Phosphorus limitation of the freshwater benthic diatom Didymosphenia geminata determined by the frequency of dividing cells

1. Unlike other nuisance algal species, the freshwater benthic diatom Didymosphenia geminata typically forms blooms in low‐nutrient rivers. The negative association between D. geminata blooming behaviour and nutrient levels appears at both catchment and smaller scales. We conducted a series of trials in streamside experimental channels colonised with D. geminata using water from the D. geminata‐affected, oligotrophic Waitaki River, South Island, New Zealand to determine how elevated nitrate and phosphate concentrations affected D. geminata cell division. Because D. geminata blooms are typically most pronounced in unshaded waters, we also investigated the growth response to shading. In all experiments, we used the frequency of dividing cells (FDC) as a metric of cell division. 2. Concentrations of nitrate and dissolved reactive phosphorus (DRP) in the Waitaki River were very low (4 mg m^?3‐N and <1 mg m^?3 DRP). In pilot trials, substrata colonised by D. geminata were subjected to enrichment by either switching the water source toN‐ and P‐rich spring water or by adding a stock solution. Both trials resulted in periods of rapid cell division lasting at least 8 days. 3. Experimental addition of alone triggered an initial cell division which was not sustained. However, addition of alone or together with resulted in prolonged elevation in cell division indicating that the cell division rate was P‐limited. 4. Reduced light levels resulted in decreased FDC in D. geminata in both ambient and N, N + P and P‐enriched river water. 5. Stimulation of D. geminata division rate by addition of above ambient levels confirms that, while blooming behaviour is often associated with oligotrophic rivers, the cells divide faster with greater levels of phosphorus enrichment.     

Contrasting effects of low‐level phosphorus and nitrogen enrichment on growth of the mat‐forming alga Didymosphenia geminata in an oligotrophic river

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Ecological eustress? Nutrient supply,bloom stimulation and competition determine dominance of the diatom Didymosphenia geminata

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Is Didymosphenia geminata an introduced species in New Zealand? Evidence from trends in water chemistry,and chloroplast DNA

Defining the geographic origins of free‐living aquatic microorganisms can be problematic because many such organisms have ubiquitous distributions, and proving absence from a region is practically impossible. Geographic origins become important if microorganisms have invasive characteristics. The freshwater diatom Didymosphenia geminata is a potentially ubiquitous microorganism for which the recent global expansion of nuisance proliferations has been attributed to environmental change. The changes may include declines in dissolved reactive phosphorus (DRP) to low levels (e.g., <2 mg/m³) and increases in dissolved inorganic nitrogen (DIN) to >10 mg/m³ because both these nutrient conditions are associated with nuisance proliferations of D. geminata. Proliferations of D. geminata have been observed in South Island, New Zealand, since 2004. We aimed to address the ubiquity hypothesis for D. geminata in New Zealand using historical river water nutrient data and new molecular analyses. We used 15 years of data at 77 river sites to assess whether trends in DRP or DIN prior to the spread of D. geminata were consistent with a transition from a rare, undetected, species to a nuisance species. We used new sequences of chloroplast regions to examine the genetic similarity of D. geminata populations from New Zealand and six overseas locations. We found no evidence for declines in DRP concentrations since 1989 that could explain the spread of proliferations since 2004. At some affected sites, lowest DRP occurred before 2004. Trends in DIN also did not indicate enhanced suitability for D. geminata. Lack of diversity in the chloroplast intergenic regions of New Zealand populations and populations from western North America is consistent with recent dispersal to New Zealand. Our analyses did not support the proposal that D. geminata was historically present in New Zealand rivers. These results provide further evidence countering proposals of general ubiquity in freshwater diatoms and indicate that, as assumed in 2004, D. geminata is a recent arrival in New Zealand.     

ENVIRONMENTAL CONTROL OF STALK LENGTH IN THE BLOOM‐FORMING,FRESHWATER BENTHIC DIATOM DIDYMOSPHENIA GEMINATA (BACILLARIOPHYCEAE)1

Blooms of the freshwater stalked diatom Didymosphenia geminata (Lyngb.) M. Schmidt in A. Schmidt typically occur in oligotrophic, unshaded streams and rivers. Observations that proliferations comprise primarily stalk material composed of extracellular polymeric substances (EPS) led us to ask whether or not the production of excessive EPS is favored under nutrient‐limited, high‐light conditions. We conducted experiments in outdoor flumes colonized by D. geminata using water from the oligotrophic, D. geminata–affected Waitaki River, South Island, New Zealand, to determine the relationship between D. geminata stalk length, cell division rates, and light intensity under ambient and nutrient‐enriched conditions. Stalk lengths were measured in situ, and cell division rates were estimated as the frequency of dividing cells (FDC). FDC responded positively to increasing light intensity and to nutrient additions (N+P and P). Under ambient conditions, stalk length increased as light level increased except at low ambient light levels and temperature. Nutrient enrichment resulted in decreased stalk length and negative correlations with FDC, with this effect most evident under high light. Our results are consistent with the hypothesis that extensive stalk production in D. geminata occurs when cell division rates are nutrient limited and light levels are high. Thus, photosynthetically driven EPS production in the form of stalks, under nutrient‐limited conditions, may explain the development of very high biomass in this species in oligotrophic rivers. The responses of FDC and stalk length under nutrient‐replete conditions are also consistent with occurrences of D. geminata as a nondominant component of mixed periphyton communities in high‐nutrient streams.     

Effects of moderate ammonium enrichment on three submersed macrophytes under contrasting light availability

1. Increased ammonium concentrations and decreased light availability in a water column have been reported to adversely affect submersed vegetation in eutrophic waters worldwide. 2. We studied the chronic effects of moderate enrichment (NH₄–N: 0.16–0.25 mg L^?1) on the growth and carbon and nitrogen metabolism of three macrophytes (Ceratophyllum demersum, Myriophyllum spicatum and Vallisneria natans) under contrasting light availability in a 2‐month experiment. 3. The enrichment greatly increased the contents of free amino acids and nitrogen in the shoot / leaf of the macrophytes. This indicates that was the dominant N source for the macrophytes. 4. Soluble carbohydrate contents remained relatively stable in the shoot / leaf of the macrophytes irrespective of the treatments. Under ambient light, the starch contents in the shoot / leaf of C. demersum and M. spicatum increased with enrichment, whereas V. natans did not exhibit any change. The starch contents decreased in C. demersum, increased in M. spicatum and remained unchanged in V. natans after the combined treatment of enrichment and reduced light. 5. The enrichment did not affect the growth of the three macrophytes under the ambient light. However, it did suppress the growth of C. demersum and M. spicatum under the reduced light. The results indicate that a moderate enrichment was not directly toxic to the macrophytes although it might change their viability in eutrophic lakes in terms of the carbon and nitrogen metabolism.     

Relationships between microcystins and environmental parameters in 30 subtropical shallow lakes along the Yangtze River, China

1. A survey of 30 subtropical shallow lakes in the middle and lower reaches of the Yangtze River area in China was conducted during July–September in 2003–2004 to study how environmental and biological variables were associated with the concentration of the cyanobacterial toxin microcystin (MC). 2. Mean MC concentration in seasonally river‐connected lakes (SL) was nearly 33 times that in permanently river‐connected lakes (RL), and more than six times that in city lakes (NC) and non‐urban lakes (NE) which were not connected to the Yangtze River. The highest MC (8.574 μg L^?1) was detected in Dianshan Lake. 3. MC‐RR and MC‐LR were the primary toxin variants in our data. MC‐RR, MC‐YR and MC‐LR were significantly correlated with Chl a, biomass of cyanobacteria, Microcystis and Anabaena, indicating that microcystins were mainly produced by Microcystis and Anabaena sp. in these lakes. 4. Nonlinear interval maxima regression indicated that the relationships of Secchi depth, total nitrogen (TN) : total phosphorus (TP) and NH with MC were characterised by negative exponential curves. The relationships between MC and TN, TP, NO + NO were fitted well with a unimodal curve. 5. Multivariate analyses by principal component and classifying analysis indicated that MC was mainly affected by Microcystis among the biological factors, and was closely related with temperature among physicochemical factors.     

A rapid technique for assessing the suitability of areas for invasive species applied to New Zealand's rivers

Early responses to incursions of non‐indigenous species (NIS) into new areas include modelling and surveillance to define the organisms’ potential and actual distributions. For well‐studied invasive species, predictive models can be developed based on quantitative data describing environmental tolerances. In late 2004, an invasive freshwater diatom Didymosphenia geminata, an NIS for which we had no such quantitative data, was detected in a New Zealand river. We describe a procedure used to rapidly develop a classification of suitability for all New Zealand's rivers, based on two sources of information. First, from a review of the limited available literature and unpublished data, we determined that temperature, hydrological and substrate stability, light availability, and water pH were the most important environmental gradients determining D. geminata's broad‐scale distribution and capacity for establishing and forming blooms in rivers. The second information source was a GIS‐based river network developed for a national classification of New Zealand's rivers, with associated data describing environmental characteristics of each section of the network. We used six variables that were available for every section of the network as surrogates for the environmental gradients that determine suitability. We then determined the environmental distance of all the river sections in the network from our assessment of the optimal conditions conducive to D. geminata blooms. The analysis suggested that > 70% of New Zealand's river sections (stream order > 3) fell into the two highest suitability categories (on a five‐point scale). At the time of writing, D. geminata had spread to 12 catchments, all of which were within these two categories. The technique is applicable in initial responses to incursions of NIS where quantitative information is limited, and makes optimal use of available qualitative information. Our assessment contributed to evaluations of the potential ecological, social, and economic impacts of D. geminata and is currently being used to stratify site selection for ongoing surveillance.     

Are geothermal streams important sites of nutrient uptake in an agricultural and urbanising landscape (Rotorua,New Zealand)?

1. Lakes in the Rotorua region of New Zealand are affected by eutrophication from urbanisation and agricultural land use. Some lake tributaries contain geothermally influenced waters, and it is currently unknown whether geothermal tributaries are active sites of nutrient cycling or represent point sources of nutrients to the lakes. 2. Using government data sets, we characterised the physicochemical conditions of geothermal and non‐geothermal streams. We then measured ecosystem metabolism and reach‐scale uptake of nitrate (), ammonium () and phosphate () in summer 2010 (n = 8 streams). Finally, we used government data to compare annual nutrient flux from geothermal and non‐geothermal surface water inputs to Lake Rotoiti. 3. As expected, geothermal streams had higher temperature, conductivity and nutrient concentrations and lower pH. However, primary production, community respiration and uptake rates in geothermal streams were not different from those in their non‐geothermal counterparts. Uptake rates of were higher in geothermal streams, and uptake was below detection in geothermal streams, probably due to the saturation by naturally high concentrations. 4. A comparison of Lake Rotoiti inputs suggested that geothermal streams are not significant sources of and , while geothermal inputs of represent an average of 46% of total flux from Lake Rotoiti tributaries. 5. Despite their high temperature and low pH, geothermal streams are active sites of photosynthesis, respiration and and cycling, indicating dynamic biofilm communities. 6. Management options for geothermal streams, if any, should focus on retention (e.g. uptake or coupled nitrification and denitrification) but could prove challenging given the persistent, naturally occurring high flux.     

Microbial activities in the burrow environment of the potamal mayfly Ephoron virgo

1. The impact of burrowing larvae of Ephoron virgo (Ephemeroptera, Polymitarcidae) on sediment microbiology has not been previously investigated because of difficulties in sampling the sediment of large rivers under in situ conditions. Therefore, we conducted experiments in the on‐ship Ecological Rhine Station of the University of Cologne (Germany), in which ambient conditions of the River Rhine can be closely mimicked. 2. In two consecutive seasons, experimental flow channels were stocked with Ephoron larvae and continuously supplied with water taken directly from the River Rhine. Sediment from the immediate vicinity of Ephoron burrows (i.e. U‐shaped cavities reaching 10–80 mm deep into the sediment) and bulk sediment samples were analysed for (i) particulate organic matter content, (ii) microscale in situ distribution of O₂, NO, and NH, and (iii) potential activities of exoenzymes. 3. Sediment surrounding the Ephoron burrows had markedly higher organic matter contents and exoenzyme activities compared with the bulk sediment. Microsensor measurements demonstrated that local O₂ and NO penetration into the sediment were greatly enhanced by larval ventilation behaviour. Volumetric O₂ and NO turnover rates that were calculated from steady state concentration profiles measured directly in the burrow lining were considerably higher than at the sediment surface. 4. In the sediment of the fast flowing River Rhine Ephoron burrows are preferential sites of organic matter accumulation and dissolved oxidant penetration. Our data suggest that the burrows are surrounded by a highly active microbial community that responds to the inputs from the water column with elevated O₂ and NO turnover, and release of exoenzymes into the sediment pore water. Especially during periods of mass occurrence, the larvae of E. virgo may thus significantly contribute (i) to the ecological connection between the water column and the sediment and (ii) to biogeochemical processing of organic matter in the riverbed.     

The fate of assimilated nitrogen in streams: an in situ benthic chamber study

1. Nitrogen (N) processing in streams has been investigated using whole‐stream ¹⁵N addition experiments that, in general, have found that a large proportion of added nitrate removed from the water column appears to be assimilated by the stream benthos. The long‐term fate of this retained N is unknown, and of particular interest is the possibility that it becomes denitrified through coupled mineralisation–nitrification–denitrification processes (indirect denitrification). 2. We used in situ chambers to produce highly ¹⁵N‐enriched benthic biofilms and removed the chambers to allow biofilms to interact with ambient stream conditions. Nitrogen assimilation and direct denitrification were estimated from the first chamber deployment. Chambers were periodically reinstalled over 4 weeks to measure tracer ¹⁵N in ammonium (), nitrate () and dinitrogen (N₂), from which we estimated subsequent rates of biotic N transformations, including N mineralisation (ammonification), nitrification and indirect denitrification. We also estimated rates of depuration of ¹⁵N tracer from benthic biomass compartments. 3. Nitrate uptake was roughly equivalent in the sand and cobble habitats that dominated the stream. Direct denitrification (denitrification of from the water column) was an order of magnitude higher in cobble habitats than in sand habitats, accounting for c. 26 and 2% of total nitrate uptake in cobble and sand, respectively. 4. Mean residence times of actively cycling organic N in stream benthos (algae and microbes) were 16 days in cobble habitats and 9 days in sand habitats. The difference between habitat types was driven by the influence of N residence time in epilithic biofilms (18 days) on cobbles. 5. Release of enriched ¹⁵ was the primary flux of remineralised N, while release of enriched ¹⁵ was an order of magnitude less. We detected slight ¹⁵N enrichment in dissolved nitrogen gas (N₂) in post‐enrichment sampling, indicating that indirect denitrification was taking place. However, indirect denitrification accounted for <0.1% of the assimilated N. 6. These experiments agree with results of whole‐stream ¹⁵N additions, in that most added N was assimilated rather than directly denitrified. Assimilation was primarily a short‐term N retention mechanism in this stream, and indirect denitrification of assimilated N accounted for only a minor proportion of the observed ¹⁵N loss over time. 7. Remaining possible fates include export of N as particulate organic matter, which may lead to additional storage of assimilated N in downstream habitats, and consumption by grazers.     

Planktonic phosphorus pool sizes and cycling efficiency in coastal and interior British Columbia lakes

1. Limnologists have long acknowledged the importance of phosphorus (P) in determining the organism biomass and productivity of lake ecosystems. Despite a relatively large number of studies that have examined P cycling in lake ecosystems, there remain several substantial methodological issues that have impeded our understanding of P cycling in limnetic plankton communities. Two critical issues confronting ecologists are (1) a lack of precise measurements of the dissolved inorganic phosphorus (PO) and (2) accurate or complete measurements of dissolved P regeneration rates by plankton communities. 2. Here, we examine patterns of epilimnetic planktonic P pool sizes and turnover rates in eight lakes in British Columbia, Canada over a 2‐year period. We determine the concentrations and turnover times of P in various planktonic compartments (dissolved and various planktonic size fractions), using recently developed methods for estimating phosphate concentration and planktonic regeneration rates. 3. The pico‐ and nanoplankton size fraction (0.2–20 μm) played a central role in planktonic P cycling in lakes examined by this study. On average across lakes, pico‐ and nanoplankton contained >60% of the planktonic P, accounted for >90% PO uptake, and contributed 50% of the plankton community dissolved P regeneration rate. 4. PO concentrations determined by steady state bioassays (ssPO) were extremely low (87–611 pmol L⁻¹) and were 2–3 orders of magnitude less than simultaneously measured colorimetric soluble reactive phosphorus estimates. Lake ssPO concentrations increased linearly with total phosphorus (TP), and the slope of this relationship was approximately 1, indicating that PO remained a consistent proportion of the TP pool across a range of TP concentrations. 5. Turnover rates of the total planktonic P pool and the <20 μm pool became more rapid with increasing lake TP, indicating that, according to this metric, planktonic P cycling efficiency increased with TP concentrations. We also detected a significant relationship between particulate phosphorus (PP) <20 μm turnover time and seston N : P ratios, with PP <20 μm turnover times becoming slower with increasing seston N : P. These findings suggest that long‐standing conceptual models of nutrient cycling that predict slower cycling rates and decreasing cycling efficiency with increasing TP concentrations require further empirical examination. We postulate that patterns in lake P turnover and cycling efficiency are a result of complex interactions between plankton biomass and composition, and the ratios of multiple nutrients (C, N, P), rather than solely a function of the TP pool.     

Extreme seasonality of litter breakdown in an arctic spring‐fed stream is driven by shredder phenology,not temperature

1. Using degree‐days to calculate ‘temperature‐corrected’ breakdown rates is a useful approach for comparing litter breakdown across sites with different thermal regimes. We used an alternative approach to investigate the importance of temperature by quantifying seasonal patterns in litter breakdown in an arctic spring‐fed stream (Ivishak Spring, North Slope, Alaska) that experiences extreme seasonality in light availability and energy inputs while fluctuations in water temperature are relatively small. 2. We incubated mesh bags of senesced Salix alaxensis litter in Ivishak Spring for successive c. 30‐day periods for 2 years. During our study, water temperature was very stable [5.7 ± 0.03 °C (daily mean ± 1 SE), range 3.7–7.8 °C]. Discharge was only slightly more variable (mean 112 ± 1 L s^?1, range 66–206 L s^?1), with lowest values occurring in late winter. Dissolved nutrient concentrations were low (52–133 μg L^?1, <1–3 μg L^?1, <1–6 μg L^?1 soluble reactive phosphorus) and also showed evidence of seasonality (i.e. highest values in winter). 3. Litter breakdown rates were sharply seasonal, ranging from <0.01 day^?1 in mid‐summer to >0.05 day^?1 in mid‐winter. Invertebrate assemblage structure in litter bags showed pronounced seasonal cyclicity; total invertebrate biomass peaked in summer. Biomass of two dominant shredders (the nemourid stonefly Zapada haysi and the limnephilid caddisfly Ecclisomyia conspersa) showed the opposite trend, however, with mid‐winter peaks in both population biomass and cohort growth rates that closely matched those we observed in litter mass loss. 4. Water temperature appeared to have negligible influence on litter breakdown rates in our study. Seasonal shifts in nutrient uptake may have increased rates of microbial activity in winter. The processing of litter inputs in Ivishak Spring, however, appeared to be most tightly coupled to shredder phenology. Our results demonstrate that extreme seasonality in the processing of allochthonous detritus can occur even in the absence of substantial temperature variation, driven by the activity of shredder taxa that have evolved to take advantage of pulsed organic matter inputs.     

Relative influence of bioturbation and predation on organic matter processing in river sediments: a microcosm experiment

1. Our objective was to measure the effects of bioturbation and predation on the physical characteristics and biogeochemical processes in river sediments. 2. We investigated the impacts of tubificid worms tested separately and together with an omnivore (Gammarus pulex), which does feed on tubificids, on sediment distribution, water flux, sediment organic carbon, biofilm biomass and microbial activities, and the concentrations of dissolved oxygen, dissolved organic carbon, PO, NO, NO and NH in slow filtration sand–gravel columns. We hypothesised that gammarids, which exploit the top 2–3 cm of the sediment, would modify the impact of worms at the sediment surface. 3. In experiments both with and without gammarids, bioturbation by the tubificids modified both the distribution of surface particles in the sediment column and water flux. In addition, microbial aerobic (oxygen consumption) and anaerobic (denitrification and fermentative decomposition of organic matter) processes in the sediment were stimulated in the presence of tubificid worms. However, G. pulex did not affect either the density or bioturbation activity of the tubificid worms. 4. Bioturbation by the benthos can be a major process in river habitats, contributing to the retention of organic matter in sediment dynamics. The presence of at least one predator had no effect on bioturbation in sediments. In such systems, physical heterogeneity may be sufficient for tubificids to escape from generalist predators, though more specialised ones might have more effect.     

Nitrogen cycling and dynamics in a macrophyte‐rich stream as determined by a release

1. A tracer release study was conducted in a macrophyte‐rich stream, the River Lilleaa in Denmark. The objectives of the study were to compare uptake rates per unit area of by primary producers and consumers in macrophyte and non‐macrophyte habitats, estimate whole‐stream uptake rates of and compare this to other stream types, and identify the pathways and estimate the rate at which enters the food web in macrophyte and non‐macrophyte habitats. 2. Macrophyte habitats had four times higher primary uptake rates and an equal uptake rate by primary consumers per unit habitat area as compared to non‐macrophyte habitats. These rates represent the lower limit of potential macrophyte effects because the rates will be highly dependent on macrophyte bed height and mean bed height in the River Lilleaa was low compared to typical bed heights in many lowland streams. Epiphytes accounted for 30% of primary uptake in macrophyte habitats, illustrating a strong indirect effect of macrophytes as habitat for epiphytes. N flux per unit habitat area from primary uptake compartments to primary consumers was four times lower in macrophyte habitats compared to non‐macrophyte habitats, reflecting much greater biomass accrual in macrophyte habitats. Thus, we did not find higher N flux from macrophyte habitats to primary consumers compared to non‐macrophyte habitats. 3. Whole‐stream uptake rate was 447 mgN m^?2 day^?1. On a habitat‐weighted basis, fine benthic organic matter (FBOM) accounted for 72% of the whole‐stream uptake rate, and macrophytes and epiphytes accounted for 19 and 8%, respectively. 4. We had expected a priori relatively high whole‐stream N uptake in our study stream compared to other stream types mainly due to generally high biomass and the macrophyte’s role as habitat for autotrophic and heterotrophic organisms, but our results did not confirm this. In comparison with other release study streams, we conclude that nutrient concentration is the overall controlling factor for N uptake rates across streams, mostly as a result of high biomass of primary uptake compartments in streams with high nutrient concentrations in general and not in macrophyte streams in particular. 5. Our results indicate that macrophytes play an important role in the longer‐term retention of N and thus a decrease in net downstream transport during the growing season compared to streams without macrophytes, through direct and indirect effects on the stream reach. Direct effects are high uptake efficiency, low turnover rate (partly due to no direct feeding on macrophytes) and high longevity. An indirect effect is increased sedimentation of FBOM in macrophytes compared to non‐macrophyte habitats and streams which possibly also increase denitrification. Increased retention with macrophyte presence would decrease downstream transport during the growing season and thus the N loading on downstream ecosystems.     

Distribution of benthic diatoms in U.S. rivers in relation to conductivity and ionic composition

• 1 We quantified the relationships between diatom relative abundance and water conductivity and ionic composition, using a dataset of 3239 benthic diatom samples collected from 1109 river sites throughout the U.S.A. [U.S. Geological Survey National Water‐Quality Assessment (NAWQA) Program dataset]. This dataset provided a unique opportunity to explore the autecology of freshwater diatoms over a broad range of environmental conditions.
• 2 Conductivity ranged from 10 to 14 500 μS cm^?1, but most of the rivers had moderate conductivity (interquartile range 180–618 μS cm^?1). Calcium and bicarbonate were the dominant ions. Ionic composition, however, varied greatly because of the influence of natural and anthropogenic factors.
• 3 Canonical correspondence analysis (CCA) and Monte Carlo permutation tests showed that conductivity and abundances of major ions (HCO + CO, Cl^?, SO, Ca²⁺, Mg²⁺, Na⁺, K⁺) all explained a statistically significant amount of the variation in assemblage composition of benthic diatoms. Concentrations of HCO + CO and Ca²⁺ were the most significant sources of environmental variance.
• 4 The CCA showed that the gradient of ionic composition explaining most variation in diatom assemblage structure ranged from waters dominated by Ca²⁺ and HCO + CO to waters with higher proportions of Na⁺, K⁺, and Cl^?. The CCA also revealed that the distributions of some diatoms correlated strongly with proportions of individual cations and anions, and with the ratio of monovalent to divalent cations.
• 5 We present species indicator values (optima) for conductivity, major ions and proportions of those ions. We also identify diatom taxa characteristic of specific major‐ion chemistries. These species optima may be useful in future interpretations of diatom ecology and as indicator values in water‐quality assessment.

     

Factors controlling streambed coverage of <Emphasis Type="Italic">Didymosphenia geminata</Emphasis> in two regulated streams in the Colorado Front Range

Didymosphenia geminata is a stalk-forming freshwater diatom which was historically found primarily in oligotrophic lakes and streams, but has recently become a nuisance species in many lotic systems worldwide. In the last 5–8 years, D. geminata has become established in Boulder Creek and South Boulder Creek, two regulated montane streams in the Front Range of the Colorado Rocky Mountains. Factors that may influence the growth of D. geminata were monitored during the summer of 2006. D. geminata abundance decreased in Boulder Creek after an unusual flood event caused by 3 days of sustained rainfall in the headwaters of the watershed. However, within a week, coverage had been restored to pre-flood levels. Variations in D. geminata abundance among sites were found to be negatively correlated with total dissolved phosphorus concentrations and bed movement, as measured by Shields stress. In contrast, D. geminata abundance was not significantly correlated with temperature, conductivity, pH, total suspended solids, or dissolved inorganic nitrogen. Our results suggest that bed movement may be a dominant scouring mechanism that acts to control the growth and distribution of D. geminata. The potential role of total dissolved phosphorus and bed movement in decreasing D. geminata coverage adds to the limited base of knowledge regarding controls on the growth and distribution of this species, and could be investigated by researchers studying D. geminata blooms in other stream ecosystems. Handling editor: J. Saros     

Disentangling the stream community impacts of <Emphasis Type="Italic">Didymosphenia geminata</Emphasis>: How are higher trophic levels affected?

Human activities frequently result in either intentional or unintentional introductions of species to new locations, and freshwater environments worldwide are particularly vulnerable to species invasions. An introduced freshwater diatom, Didymosphenia geminata, was first discovered in New Zealand in 2004 but there was limited research available to predict the drivers of D. geminata biomass and how biomass variability might influence higher trophic levels (e.g. invertebrates and fish). We examined the effect of D. geminata biomass on benthic invertebrates, invertebrate drift and fish communities in 20 rivers in New Zealand with variable hydrology, physical habitat and water chemistry. Variation in D. geminata biomass was best explained by a model that showed D. geminata biomass increased with time since the last flow event exceeding three times the median annual discharge and decreasing concentration of dissolved reactive phosphorus. Analyses of biotic responses showed that high D. geminata biomass did not affect either invertebrate or fish diversity but altered the structure of benthic communities, changed the composition of drifting invertebrate communities and reduced fish biomass by 90 %, particularly trout. A partial least squares path model was used to disentangle both direct and indirect effects of D. geminata on fish communities and showed D. geminata had a significant negative direct effect on fish communities. This is the first study to show how the potential effects of the introduced diatom D. geminata can impact fish communities and has shown that D. geminata impacts fish both directly and indirectly through changes in their invertebrate prey community.     

Periphyton‐based transfer functions to assess ecological imbalance and management of a subtropical ombrotrophic peatland

1. To assess the biological status and response of aquatic resources to management actions, managers and decision‐makers require accurate and precise metrics. This is especially true for some parts of the Florida Everglades where multiple stressors (e.g. hydrologic alterations and eutrophication) have resulted in a highly degraded and fragmented ecosystem. Biological assessments are required that directly allow for the evaluation of historical and current status and responses to implantation of large‐scale restoration projects. 2. Utilising periphyton composition and water‐quality data obtained from long‐term (15 years) monitoring programmes, we developed calibrated and verified periphyton‐based numerical models (transfer functions) that could be used to simultaneously assess multiple stressors affecting the Everglades peatland (e.g. salinity, nitrogen and phosphorus). Periphyton is an ideal indicator because responses to stressors are rapid and predictable and possess valued ecological attributes. 3. Weighted averaging partial least squares regression was used to develop models to infer water‐quality concentrations from 456 samples comprising 319 periphyton taxa. Measured versus periphyton‐inferred concentrations were strongly related for log‐transformed salinity ( = 0.81; RMSEP 0.15 mg L^?1) and log‐transformed total phosphorus (TP; = 0.70; RMSEP 0.18 mg L^?1), but weakly related for total Kjeldahl nitrogen (TKN) ( = 0.46; RMSEP 0.12 mg L^?1). Validation results using an independent 455 sample data set were similar (log(salinity) r² = 0.78, log(TP) r² = 0.65 and log(TKN) r² = 0.38). 4. Water Conservation Area 1 (WCA‐1), a large ombrotrophic subtropical peatland impacted by multiple water‐quality stressors that has undergone major changes in water management, was used as a case study. The models were applied to a long‐term periphyton data set to reconstruct water‐quality trends in relation to restoration efforts to reduce nutrient loading to the Everglades. The combination of biologically inferred TP and salinity was used to identify the ecological status of periphyton assemblages. Periphyton assemblages were ecologically imbalanced with respect to salinity and TP. Salinity imbalance varied spatially and temporally, whereas TP was spatially restricted. Imbalances caused by water management were owing to salinity more so than to TP. 5. The transfer functions developed for the Everglades are trait‐based quantitative numerical methods and are ideal because the abundances of species are modelled numerically in relation to a stressor. The resulting inferred value is a numerical representation of the stressor’s effect on biological condition that can be compared against the management of the stressor independent of other factors. The benefits are that biological lags or hysteresis events can easily be identified and environmental conditions can be estimated when measurements are lacking. Reporting biological assessments in terms of well‐defined water‐quality metrics (e.g. numeric criterion) increased the communicative ability of the assessment. The use of multiple metrics to assess ecological imbalance increased the ability to identify probable causes.     

Regulation of nitrous oxide emission associated with benthic invertebrates

1. A number of freshwater invertebrate species emit N₂O, a greenhouse gas that is produced in their gut by denitrifying bacteria (direct N₂O emission). Additionally, benthic invertebrate species may contribute to N₂O emission from sediments by stimulating denitrification because of their bioirrigation behaviour (indirect N₂O emission). 2. Two benthic invertebrate species were studied to determine (i) the dependence of direct N₂O emission on the preferred diet of the animals, (ii) the regulation of direct N₂O emission by seasonally changing factors, such as body size, temperature and availability and (iii) the quantitative relationship between direct and indirect N₂O emission. 3. Larvae of the mayfly Ephemera danica, which prefer a bacteria‐rich detritus diet, emitted N₂O at rates of up to 90 pmol Ind.^?1 h^?1 under in situ conditions and 550 pmol Ind.^?1 h^?1 under laboratory conditions. In contrast, larvae of the alderfly Sialis lutaria, which prefer a bacteria‐poor carnivorous diet, emitted N₂O at invariably low rates of 0–20 pmol Ind.^?1 h^?1. The N₂O emission rate of E. danica larvae was positively correlated with seasonally changing factors (body size, temperature and availability). Direct N₂O emission by E. danica larvae was limited by low temperature in winter, larval development in spring and low availability in summer. 4. Both E. danica and the non‐emitting S. lutaria increased the total N₂O and N₂ emission from sediment in a density‐dependent manner. While N₂O directly emitted by benthic invertebrates can be partially consumed in the sediment (E. danica), non‐emitting species can still indirectly contribute to total N₂O emission from sediment (S. lutaria).