use Change model also showed negative total effects of all types of contemporary developed land (developed, agricultural or forested in 1952) on benthic condition. Developed land that was forested in 1952 had negative effects on fish condition. Forest sites that were agricultural in 1952 had negative effects on fish and macroinvertebrate conditions, suggesting a long‐term imprint of abandoned agriculture in stream communities. 5. Our analyses (i) identified multiple indirect effects of contemporary land use on streams, (ii) showed that current land uses with different land use histories can exhibit different effects on streams and (iii) demonstrated an imprint of land use lasting >50 years. Knowledge of these indirect and long‐term effects of land use will help to conserve and restore streams.     

Environmental context determines multi‐trophic effects of consumer species loss

Loss of biodiversity and nutrient enrichment are two of the main human impacts on ecosystems globally, yet we understand very little about the interactive effects of multiple stressors on natural communities and how this relates to biodiversity and ecosystem functioning. Advancing our understanding requires the following: (1) incorporation of processes occurring within and among trophic levels in natural ecosystems and (2) tests of context‐dependency of species loss effects. We examined the effects of loss of a key predator and two groups of its prey on algal assemblages at both ambient and enriched nutrient conditions in a marine benthic system and tested for interactions between the loss of functional diversity and nutrient enrichment on ecosystem functioning. We found that enrichment interacted with food web structure to alter the effects of species loss in natural communities. At ambient conditions, the loss of primary consumers led to an increase in biomass of algae, whereas predator loss caused a reduction in algal biomass (i.e. a trophic cascade). However, contrary to expectations, we found that nutrient enrichment negated the cascading effect of predators on algae. Moreover, algal assemblage structure varied in distinct ways in response to mussel loss, grazer loss, predator loss and with nutrient enrichment, with compensatory shifts in algal abundance driven by variation in responses of different algal species to different environmental conditions and the presence of different consumers. We identified and characterized several context‐dependent mechanisms driving direct and indirect effects of consumers. Our findings highlight the need to consider environmental context when examining potential species redundancies in particular with regard to changing environmental conditions. Furthermore, non‐trophic interactions based on empirical evidence must be incorporated into food web‐based ecological models to improve understanding of community responses to global change.     

Artificial light as a disturbance to light‐naïve streams

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In‐stream habitat and macroinvertebrate responses to riparian corridor length in rangeland streams

Conservation and restoration of riparian vegetation in agricultural landscapes has had mixed success at protecting in‐stream habitat, potentially due to the mismatch between watershed‐scale impacts and reach‐scale restoration. Prioritizing contiguous placement of small‐scale restoration interventions may effectively create larger‐scale restoration projects and improve ecological outcomes. We performed a multi‐site field study to evaluate whether greater linear length of narrow riparian tree corridors resulted in measurable benefits to in‐stream condition. We collected data at 41 sites with varying upstream tree cover nested within 13 groups in rangeland streams in coastal northern California, United States. We evaluated the effect of riparian tree corridor length on benthic macroinvertebrate communities, as well as food resources, water temperature, and substrate size. Sites with longer riparian corridors had higher percentages of invertebrates sensitive to disturbance (including clingers and EPT taxa) as well as lower water temperatures and less fine sediment, two of the most important aquatic stressors. Despite marked improvement, we found no evidence that macroinvertebrate communities fully recovered, suggesting that land use continued to constrain conditions. The restoration of long riparian corridors may be an economically viable and rapidly implementable technique to improve habitat, control sediment, and counter increasing water temperatures expected with climate change within the context of ongoing land use.     

Discriminating trait‐convergence and trait‐divergence assembly patterns in ecological community gradients

Question: Whereas similar ecological requirements lead to trait‐convergence assembly patterns (TCAP) of species in communities, the interactions controlling how species associate produce trait‐divergence assembly patterns (TDAP). Yet, the linking of the latter to community processes has so far only been suggested. We offer a method to elucidate TCAP and TDAP in ecological community gradients that will help fill this gap. Method: We evaluated the correlation between trait‐based described communities and ecological gradients, and using partial correlation, we separated the fractions reflecting TCAP and TDAP. The required input data matrices describe operational taxonomic units (OTUs) by traits, communities by the quantities or presence‐absence of these OTUs, and community sites by ecological variables. We defined plant functional types (PFTs) or species as community components after fuzzy weighting by the traits. The measured correlations for TCAP and TDAP were tested by permutation. The null model for TDAP preserves the trait convergence, the structure intrinsic in the fuzzy types, and community total abundances and autocorrelation. Results: We applied the method to trait‐based data from plant communities in south Brazil, one set in natural grassland experimental plots under different nitrogen and grazing levels, and another in sapling communities colonizing Araucaria forest patches of increasing size in a forest‐grassland mosaic. In these cases, depending on the traits considered, we found strong evidence of either TCAP or TDAP, or both, that was related to the environmental gradients. Conclusions: The method developed is able to reveal TCAP and TDAP that are more likely to be functional for specified ecological gradients, allowing establishment of objective hypotheses on their links to community processes.     

Direct and plant trait‐mediated effects of the local environmental context on butterfly oviposition patterns

Variation in the intensity of plant–animal interactions over different spatial scales is widespread and might strongly influence fitness and trait selection in plants. Differences in traits among plant individuals have been shown to influence variation in interaction intensities within populations, while differences in environmental factors and community composition are shown to be important for variation over larger scales. However, little is still known about the relative importance of the local environmental context vs. plant traits for the outcome of interactions within plant populations. We investigated how oviposition by the seed‐predator butterfly Phengaris alcon on its host plant Gentiana pneumonanthe was related to host plant traits and to local environmental variation, as well as how oviposition patterns translated into effects on host plant fruit set. We considered the local environmental context in terms of height of the surrounding vegetation and abundance of the butterfly's second host, Myrmica ants. The probability of oviposition was higher in plants that were surrounded by lower vegetation, and both the probability of oviposition and the number of eggs increased in early‐flowering and tall plants with many flowers in the three study populations. Flowering phenology, shoot height and flower production were, in turn, related to higher surrounding vegetation. Myrmica abundance was correlated with vegetation height, but had no effect on oviposition patterns. Oviposition and subsequent seed predation by the caterpillars strongly reduced host plant fruit set. Our results show that plant–animal interactions are context‐dependent not only because the context influences the abundance or the behavior of the animal interactor, but also because it influences the expression of plant traits that affect the outcome of the interaction. The results also demonstrate that heterogeneity in environmental conditions at a very local scale can be important for the outcomes of interactions.     

Environmental harshness and global richness patterns in glacier‐fed streams

Aim To test for a possible effect of environmental harshness on large‐scale latitudinal and elevational patterns in taxon richness of macrofauna in arctic and alpine glacier‐fed streams. Location Svalbard (79° N), Iceland (65° N), Norway (62° N), Switzerland and Italy (46° N), France (43° N), New Zealand (43° S) and Ecuador (0°), covering an elevational gradient from sea level to 4800 m a.s.l. Methods We gathered data from 63 sites along 13 streams and created an index of glacial influence (the glacial index, GI) as an integrative proxy for environmental harshness. The explicative power of the GI, environmental variables, latitude and elevation on taxon richness was tested in generalized linear models. Taxon richness along geographical gradients was analysed at standardized levels of GI in contour plots. Beta diversity and assemblage similarity was calculated at different GI intervals and compared with a null‐model. Results Overall, taxon richness decreased exponentially with increased GI (r²= 0.64), and of all included factors, GI had the highest explicative power. At low values of GI we found that local taxon richness varied along the coupled gradients of latitude and elevation in a hump‐shaped manner. However, this pattern disappeared at high values of GI, i.e. when environmental harshness increased. Beta diversity increased, while similarity among assemblages decreased towards high GI values. Main conclusions In our study system, the number of taxa able to cope with the harshest conditions was largely independent of the regional taxon pool, and environmental harshness constituted a ‘fixed’ constraint for local richness, irrespective of latitude and elevation. Contrary to expectations, we found that beta diversity was highest and similarity lowest among the harshest sites, suggesting that taxon richness was not solely driven by niche selection based on environmental tolerances, but also stochastic ecological drift, leading to dispersal‐limited communities.     

Identifying models of trait‐mediated community assembly using random forests and approximate Bayesian computation

Ecologists often use dispersion metrics and statistical hypothesis testing to infer processes of community formation such as environmental filtering, competitive exclusion, and neutral species assembly. These metrics have limited power in inferring assembly models because they rely on often‐violated assumptions. Here, we adapt a model of phenotypic similarity and repulsion to simulate the process of community assembly via environmental filtering and competitive exclusion, all while parameterizing the strength of the respective ecological processes. We then use random forests and approximate Bayesian computation to distinguish between these models given the simulated data. We find that our approach is more accurate than using dispersion metrics and accounts for uncertainty in model selection. We also demonstrate that the parameter determining the strength of the assembly processes can be accurately estimated. This approach is available in the R package CAMI; Community Assembly Model Inference. We demonstrate the effectiveness of CAMI using an example of plant communities living on lava flow islands.     

Multi‐scale responses of plant species diversity in semi‐natural buffer strips to agricultural landscapes

Question: How does agricultural land usage affect plant species diversity in semi‐natural buffer strips at multiple scales? Location: Lepsämä River watershed, Nurmijärvi, Southern Finland. Methods: Species diversity indicators included both richness and evenness. Plant communities in buffer strips were surveyed in 29 sampling sites. Using ArcGIS Desktop 9.0 (ArcInfo) and Fragstats 3.3 for GIS analysis, the landscape composition around each sampling site was characterized by seven parameters in square sectors at five scales: 4, 36, 100, 196, and 324ha. For each scale, Principle Component Analysis was used to examine the importance of each structural metric to diversity indicators using multiple regression and other simple analyses. Results: For all but the smallest scales (4 ha), two structural metrics including the diversity of land cover types and percentage of arable land were positively and negatively correlated with species richness, respectively. Both metrics had the highest correlation coefficients for species richness at the second largest scale (196 ha). The density of arable field edges between the fields was the only metric that correlated with species evenness for all scales, which had highest predictive power at the second smallest scale (36 ha). Conclusions: Species richness and evenness of buffer strips had scale‐dependent relationships to land use in agricultural ecosystems. The results of this study indicated that species richness depends on the pattern of arable land use at large scales, which may relate to the regional species pool. Meanwhile, species evenness depended on the level of field edge density at small scales, which relates to how the nearby farmland was divided by the edges (e.g. many small‐scale fields with high edge density or a few big‐scale fields with low edge density). This implies that it is important to manage the biodiversity of buffer strips within a landscape context at multiple scales.     

Impacts of multiple anthropogenic stressors on stream macroinvertebrate community composition and functional diversity

Ensuring the provision of essential ecosystem services in systems affected by multiple stressors is a key challenge for theoretical and applied ecology. Trait‐based approaches have increasingly been used in multiple‐stressor research in freshwaters because they potentially provide a powerful method to explore the mechanisms underlying changes in populations and communities. Individual benthic macroinvertebrate traits associated with mobility, life history, morphology, and feeding habits are often used to determine how environmental drivers structure stream communities. However, to date multiple‐stressor research on stream invertebrates has focused more on taxonomic than on functional metrics. We conducted a fully crossed, 4‐factor experiment in 64 stream mesocosms fed by a pristine montane stream (21 days of colonization, 21 days of manipulations) and investigated the effects of nutrient enrichment, flow velocity reduction and sedimentation on invertebrate community, taxon, functional diversity and trait variables after 2 and 3 weeks of stressor exposure. 89% of the community structure metrics, 59% of the common taxa, 50% of functional diversity metrics, and 79% of functional traits responded to at least one stressor each. Deposited fine sediment and flow velocity reduction had the strongest impacts, affecting invertebrate abundances and diversity, and their effects translated into a reduction of functional redundancy. Stressor effects often varied between sampling occasions, further complicating the prediction of multiple‐stressor effects on communities. Overall, our study suggests that future research combining community, trait, and functional diversity assessments can improve our understanding of multiple‐stressor effects and their interactions in running waters.     

Exploring Core Response Mechanisms to Multiple Environmental Stressors Via A Genome‐Wide Study in the Brown Alga Saccharina japonica (Laminariales,Phaeophyceae)

Saccharina japonica is an important large brown alga and a major component of productive beds on the northwest coast of the Pacific Ocean. Abiotic stress response mechanisms are receiving considerable attention because global climate change is increasing their abiotic stress levels. However, our knowledge of how S. japonica broadly responds to stress is limited. In this study, we investigated the S. japonica responsive genes underlying acclimation to diverse stressors of acidification, high light, high temperature, hypersalinity, and hyposalinity and identified 408 core genes constantly and differentially expressed in response to all stressors. Our results confirm that stressors had strong effects on genes participating in photosynthesis, amino acid metabolism, carbohydrate metabolism, halogen metabolism, and reactive oxygen species defense. These findings will improve our understanding of brown algal response mechanisms linked to environmental stress and provide a list of candidate genes for improving algal stress tolerance in light of environmental stress in future studies.     

Environmental predictability of taxonomic and functional community composition in high‐latitude streams

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Macroinvertebrate community response to natural and forest harvest gradients in western Oregon headwater streams

1. To examine the effects of forest harvest practices on headwater stream macroinvertebrates, we compiled a 167 site database with macroinvertebrate, fish, physical habitat and catchment land cover data from the three forested ecoregions in western Oregon. For our analysis, headwater streams were defined by catchment areas <10 km² and perennial water during summer low flows. Almost all sites in the database were selected using a randomised survey design, constituting a representative sample of headwater streams in these ecoregions. 2. Macroinvertebrate taxonomic and functional feeding group composition were very similar among the three ecoregions in the study area (Coast Range, Cascades and Klamath Mountains). On average, 55% of the individuals at each site were in the orders Ephemeroptera, Plecoptera or Trichoptera. Dipteran taxa (mostly chironomids) accounted for another 34%. At almost all sites, non‐insects made up <10% of the macroinvertebrate assemblage. Almost half (49%) of the assemblages were collectors; remaining individuals were about evenly divided among scrapers, shredders and predators. 3. There were 189 different macroinvertebrate taxa at the 167 sites with richness at individual sites ranging from 7 to 71 taxa. Ordination by non‐metric multidimensional scaling revealed a strong association between % Ephemeroptera, especially Baetis, and site scores along the first axis. This axis was also strongly related to % coarse substratum and fast water habitat. The second axis was strongly related to % intolerant individuals, site slope and altitude. No strong relationships were evident between any ordination axis and either logging activity, presence/absence of fish, catchment size or ecoregion. 4. Based on macroinvertebrate index of biotic integrity (IBI) scores, 62% of the sites had no impairment, 31% of the sites had slight impairment and only 6% of the sites had moderate or severe impairment. IBI scores were not strongly related to forest harvest history. All four severely impaired sites and five of the seven sites with moderate impairment were lower altitude, shallower slope stream reaches located in the Coast Range with evidence of agricultural activity in their catchment or riparian zone. % sand + fine substratum was the environmental variable most strongly related to macroinvertebrate IBI.     

Habitat heterogeneity and disturbance influence patterns of community temporal variability in a small temperate stream

Both habitat heterogeneity and disturbance can profoundly influence ecological systems at many levels of biological and ecological organization. However, the joint influences of heterogeneity and disturbance on temporal variability in communities have received little attention despite the intense homogenizing influence of human activity. I performed a field manipulation of substrate heterogeneity in a small New England stream, and measured changes in benthic macroinvertebrate communities for 100 days—a period that included both a severe drought and a flood. Generally, community variability decreased with increasing substrate heterogeneity. However, within sampling intervals, this relationship tended to fluctuate through time, apparently tracking changes in hydrology. At the beginning of the experiment, community temporal variability clearly decreased along a gradient of increasing substrate heterogeneity—a result consistent with an observational study performed the previous year. During the subsequent weeks, droughts and flooding created exceptionally high variability in both hydrology and benthic macroinvertebrate community structure resulting in the disappearance of this relationship. However, during the last weeks of the experiment when hydrologic conditions were relatively more stable, the negatively sloped relationship between community temporal variability and habitat heterogeneity reemerged and mimicked relationships observed both early in the experiment and in the previous year’s study. High habitat heterogeneity may promote temporal stability through several mechanisms including stabilization of resources and increased refugia from minor disturbances or predation. However, the results of this experiment suggest that severe disturbance events can create large-scale environmental variability that effectively swamps the influence of habitat heterogeneity, illustrating that a thorough understanding of community temporal variability in natural systems will necessarily consider sources of environmental variability at multiple spatial and temporal scales. Handling editor: L. M. Bini     

Impacts of fine sediment addition to tussock, pasture, dairy and deer farming streams in New Zealand

1. Increased fine sediment input caused by agricultural development is expected to act as a stressor for stream ecosystems. In a large‐scale field experiment, we added fine river sand to 50‐m reaches of three second‐order streams in each of four categories of catchment development (ungrazed tussock grasslands, grazed pasture, dairying and deer farming) and measured the responses of macroinvertebrates and aquatic moss. 2. Before addition, fine sediment cover differed between land uses, being lowest in tussock (7%), intermediate in pasture (30%) and dairy (47%) and highest in deer streams (88%). Sediment addition increased cover by one land‐use category (e.g. augmented sediment cover in tussock streams was similar to pre‐existing cover in pasture streams), and cover remained high in impact reaches (compared with controls) throughout the 5‐week experiment. Sediment addition did not change concentrations of phosphate, nitrate and ammonium, which were generally highest in dairy streams and lowest in tussock streams. 3. Aquatic mosses (most common in tussock, absent in dairy and deer), invertebrate density (highest in deer, lowest in tussock), taxon richness (highest in pasture, lowest in deer) and diversity (highest in pasture and tussock, lowest in dairy and deer) all differed between land uses. Sediment addition resulted in reductions of moss cover, invertebrate taxon richness and richness of Ephemeroptera, Plecoptera and Trichoptera in impact relative to control reaches. 4. The impact of sediment addition was strongest in pasture streams where pre‐existing sediment cover was moderate and richness and diversity of the invertebrate community highest. However, even in the already sediment‐rich and species‐poor deer streams, density of one common taxon was reduced significantly by sediment addition, and another two were affected in the same way in dairy streams, the second‐most intense land use. 5. Our experiment has disentangled the impact of sediment addition from other concomitant land‐use effects that could not be reliably distinguished in previous research, which has mainly consisted of correlative studies or unrealistically small‐scale experiments.