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1.  Forecasting the combined effects of urbanization and climate change on stream ecosystems: from impacts to management options 
The Journal of Applied Ecology  2009;46(1):154-163.
Streams collect runoff, heat, and sediment from their watersheds, making them highly vulnerable to anthropogenic disturbances such as urbanization and climate change. Forecasting the effects of these disturbances using process-based models is critical to identifying the form and magnitude of likely impacts. Here, we integrate a new biotic model with four previously developed physical models (downscaled climate projections, stream hydrology, geomorphology, and water temperature) to predict how stream fish growth and reproduction will most probably respond to shifts in climate and urbanization over the next several decades.The biotic submodel couples dynamics in fish populations and habitat suitability to predict fish assemblage composition, based on readily available biotic information (preferences for habitat, temperature, and food, and characteristics of spawning) and day-to-day variability in stream conditions.We illustrate the model using Piedmont headwater streams in the Chesapeake Bay watershed of the USA, projecting ten scenarios: Baseline (low urbanization; no on-going construction; and present-day climate); one Urbanization scenario (higher impervious surface, lower forest cover, significant construction activity); four future climate change scenarios [Hadley CM3 and Parallel Climate Models under medium-high (A2) and medium-low (B2) emissions scenarios]; and the same four climate change scenarios plus Urbanization.Urbanization alone depressed growth or reproduction of 8 of 39 species, while climate change alone depressed 22 to 29 species. Almost every recreationally important species (i.e. trouts, basses, sunfishes) and six of the ten currently most common species were predicted to be significantly stressed. The combined effect of climate change and urbanization on adult growth was sometimes large compared to the effect of either stressor alone. Thus, the model predicts considerable change in fish assemblage composition, including loss of diversity.Synthesis and applications. The interaction of climate change and urban growth may entail significant reconfiguring of headwater streams, including a loss of ecosystem structure and services, which will be more costly than climate change alone. On local scales, stakeholders cannot control climate drivers but they can mitigate stream impacts via careful land use. Therefore, to conserve stream ecosystems, we recommend that proactive measures be taken to insure against species loss or severe population declines. Delays will inevitably exacerbate the impacts of both climate change and urbanization on headwater systems.
doi:10.1111/j.1365-2664.2008.01599.x
PMCID: PMC2695864  PMID: 19536343
multiple stressors; urbanization; fish assemblage; headwater stream; siltation; flow regime; temperature regime; urban stream syndrome
2.  Response of biotic communities to salinity changes in a Mediterranean hypersaline stream 
Saline Systems  2006;2:12.
Background
This study investigates the relationship between salinity and biotic communities (primary producers and macroinvertebrates) in Rambla Salada, a Mediterranean hypersaline stream in SE Spain. Since the 1980's, the mean salinity of the stream has fallen from about 100 g L-1 to 35.5 g L-1, due to intensive irrigated agriculture in the watershed. Furthermore, large dilutions occur occasionally when the water irrigation channel suffers cracks.
Results
Along the salinity gradient studied (3.5 – 76.4 g L-1) Cladophora glomerata and Ruppia maritima biomass decreased with increasing salinity, while the biomass of epipelic algae increased. Diptera and Coleoptera species dominated the community both in disturbed as in re-established conditions. Most macroinvertebrates species found in Rambla Salada stream are euryhaline species with a broad range of salinity tolerance. Eight of them were recorded in natural hypersaline conditions (~100 g L-1) prior to important change in land use of the watershed: Ephydra flavipes, Stratyomis longicornis, Nebrioporus ceresyi, N. baeticus, Berosus hispanicus, Enochrus falcarius, Ochthebius cuprescens and Sigara selecta. However, other species recorded in the past, such as Ochthebius glaber, O. notabilis and Enochrus politus, were restricted to a hypersaline source or absent from Rambla Salada. The dilution of salinity to 3.5 – 6.8 gL-1 allowed the colonization of species with low salininty tolerance, such as Melanopsis praemorsa, Anax sp., Simulidae, Ceratopogonidae and Tanypodinae. The abundance of Ephydra flavipes and Ochthebius corrugatus showed a positive significant response to salinity, while Anax sp., Simulidae, S. selecta, N. ceresyi, N. baeticus, and B. hispanicus showed significant negative correlations. The number of total macroinvertebrate taxa, Diptera and Coleoptera species, number of families, Margalef's index and Shannon's diversity index decreased with increasing salinity. However, the rest of community parameters, such as the abundance of individuals, evenness and Simpson's index, showed no significant response to changes in salinity. Classification and ordination analysis revealed major differences in macroinvertebrate community structure between hypersaline conditions (76.4 g L-1) and the rest of the communities observed at the lower salinity levels, and revealed that below ~75 g L-1, dissimilarities in the communities were greater between the two habitats studied (runs and pools) than between salinity levels.
Conclusion
Salinity was the first factor determining community composition and structure in Rambla Salada stream followed by the type of habitat.
doi:10.1186/1746-1448-2-12
PMCID: PMC1609171  PMID: 17014701
3.  Re-Meandering of Lowland Streams: Will Disobeying the Laws of Geomorphology Have Ecological Consequences? 
PLoS ONE  2014;9(9):e108558.
We evaluated the restoration of physical habitats and its influence on macroinvertebrate community structure in 18 Danish lowland streams comprising six restored streams, six streams with little physical alteration and six channelized streams. We hypothesized that physical habitats and macroinvertebrate communities of restored streams would resemble those of natural streams, while those of the channelized streams would differ from both restored and near-natural streams. Physical habitats were surveyed for substrate composition, depth, width and current velocity. Macroinvertebrates were sampled along 100 m reaches in each stream, in edge habitats and in riffle/run habitats located in the center of the stream. Restoration significantly altered the physical conditions and affected the interactions between stream habitat heterogeneity and macroinvertebrate diversity. The substrate in the restored streams was dominated by pebble, whereas the substrate in the channelized and natural streams was dominated by sand. In the natural streams a relationship was identified between slope and pebble/gravel coverage, indicating a coupling of energy and substrate characteristics. Such a relationship did not occur in the channelized or in the restored streams where placement of large amounts of pebble/gravel distorted the natural relationship. The analyses revealed, a direct link between substrate heterogeneity and macroinvertebrate diversity in the natural streams. A similar relationship was not found in either the channelized or the restored streams, which we attribute to a de-coupling of the natural relationship between benthic community diversity and physical habitat diversity. Our study results suggest that restoration schemes should aim at restoring the natural physical structural complexity in the streams and at the same time enhance the possibility of re-generating the natural geomorphological processes sustaining the habitats in streams and rivers. Documentation of restoration efforts should be intensified with continuous monitoring of geomorphological and ecological changes including surveys of reference river systems.
doi:10.1371/journal.pone.0108558
PMCID: PMC4180926  PMID: 25264627
4.  The Role of Headwater Streams in Downstream Water Quality1 
Knowledge of headwater influences on the water-quality and flow conditions of downstream waters is essential to water-resource management at all governmental levels; this includes recent court decisions on the jurisdiction of the Federal Clean Water Act (CWA) over upland areas that contribute to larger downstream water bodies. We review current watershed research and use a water-quality model to investigate headwater influences on downstream receiving waters. Our evaluations demonstrate the intrinsic connections of headwaters to landscape processes and downstream waters through their influence on the supply, transport, and fate of water and solutes in watersheds. Hydrological processes in headwater catchments control the recharge of subsurface water stores, flow paths, and residence times of water throughout landscapes. The dynamic coupling of hydrological and biogeochemical processes in upland streams further controls the chemical form, timing, and longitudinal distances of solute transport to downstream waters. We apply the spatially explicit, mass-balance watershed model SPARROW to consider transport and transformations of water and nutrients throughout stream networks in the northeastern United States. We simulate fluxes of nitrogen, a primary nutrient that is a water-quality concern for acidification of streams and lakes and eutrophication of coastal waters, and refine the model structure to include literature observations of nitrogen removal in streams and lakes. We quantify nitrogen transport from headwaters to downstream navigable waters, where headwaters are defined within the model as first-order, perennial streams that include flow and nitrogen contributions from smaller, intermittent and ephemeral streams. We find that first-order headwaters contribute approximately 70% of the mean-annual water volume and 65% of the nitrogen flux in second-order streams. Their contributions to mean water volume and nitrogen flux decline only marginally to about 55% and 40% in fourth- and higher-order rivers that include navigable waters and their tributaries. These results underscore the profound influence that headwater areas have on shaping downstream water quantity and water quality. The results have relevance to water-resource management and regulatory decisions and potentially broaden understanding of the spatial extent of Federal CWA jurisdiction in U.S. waters.
doi:10.1111/j.1752-1688.2007.00005.x
PMCID: PMC3307624  PMID: 22457565
rivers/streams; nitrogen; transport and fate; streamflow; headwaters; SWANCC; Rapanos
5.  Denitrification in Agriculturally Impacted Streams: Seasonal Changes in Structure and Function of the Bacterial Community 
PLoS ONE  2014;9(8):e105149.
Denitrifiers remove fixed nitrogen from aquatic environments and hydrologic conditions are one potential driver of denitrification rate and denitrifier community composition. In this study, two agriculturally impacted streams in the Sugar Creek watershed in Indiana, USA with different hydrologic regimes were examined; one stream is seasonally ephemeral because of its source (tile drainage), whereas the other stream has permanent flow. Additionally, a simulated flooding experiment was performed on the riparian benches of the ephemeral stream during a dry period. Denitrification activity was assayed using the chloramphenicol amended acetylene block method and bacterial communities were examined based on quantitative PCR and terminal restriction length polymorphisms of the nitrous oxide reductase (nosZ) and 16S rRNA genes. In the stream channel, hydrology had a substantial impact on denitrification rates, likely by significantly lowering water potential in sediments. Clear patterns in denitrification rates were observed among pre-drying, dry, and post-drying dates; however, a less clear scenario was apparent when analyzing bacterial community structure suggesting that denitrifier community structure and denitrification rate were not strongly coupled. This implies that the nature of the response to short-term hydrologic changes was physiological rather than increases in abundance of denitrifiers or changes in composition of the denitrifier community. Flooding of riparian bench soils had a short-term, transient effect on denitrification rate. Our results imply that brief flooding of riparian zones is unlikely to contribute substantially to removal of nitrate (NO3-) and that seasonal drying of stream channels has a negative impact on NO3- removal, particularly because of the time lag required for denitrification to rebound. This time lag is presumably attributable to the time required for the denitrifiers to respond physiologically rather than a change in abundance or community composition.
doi:10.1371/journal.pone.0105149
PMCID: PMC4149370  PMID: 25171209
6.  Coupled Effects of Natural and Anthropogenic Controls on Seasonal and Spatial Variations of River Water Quality during Baseflow in a Coastal Watershed of Southeast China 
PLoS ONE  2014;9(3):e91528.
Surface water samples of baseflow were collected from 20 headwater sub-watersheds which were classified into three types of watersheds (natural, urban and agricultural) in the flood, dry and transition seasons during three consecutive years (2010–2012) within a coastal watershed of Southeast China. Integrating spatial statistics with multivariate statistical techniques, river water quality variations and their interactions with natural and anthropogenic controls were examined to identify the causal factors and underlying mechanisms governing spatiotemporal patterns of water quality. Anthropogenic input related to industrial effluents and domestic wastewater, agricultural activities associated with the precipitation-induced surface runoff, and natural weathering process were identified as the potential important factors to drive the seasonal variations in stream water quality for the transition, flood and dry seasons, respectively. All water quality indicators except SRP had the highest mean concentrations in the dry and transition seasons. Anthropogenic activities and watershed characteristics led to the spatial variations in stream water quality in three types of watersheds. Concentrations of NH4+-N, SRP, K+, CODMn, and Cl− were generally highest in urban watersheds. NO3–N Concentration was generally highest in agricultural watersheds. Mg2+ concentration in natural watersheds was significantly higher than that in agricultural watersheds. Spatial autocorrelations analysis showed similar levels of water pollution between the neighboring sub-watersheds exhibited in the dry and transition seasons while non-point source pollution contributed to the significant variations in water quality between neighboring sub-watersheds. Spatial regression analysis showed anthropogenic controls played critical roles in variations of water quality in the JRW. Management implications were further discussed for water resource management. This research demonstrates that the coupled effects of natural and anthropogenic controls involved in watershed processes, contribute to the seasonal and spatial variation of headwater stream water quality in a coastal watershed with high spatial variability and intensive anthropogenic activities.
doi:10.1371/journal.pone.0091528
PMCID: PMC3950248  PMID: 24618771
7.  Watershed responses to Amazon soya bean cropland expansion and intensification 
The expansion and intensification of soya bean agriculture in southeastern Amazonia can alter watershed hydrology and biogeochemistry by changing the land cover, water balance and nutrient inputs. Several new insights on the responses of watershed hydrology and biogeochemistry to deforestation in Mato Grosso have emerged from recent intensive field campaigns in this region. Because of reduced evapotranspiration, total water export increases threefold to fourfold in soya bean watersheds compared with forest. However, the deep and highly permeable soils on the broad plateaus on which much of the soya bean cultivation has expanded buffer small soya bean watersheds against increased stormflows. Concentrations of nitrate and phosphate do not differ between forest or soya bean watersheds because fixation of phosphorus fertilizer by iron and aluminium oxides and anion exchange of nitrate in deep soils restrict nutrient movement. Despite resistance to biogeochemical change, streams in soya bean watersheds have higher temperatures caused by impoundments and reduction of bordering riparian forest. In larger rivers, increased water flow, current velocities and sediment flux following deforestation can reshape stream morphology, suggesting that cumulative impacts of deforestation in small watersheds will occur at larger scales.
doi:10.1098/rstb.2012.0425
PMCID: PMC3638438  PMID: 23610178
soya beans; watersheds; nitrogen; phosphorus; soil
8.  Dissolved Organic Carbon in Headwater Streams and Riparian Soil Organic Carbon along an Altitudinal Gradient in the Wuyi Mountains, China 
PLoS ONE  2013;8(11):e78973.
Stream water dissolved organic carbon (DOC) correlates positively with soil organic carbon (SOC) in many biomes. Does this relationship hold in a small geographic region when variations of temperature, precipitation and vegetation are driven by a significant altitudinal gradient? We examined the spatial connectivity between concentrations of DOC in headwater stream and contents of riparian SOC and water-soluble soil organic carbon (WSOC), riparian soil C:N ratio, and temperature in four vegetation types along an altitudinal gradient in the Wuyi Mountains, China. Our analyses showed that annual mean concentrations of headwater stream DOC were lower in alpine meadow (AM) than in subtropical evergreen broadleaf forest (EBF), coniferous forest (CF), and subalpine dwarf forest (SDF). Headwater stream DOC concentrations were negatively correlated with riparian SOC as well as WSOC contents, and were unrelated to riparian soil C:N ratio. Our findings suggest that DOC concentrations in headwater streams are affected by different factors at regional and local scales. The dilution effect of higher precipitation and adsorption of soil DOC to higher soil clay plus silt content at higher elevation may play an important role in causing lower DOC concentrations in AM stream of the Wuyi Mountains. Our results suggest that upscaling and downscaling of the drivers of DOC export from forested watersheds when exploring the response of carbon flux to climatic change or other drivers must done with caution.
doi:10.1371/journal.pone.0078973
PMCID: PMC3827275  PMID: 24265737
9.  Functional Changes in Littoral Macroinvertebrate Communities in Response to Watershed-Level Anthropogenic Stress 
PLoS ONE  2014;9(7):e101499.
Watershed-scale anthropogenic stressors have profound effects on aquatic communities. Although several functional traits of stream macroinvertebrates change predictably in response to land development and urbanization, little is known about macroinvertebrate functional responses in lakes. We assessed functional community structure, functional diversity (Rao’s quadratic entropy) and voltinism in macroinvertebrate communities sampled across the full gradient of anthropogenic stress in Laurentian Great Lakes coastal wetlands. Functional diversity and voltinism significantly decreased with increasing development, whereas agriculture had smaller or non-significant effects. Functional community structure was affected by watershed-scale development, as demonstrated by an ordination analysis followed by regression. Because functional community structure affects energy flow and ecosystem function, and functional diversity is known to have important implications for ecosystem resilience to further environmental change, these results highlight the necessity of finding ways to remediate or at least ameliorate these effects.
doi:10.1371/journal.pone.0101499
PMCID: PMC4090147  PMID: 25006811
10.  Fire and Grazing Influences on Rates of Riparian Woody Plant Expansion along Grassland Streams 
PLoS ONE  2014;9(9):e106922.
Grasslands are threatened globally due to the expansion of woody plants. The few remaining headwater streams within tallgrass prairies are becoming more like typical forested streams due to rapid conversion of riparian zones from grassy to wooded. Forestation can alter stream hydrology and biogeochemistry. We estimated the rate of riparian woody plant expansion within a 30 m buffer zone surrounding the stream bed across whole watersheds at Konza Prairie Biological Station over 25 years from aerial photographs. Watersheds varied with respect to experimentally-controlled fire and bison grazing. Fire frequency, presence or absence of grazing bison, and the historical presence of woody vegetation prior to the study time period (a proxy for proximity of propagule sources) were used as independent variables to predict the rate of riparian woody plant expansion between 1985 and 2010. Water yield was estimated across these years for a subset of watersheds. Riparian woody encroachment rates increased as burning became less frequent than every two years. However, a higher fire frequency (1–2 years) did not reverse riparian woody encroachment regardless of whether woody vegetation was present or not before burning regimes were initiated. Although riparian woody vegetation cover increased over time, annual total precipitation and average annual temperature were variable. So, water yield over 4 watersheds under differing burn frequencies was quite variable and with no statistically significant detected temporal trends. Overall, burning regimes with a frequency of every 1–2 years will slow the conversion of tallgrass prairie stream ecosystems to forested ones, yet over long time periods, riparian woody plant encroachment may not be prevented by fire alone, regardless of fire frequency.
doi:10.1371/journal.pone.0106922
PMCID: PMC4156405  PMID: 25192194
11.  Ecosystem Function in Appalachian Headwater Streams during an Active Invasion by the Hemlock Woolly Adelgid 
PLoS ONE  2013;8(4):e61171.
Forested ecosystems in the southeastern United States are currently undergoing an invasion by the hemlock woolly adelgid (HWA). Previous studies in this area have shown changes to forest structure, decreases in canopy cover, increases in organic matter, and changes to nutrient cycling on the forest floor and soil. Here, we were interested in how the effects of canopy loss and nutrient leakage from terrestrial areas would translate into functional changes in streams draining affected watersheds. We addressed these questions in HWA-infested watersheds at the Coweeta Hydrologic Laboratory in North Carolina. Specifically, we measured stream metabolism (gross primary production and ecosystem respiration) and nitrogen uptake from 2008 to 2011 in five streams across the Coweeta basin. Over the course of our study, we found no change to in-stream nutrient concentrations. While canopy cover decreased annually in these watersheds, this change in light penetration did not translate to higher rates of in-stream primary production during the summer months of our study. We found a trend towards greater heterotrophy within our watersheds, where in-stream respiration accounted for a much larger component of net ecosystem production than GPP. Additionally, increases in rhododendron cover may counteract changes in light and nutrient availability that occurred with hemlock loss. The variability in our metabolic and uptake parameters suggests an actively-infested ecosystem in transition between steady states.
doi:10.1371/journal.pone.0061171
PMCID: PMC3632569  PMID: 23613803
12.  Temporal Patterns and Environmental Correlates of Macroinvertebrate Communities in Temporary Streams 
PLoS ONE  2015;10(11):e0142370.
Temporary streams are characterised by short periods of seasonal or annual stream flow after which streams contract into waterholes or pools of varying hydrological connectivity and permanence. Although these streams are widespread globally, temporal variability of their ecology is understudied, and understanding the processes that structure community composition in these systems is vital for predicting and managing the consequences of anthropogenic impacts. We used multivariate and univariate approaches to investigate temporal variability in macroinvertebrate compositional data from 13 years of sampling across multiple sites from autumn and spring, in South Australia, the driest state in the driest inhabited continent in the world. We examined the potential of land-use, geographic and environmental variables to predict the temporal variability in macroinvertebrate assemblages, and also identified indicator taxa, that is, those highly correlated with the most significantly associated physical variables. Temporal trajectories of macroinvertebrate communities varied within site in both seasons and across years. A combination of land-use, geographic and environmental variables accounted for 24% of the variation in community structure in autumn and 27% in spring. In autumn, community composition among sites were more closely clustered together relative to spring suggesting that communities were more similar in autumn than in spring. In both seasons, community structure was most strongly correlated with conductivity and latitude, and community structure was more associated with cover by agriculture than urban land-use. Maintaining temporary streams will require improved catchment management aimed at sustaining seasonal flows and critical refuge habitats, while also limiting the damaging effects from increased agriculture and urban developments.
doi:10.1371/journal.pone.0142370
PMCID: PMC4640519  PMID: 26556711
13.  Multi-Scale, Direct and Indirect Effects of the Urban Stream Syndrome on Amphibian Communities in Streams 
PLoS ONE  2013;8(7):e70262.
Urbanization affects streams by modifying hydrology, increasing pollution and disrupting in-stream and riparian conditions, leading to negative responses by biotic communities. Given the global trend of increasing urbanization, improved understanding of its direct and indirect effects at multiple scales is needed to assist management. The theory of stream ecology suggests that the riverscape and the surrounding landscape are inextricably linked, and watershed-scale processes will also affect in-stream conditions and communities. This is particularly true for species with semi-aquatic life cycles, such as amphibians, which transfer energy between streams and surrounding terrestrial areas. We related measures of urbanization at different scales to frog communities in streams along an urbanization gradient in Melbourne, Australia. We used boosted regression trees to determine the importance of predictors and the shape of species responses. We then used structural equation models to investigate possible indirect effects of watershed imperviousness on in-stream parameters. The proportion of riparian vegetation and road density surrounding the site at the reach scale (500-m radius) had positive and negative effects, respectively, on species richness and on the occurrence of the two most common species in the area (Criniasignifera and Limnodynastesdumerilii). Road density and local aquatic vegetation interacted in influencing species richness, suggesting that isolation of a site can prevent colonization, in spite of apparently good local habitat. Attenuated imperviousness at the catchment scale had a negative effect on local aquatic vegetation, indicating possible indirect effects on frog species not revealed by single-level models. Processes at the landscape scale, particularly related to individual ranging distances, can affect frog species directly and indirectly. Catchment imperviousness might not affect adult frogs directly, but by modifying hydrology it can disrupt local vegetation and prove indirectly detrimental. Integrating multiple-scale management actions may help to meet conservation targets for streams in the face of urbanization.
doi:10.1371/journal.pone.0070262
PMCID: PMC3726379  PMID: 23922963
14.  Deforestation and Benthic Indicators: How Much Vegetation Cover Is Needed to Sustain Healthy Andean Streams? 
PLoS ONE  2014;9(8):e105869.
Deforestation in the tropical Andes is affecting ecological conditions of streams, and determination of how much forest should be retained is a pressing task for conservation, restoration and management strategies. We calculated and analyzed eight benthic metrics (structural, compositional and water quality indices) and a physical-chemical composite index with gradients of vegetation cover to assess the effects of deforestation on macroinvertebrate communities and water quality of 23 streams in southern Ecuadorian Andes. Using a geographical information system (GIS), we quantified vegetation cover at three spatial scales: the entire catchment, the riparian buffer of 30 m width extending the entire stream length, and the local scale defined for a stream reach of 100 m in length and similar buffer width. Macroinvertebrate and water quality metrics had the strongest relationships with vegetation cover at catchment and riparian scales, while vegetation cover did not show any association with the macroinvertebrate metrics at local scale. At catchment scale, the water quality metrics indicate that ecological condition of Andean streams is good when vegetation cover is over 70%. Further, macroinvertebrate community assemblages were more diverse and related in catchments largely covered by native vegetation (>70%). Our results suggest that retaining an important quantity of native vegetation cover within the catchments and a linkage between headwater and riparian forests help to maintain and improve stream biodiversity and water quality in Andean streams affected by deforestation. This research proposes that a strong regulation focused to the management of riparian buffers can be successful when decision making is addressed to conservation/restoration of Andean catchments.
doi:10.1371/journal.pone.0105869
PMCID: PMC4141824  PMID: 25147941
15.  Effects of Temperature, Salinity and Fish in Structuring the Macroinvertebrate Community in Shallow Lakes: Implications for Effects of Climate Change 
PLoS ONE  2012;7(2):e30877.
Climate warming may lead to changes in the trophic structure and diversity of shallow lakes as a combined effect of increased temperature and salinity and likely increased strength of trophic interactions. We investigated the potential effects of temperature, salinity and fish on the plant-associated macroinvertebrate community by introducing artificial plants in eight comparable shallow brackish lakes located in two climatic regions of contrasting temperature: cold-temperate and Mediterranean. In both regions, lakes covered a salinity gradient from freshwater to oligohaline waters. We undertook day and night-time sampling of macroinvertebrates associated with the artificial plants and fish and free-swimming macroinvertebrate predators within artificial plants and in pelagic areas. Our results showed marked differences in the trophic structure between cold and warm shallow lakes. Plant-associated macroinvertebrates and free-swimming macroinvertebrate predators were more abundant and the communities richer in species in the cold compared to the warm climate, most probably as a result of differences in fish predation pressure. Submerged plants in warm brackish lakes did not seem to counteract the effect of fish predation on macroinvertebrates to the same extent as in temperate freshwater lakes, since small fish were abundant and tended to aggregate within the macrophytes. The richness and abundance of most plant-associated macroinvertebrate taxa decreased with salinity. Despite the lower densities of plant-associated macroinvertebrates in the Mediterranean lakes, periphyton biomass was lower than in cold temperate systems, a fact that was mainly attributed to grazing and disturbance by fish. Our results suggest that, if the current process of warming entails higher chances of shallow lakes becoming warmer and more saline, climatic change may result in a decrease in macroinvertebrate species richness and abundance in shallow lakes.
doi:10.1371/journal.pone.0030877
PMCID: PMC3290630  PMID: 22393354
16.  Land Cover and Rainfall Interact to Shape Waterbird Community Composition 
PLoS ONE  2012;7(4):e35969.
Human land cover can degrade estuaries directly through habitat loss and fragmentation or indirectly through nutrient inputs that reduce water quality. Strong precipitation events are occurring more frequently, causing greater hydrological connectivity between watersheds and estuaries. Nutrient enrichment and dissolved oxygen depletion that occur following these events are known to limit populations of benthic macroinvertebrates and commercially harvested species, but the consequences for top consumers such as birds remain largely unknown. We used non-metric multidimensional scaling (MDS) and structural equation modeling (SEM) to understand how land cover and annual variation in rainfall interact to shape waterbird community composition in Chesapeake Bay, USA. The MDS ordination indicated that urban subestuaries shifted from a mixed generalist-specialist community in 2002, a year of severe drought, to generalist-dominated community in 2003, of year of high rainfall. The SEM revealed that this change was concurrent with a sixfold increase in nitrate-N concentration in subestuaries. In the drought year of 2002, waterbird community composition depended only on the direct effect of urban development in watersheds. In the wet year of 2003, community composition depended both on this direct effect and on indirect effects associated with high nitrate-N inputs to northern parts of the Bay, particularly in urban subestuaries. Our findings suggest that increased runoff during periods of high rainfall can depress water quality enough to alter the composition of estuarine waterbird communities, and that this effect is compounded in subestuaries dominated by urban development. Estuarine restoration programs often chart progress by monitoring stressors and indicators, but rarely assess multivariate relationships among them. Estuarine management planning could be improved by tracking the structure of relationships among land cover, water quality, and waterbirds. Unraveling these complex relationships may help managers identify and mitigate ecological thresholds that occur with increasing human land cover.
doi:10.1371/journal.pone.0035969
PMCID: PMC3338777  PMID: 22558286
17.  Studies of Giardia spp. and Cryptosporidium spp. in two adjacent watersheds. 
Two adjacent British Columbia, Canada, watersheds with similar topographical features were studied. Both the Black Mountain Irrigation District (BMID) and the Vernon Irrigation District (VID) serve rural agricultural communities which are active in cattle ranching. The present study was carried out in five phases, during which a total of 249 surface water samples were tested in the study watersheds. The aims of these phases were to determine levels of parasite contamination in raw water samples collected from the intakes as well as from other sites in each watershed and to investigate cattle in the watersheds as potential sources of parasite contamination of surface drinking water supplies. Giardia cysts were not detected in the raw water samples collected from lake sources at the headwaters of both watersheds but were found in 100% (70 or 70) of water samples collected at the BMID intake and 97% (68 of 70) of water samples collected at the VID intake. Significantly higher levels (P < 0.05) of Giardia cysts were found at the BMID intake (phase 1, 7 to 2,215 cysts per 100 liters; phase 3, 4.6 to 1,880 cysts per 100 liters) when compared with that of the VID intake (2 to 114 cysts per 100 liters). The BMID watershed has a more complex system of surface water sources than the VID watershed. Cattle have access to creeks in the BMID watershed, whereas access is restricted in the VID watershed. Collection of raw water samples from a creek upstream and downstream of a cattle ranch in the BMID watershed showed that the downstream location had significantly higher (P < 0.05) levels (0.6 to 42.9 cysts per 100 liters and 1.4 to 300.0 oocysts per 100 liters) of both Giardia cysts and Cryptosporidium oocysts than those of the upstream location (0.5 to 34.4 cysts per 100 liters and 0.5 to 34.4 oocysts per 100 liters). Peak concentrations of both parasites coincided with calving activity. Fecal samples, collected from cattle in both watersheds, showed 10% (3 of 30) in the BMID and 50% (5 of 10) in the VID watersheds to be Giardia positive. No Cryptosporidium-positive fecal samples were found. Giardia cysts isolated from the BMID watershed were repeatedly infective to gerbils in contrast to those from the VID watershed. The 10 BMID drinking water Giardia isolates retrieved into culture and biotyped showed zymodeme and karyotype heterogeneity. The differences in patterns of parasite contamination and cattle management practices contribute to the unique watershed characteristics observed between two areas which are topographically similar and geographically adjacent.
PMCID: PMC168065  PMID: 8702272
18.  Detritus-based assemblage responses under salinity stress conditions in a disused aquatic artificial ecosystem 
Aquatic Biosystems  2013;9:22.
Background
Despite the plethora of approaches, the sensitivity of the methods to measure the relationship between the abundance and biomass curves in stressed detritus-based ecosystems still remain to be refined. In this work, we report the comparison between biomass and abundance in a set of detritus-based macrozoobenthic assemblages located in six sampling pools with different salinity in an artificial aquatic ecosystem (disused Tarquinia Saltworks), using two diversity/dominance approaches (Abundance/Biomass Comparisons, or ABC, and Whittaker plots). We also evaluated the contribution of abundances and biomasses diversity (Simpson index) and nestedness, which measures the order by which macroinvertebrates colonized the detrital resource.
Results
The outputs obtained by both ABC curves and Whittaker plots highlight two different thresholds in assemblage structure: between about 44 and 50 practical salinity unit (psu) and between 50 and 87 psu, respectively. The first threshold was due to a turnover in taxon composition between assemblages, the second threshold (evidenced by Whittaker plots) was due to a change in taxon richness (lower in pools with higher salinity: i.e. > 50 psu). Moreover, a normal-shaped pattern in diversity (Simpson index) emerged, suggestive of an intermediate disturbance effect. The nested pattern did not show significant differences when considering the density and biomass of the sampled taxa, providing similar threshold of salinity in the relative contribution of macrozoobenthos on nestedness.
Conclusions
The use of detailed (ABC and Whittaker plots) and macroscopic (Simpson index and nestedness) approaches is proposed to identify thresholds in the structuring and functioning of detritus-based community of disused aquatic ecosystems: in particular, the inclusion of the parameter of biomass (scarcely utilized in community-based research) appears crucial. The responses of macrozoobenthic assemblages to the salinity stress conditions, in term of abundance and biomass, using a detritus food source (Phragmites australis leaves), may also highlight, by comparing macroscopic and detailed approaches, structuring and functioning patterns to consider for the management of disused artificial ecosystems.
doi:10.1186/2046-9063-9-22
PMCID: PMC4028869  PMID: 24308820
Leaf-detritus; Macrozoobenthos; Abundance/Biomass Comparisons; Whittaker plots; Simpson index; Nestedness; Patchy environment
19.  Invertebrate Metacommunity Structure and Dynamics in an Andean Glacial Stream Network Facing Climate Change 
PLoS ONE  2015;10(8):e0136793.
Under the ongoing climate change, understanding the mechanisms structuring the spatial distribution of aquatic species in glacial stream networks is of critical importance to predict the response of aquatic biodiversity in the face of glacier melting. In this study, we propose to use metacommunity theory as a conceptual framework to better understand how river network structure influences the spatial organization of aquatic communities in glacierized catchments. At 51 stream sites in an Andean glacierized catchment (Ecuador), we sampled benthic macroinvertebrates, measured physico-chemical and food resource conditions, and calculated geographical, altitudinal and glaciality distances among all sites. Using partial redundancy analysis, we partitioned community variation to evaluate the relative strength of environmental conditions (e.g., glaciality, food resource) vs. spatial processes (e.g., overland, watercourse, and downstream directional dispersal) in organizing the aquatic metacommunity. Results revealed that both environmental and spatial variables significantly explained community variation among sites. Among all environmental variables, the glacial influence component best explained community variation. Overland spatial variables based on geographical and altitudinal distances significantly affected community variation. Watercourse spatial variables based on glaciality distances had a unique significant effect on community variation. Within alpine catchment, glacial meltwater affects macroinvertebrate metacommunity structure in many ways. Indeed, the harsh environmental conditions characterizing glacial influence not only constitute the primary environmental filter but also, limit water-borne macroinvertebrate dispersal. Therefore, glacier runoff acts as an aquatic dispersal barrier, isolating species in headwater streams, and preventing non-adapted species to colonize throughout the entire stream network. Under a scenario of glacier runoff decrease, we expect a reduction in both environmental filtering and dispersal limitation, inducing a taxonomic homogenization of the aquatic fauna in glacierized catchments as well as the extinction of specialized species in headwater groundwater and glacier-fed streams, and consequently an irreversible reduction in regional diversity.
doi:10.1371/journal.pone.0136793
PMCID: PMC4550352  PMID: 26308853
20.  The Effects of Spatial Scale on Breakdown of Leaves in a Tropical Watershed 
PLoS ONE  2014;9(5):e97072.
The objective was to assess the effects of natural variation in the physical structure of the environment on biological communities and on the processing of Eucalyptus cloeziana and Inga laurina and to identify the controlling factors at different scales along stream order gradients. The study area consisted of 14 sampling sites distributed within a tropical watershed (1st, 2nd, 3rd and 4th order streams replicated in 4 sub-basins). Our samples consisted of 3 g of leaves of E. cloeziana (high-quality) and I. laurina (low-quality) placed in 252 bags with 10mm mesh (measured by the chemical composition of the detritus). Four samples of each leaf type were collected periodically (three times) over a period of 75–125 days and washed on a sieve to separate the invertebrates. A series of leaf disks were cut to determine ash-free dry mass, polyphenol, lignin, cellulose, total microbial biomass and fungal biomass, and the remaining material was oven-dried to determine the dry weight. We performed analyses within and between spatial scales (regional and local) to assess which watershed scale was the more import determinant of the leaf breakdown rate (k). The microbial and shredder were most influenced at the local scale (stream order). Shredders were influenced by microorganisms, with stronger interactions between them than were found to drive the k at the local scale. Moreover, differences in the overall k and abiotic variables were more strongly influenced at the regional scale (sub-basin), showing that the study scale alters the response of the studied variables. We found higher k values at higher values of water velocity, dissolved oxygen and temperature, all of which accelerate biological metabolism in response to variations on the regional scale. Watersheds with warmer microclimates and streams with higher nutrient levels and oxygen could be accelerating the ecosystem metabolism, independent of the detritus quality.
doi:10.1371/journal.pone.0097072
PMCID: PMC4014586  PMID: 24810918
21.  Differences Found in the Macroinvertebrate Community Composition in the Presence or Absence of the Invasive Alien Crayfish, Orconectes hylas 
PLoS ONE  2016;11(3):e0150199.
Introductions of alien species into aquatic ecosystems have been well documented, including invasions of crayfish species; however, little is known about the effects of these introductions on macroinvertebrate communities. The woodland crayfish (Orconectes hylas (Faxon)) has been introduced into the St. Francis River watershed in southeast Missouri and has displaced populations of native crayfish. The effects of O. hylas on macroinvertebrate community composition were investigated in a fourth-order Ozark stream at two locations, one with the presence of O. hylas and one without. Significant differences between sites and across four sampling periods and two habitats were found in five categories of benthic macroinvertebrate metrics: species richness, percent/composition, dominance/diversity, functional feeding groups, and biotic indices. In most seasons and habitat combinations, the invaded site had significantly higher relative abundance of riffle beetles (Coleoptera: Elmidae), and significantly lower Missouri biotic index values, total taxa richness, and both richness and relative abundance of midges (Diptera: Chironomidae). Overall study results indicate that some macroinvertebrate community differences due to the O. hylas invasion were not consistent between seasons and habitats, suggesting that further research on spatial and temporal habitat use and feeding ecology of Ozark crayfish species is needed to improve our understanding of the effects of these invasions on aquatic communities.
doi:10.1371/journal.pone.0150199
PMCID: PMC4795676  PMID: 26986207
22.  Hydrologic Response and Watershed Sensitivity to Climate Warming in California's Sierra Nevada 
PLoS ONE  2010;5(4):e9932.
This study focuses on the differential hydrologic response of individual watersheds to climate warming within the Sierra Nevada mountain region of California. We describe climate warming models for 15 west-slope Sierra Nevada watersheds in California under unimpaired conditions using WEAP21, a weekly one-dimensional rainfall-runoff model. Incremental climate warming alternatives increase air temperature uniformly by 2°, 4°, and 6°C, but leave other climatic variables unchanged from observed values. Results are analyzed for changes in mean annual flow, peak runoff timing, and duration of low flow conditions to highlight which watersheds are most resilient to climate warming within a region, and how individual watersheds may be affected by changes to runoff quantity and timing. Results are compared with current water resources development and ecosystem services in each watershed to gain insight into how regional climate warming may affect water supply, hydropower generation, and montane ecosystems. Overall, watersheds in the northern Sierra Nevada are most vulnerable to decreased mean annual flow, southern-central watersheds are most susceptible to runoff timing changes, and the central portion of the range is most affected by longer periods with low flow conditions. Modeling results suggest the American and Mokelumne Rivers are most vulnerable to all three metrics, and the Kern River is the most resilient, in part from the high elevations of the watershed. Our research seeks to bridge information gaps between climate change modeling and regional management planning, helping to incorporate climate change into the development of regional adaptation strategies for Sierra Nevada watersheds.
doi:10.1371/journal.pone.0009932
PMCID: PMC2848572  PMID: 20368984
23.  Relative Diversity and Community Structure of Ciliates in Stream Biofilms According to Molecular and Microscopy Methods▿ † 
Applied and Environmental Microbiology  2009;75(16):5261-5272.
Ciliates are an important component of aquatic ecosystems, acting as predators of bacteria and protozoa and providing nutrition for organisms at higher trophic levels. Understanding of the diversity and ecological role of ciliates in stream biofilms is limited, however. Ciliate diversity in biofilm samples from four streams subject to different impacts by human activity was assessed using microscopy and terminal restriction fragment length polymorphism (T-RFLP) analysis of 18S rRNA sequences. Analysis of 3′ and 5′ terminal fragments yielded very similar estimates of ciliate diversity. The diversity detected using microscopy was consistently lower than that suggested by T-RFLP analysis, indicating the existence of genetic diversity not apparent by morphological examination. Microscopy and T-RFLP analyses revealed similar relative trends in diversity between different streams, with the lowest level of biofilm-associated ciliate diversity found in samples from the least-impacted stream and the highest diversity in samples from moderately to highly impacted streams. Multivariate analysis provided evidence of significantly different ciliate communities in biofilm samples from different streams and seasons, particularly between a highly degraded urban stream and less impacted streams. Microscopy and T-RFLP data both suggested the existence of widely distributed, resilient biofilm-associated ciliates as well as ciliate taxa restricted to sites with particular environmental conditions, with cosmopolitan taxa being more abundant than those with restricted distributions. Differences between ciliate assemblages were associated with water quality characteristics typical of urban stream degradation and may be related to factors such as nutrient availability and macroinvertebrate communities. Microscopic and molecular techniques were considered to be useful complementary approaches for investigation of biofilm ciliate communities.
doi:10.1128/AEM.00412-09
PMCID: PMC2725482  PMID: 19561192
24.  Land Use Influences Niche Size and the Assimilation of Resources by Benthic Macroinvertebrates in Tropical Headwater Streams 
PLoS ONE  2016;11(3):e0150527.
It is well recognized that assemblage structure of stream macroinvertebrates changes with alterations in catchment or local land use. Our objective was to understand how the trophic ecology of benthic macroinvertebrate assemblages responds to land use changes in tropical streams. We used the isotope methodology to assess how energy flow and trophic relations among macroinvertebrates were affected in environments affected by different land uses (natural cover, pasture, sugar cane plantation). Macroinvertebrates were sampled and categorized into functional feeding groups, and available trophic resources were sampled and evaluated for the isotopic composition of 13C and 15N along streams located in the Cerrado (neotropical savanna). Streams altered by pasture or sugar cane had wider and more overlapped trophic niches, which corresponded to more generalist feeding habits. In contrast, trophic groups in streams with native vegetation had narrower trophic niches with smaller overlaps, suggesting greater specialization. Pasture sites had greater ranges of resources exploited, indicating higher trophic diversity than sites with natural cover and sugar cane plantation. We conclude that agricultural land uses appears to alter the food base and shift macroinvertebrate assemblages towards more generalist feeding behaviors and greater overlap of the trophic niches.
doi:10.1371/journal.pone.0150527
PMCID: PMC4774910  PMID: 26934113
25.  Topography and Land Cover of Watersheds Predicts the Distribution of the Environmental Pathogen Mycobacterium ulcerans in Aquatic Insects 
Background
An understanding of the factors driving the distribution of pathogens is useful in preventing disease. Often we achieve this understanding at a local microhabitat scale; however the larger scale processes are often neglected. This can result in misleading inferences about the distribution of the pathogen, inhibiting our ability to manage the disease. One such disease is Buruli ulcer, an emerging neglected tropical disease afflicting many thousands in Africa, caused by the environmental pathogen Mycobacterium ulcerans. Herein, we aim to describe the larger scale landscape process describing the distribution of M. ulcerans.
Methodology
Following extensive sampling of the community of aquatic macroinvertebrates in Cameroon, we select the 5 dominant insect Orders, and conduct an ecological niche model to describe how the distribution of M. ulcerans positive insects changes according to land cover and topography. We then explore the generalizability of the results by testing them against an independent dataset collected in a second endemic region, French Guiana.
Principal Findings
We find that the distribution of the bacterium in Cameroon is accurately described by the land cover and topography of the watershed, that there are notable seasonal differences in distribution, and that the Cameroon model does not predict the distribution of M. ulcerans in French Guiana.
Conclusions/Significance
Future studies of M. ulcerans would benefit from consideration of local structure of the local stream network in future sampling, and further work is needed on the reasons for notable differences in the distribution of this species from one region to another. This work represents a first step in the identification of large-scale environmental drivers of this species, for the purposes of disease risk mapping.
Author Summary
Many pathogens persist in the environment, and an understanding of where they are can assist in disease control, allowing us to identify areas of risk to local human populations. Herein, we use general linear models to describe the distribution of a particular environmental pathogen, Mycobacterium ulcerans, describing the landscape conditions correlated with the presence of this pathogen in local biota, and mapping the distribution of these habitats in a region of Cameroon, Africa. Our findings identify the importance of the watershed as a factor determining the distribution of the bacterium, where landscape conditions upstream of the sample site can influence the abundance of the bacterium in downstream sites. We find that the bacterium has notable seasonal changes in its distribution, between the wet and dry seasons, which may have implications for human health. We also discuss sensitivity of these models to extrapolation, finding that they work well in the African region and underperforming when extrapolated to another region in South America.
doi:10.1371/journal.pntd.0003298
PMCID: PMC4222759  PMID: 25375173

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