Effects of shading on calcareous benthic periphyton in a short-hydroperiod oligotrophic wetland (Everglades, FL, USA)

The effects of shade on benthic calcareous periphyton were tested in a short-hydroperiod oligotrophic subtropical wetland (freshwater Everglades). The experiment was a split-plot design set in three sites with similar environmental characteristics. At each site, eight randomly selected 1-m² areas were isolated individually in a shade house, which did not spectrally change the incident irradiance but reduced it quantitatively by 0, 30, 50, 60, 70, 80, 90 and 98%. Periphyton mat was sampled monthly under each shade house for a 5 month period while the wetland was flooded. Periphyton was analyzed for thickness, DW, AFDW, chlorophyll a (chl a) and incubated in light and dark BOD bottles at five different irradiances to assess its photosynthesis–irradiance (PI) curve and respiration. The PI curves parameters P _max, I _k and eventually the photoinhibition slope (β) were determined following non-linear regression analyses. Taxonomic composition and total algal biovolume were determined at the end of the experiment. The periphyton composition did not change with shade but the PI curves were significantly affected by it. I _k increased linearly with increasing percent irradiance transmittance (%IT = 1−%shade). P _max could be fitted with a PI curve equation as it increased with %IT and leveled off after 10%IT. For each shade level, the PI curve was used to integrate daily photosynthesis for a day of average irradiance. The daily photosynthesis followed a PI curve equation with the same characteristics as P _max vs. %IT. Thus, periphyton exhibited a high irradiance plasticity under 0–80% shade but could not keep up the same photosynthetic level at higher shade, causing a decrease in daily GPP at 98% shade levels. The plasticity was linked to an increase in the chl a content per cell in the 60–80% shade, while this increase was not observed at lower shade likely because it was too demanding energetically. Thus, chl a is not a good metric for periphyton biomass assessment across variously shaded habitats. It is also hypothesized that irradiance plasticity is linked to photosynthetic coupling between differently comprised algal layers arranged vertically within periphyton mats that have different PI curves.     

Diatom Assemblages and their Associations with Environmental Variables in Oregon Coast Range Streams, USA

The Oregon Coast Range, rich in natural resources, is under increasing pressure from rapid development. The purpose of this study was to examine diatom species patterns in relation to environmental variables in streams of this region. Diatoms, water quality, physical habitat and watershed characteristics were assessed for 33 randomly selected stream sites. Watershed size, elevation, geology, vegetation and stream morphology varied substantially among sites. Streams were characterized by dilute water chemistry and a low percent of fine substrate. A total of 80 diatom taxa were identified. Taxa richness was low throughout the region (median 15, range 10–26). Assemblages were dominated by two adnate species, Achnanthidium minutissimum and Achnanthes pyrenaicum. Diatoms sensitive to organic pollution dominated the assemblages at all sites (median 85%). Non-metric multidimensional scaling (NMDS) and correlational analysis showed quantitative relationships between diatom assemblages and environmental variables. NMDS axes were significantly correlated with watershed area, watershed geology, conductivity, total nitrogen, total solids and stream width. Diatom-based site classification (Two-way Indicators Species Analysis, (TWINSPAN)) yielded 4 discrete groups that displayed weak correlations with environmental variables. When stream sites were classified by dominant watershed geology, overall diatom assemblages between groups were significantly different (Analysis of Similarity (ANOSIM) global R = 0.19, p < 0.05). Our results suggest that streams in the coastal region are in relatively good condition. High natural variability in stream conditions in the Oregon Coast Range ecoregion may obscure quantitative relationships between environmental variables and diatom assemblages. A bioassessment protocol that classifies sites by major landscape variables and selects streams along the major human disturbance gradient might allow for detection of early signs of human disturbance in environmentally heterogeneous regions, such as the Pacific Northwest.     

The non-indigenous diatom Didymosphenia geminata alters benthic communities in New Zealand rivers

1. Blooms of the benthic, stalked diatom Didymosphenia geminata were first observed in New Zealand in 2004. Since then, D. geminata has spread to numerous catchments in the South Island and is also spreading in its native range. The species is a rare example of an invasive alga in lotic systems.
2. Ecosystem effects may be expected as D. geminata attains unusually high biomass in rivers. We examined data from three independent studies in three South Island, New Zealand, rivers for evidence of effects on periphyton biomass and benthic invertebrate communities.
3. The combined results confirmed that the presence of D. geminata was associated with greatly increased periphyton biomass and, in most cases, increased invertebrate densities. We also recorded shifts in community composition, dominated by increased densities of Oligochaeta. Chironomidae, Cladocera and Nematoda also generally increased in density with D. geminata . Significant increases or declines in other invertebrate taxa were inconsistent among rivers.
4. In all three studies, increased spatial invertebrate community homogeneity was associated with high D. geminata biomass at the within-river scale. However, no declines in taxon richness or diversity were detected.
5. Although ecosystem effects of D. geminata on existing periphyton biomass and invertebrate communities are measurable, no inferences can be made from the present data about effects on higher trophic levels (fish).     

Stream geomorphology regulates the effects on periphyton of ecosystem engineering and nutrient enrichment by Pacific salmon

1. Pacific salmon (Oncorhynchus spp.) returning to streams deliver substantial quantities of nutrients (nitrogen and phosphorus) that may stimulate primary production. Salmon can also affect the phytobenthos negatively via physical disturbance during nest excavation, a process that may counteract the positive effects of salmon‐derived nutrients on benthic algae. The ability of salmon to disturb benthic habitats may be a function of substratum particle size, and therefore, the geomorphology of streams could determine the net effect of salmon on benthic communities. 2. Based on surveys of 17 streams in southwest Alaska before the salmon run and during peak salmon density, we identified size thresholds for the disturbance of substratum particles by salmon and classified particles as vulnerable (<60 mm B‐axis), invulnerable (>110 mm) or transitional (61–110 mm). At the scale of individual rocks, algal biomass on vulnerable substrata decreased at peak spawning (relative to values before the run) as a power function of salmon density; transitional and invulnerable substrata showed no quantifiable pattern. However, invulnerable substrata in streams with more than 0.11 salmon m^?2 showed net algal accrual, or relatively smaller declines in algal biomass, than vulnerable substrata, indicating that large rocks provide refuge for benthic algae from salmon disturbance. 3. We expected that streams with proportionally larger rocks would respond positively to salmon at the whole‐stream scale, after accounting for the relative abundance of rocks of different sizes within streams. Invulnerable rocks made up only 0–12% of the total substratum particle size distribution in salmon‐bearing streams, however, and algal accrual on invulnerable substrata did not outweigh the strong disturbance effects on the more spatially extensive vulnerable substrata. The change in whole‐stream benthic algal biomass among streams was negatively related to salmon density. 4. Stable isotopes of nitrogen (δ¹⁵N) were used to track nutrients from salmon into benthic biota. Periphyton δ¹⁵N on rocks of all size classes was higher at peak salmon spawning than before the salmon run, indicating the uptake of salmon‐derived nitrogen. Peak δ¹⁵N values were positively related to salmon abundance and followed a two‐isotope mixing relationship. The per cent of N from salmon in periphyton was also related to salmon density and was best explained by a saturating relationship. Spring δ¹⁵N was unrelated to salmon returns in the previous year, suggesting little annual carryover of salmon nutrients.     

Applying the light : nutrient hypothesis to stream periphyton

1. The light : nutrient hypothesis (LNH) states that algal nutrient content is determined by the balance of light and dissolved nutrients available to algae during growth. Light and phosphorus gradients in both laboratory and natural streams were used to examine the relevance of the LNH to stream periphyton. Controlled gradients of light (12–426 μmol photons m^?2 s^?1) and dissolved reactive phosphorus (DRP, 3–344 μg L^?1) were applied experimentally to large flow‐through laboratory streams, and natural variability in canopy cover and discharge from a wastewater treatment facility created gradients of light (0.4–35 mol photons m^?2 day^?1) and DRP (10–1766 μg L^?1) in a natural stream. 2. Periphyton phosphorus content was strongly influenced by the light and DRP gradients, ranging from 1.8 to 10.7 μg mg AFDM^?1 in the laboratory streams and from 2.3 to 36.9 μg mg AFDM^?1 in the natural stream. Phosphorus content decreased with increasing light and increased with increasing water column phosphorus. The simultaneous effects of light and phosphorus were consistent with the LNH that the balance between light and nutrients determines algal nutrient content. 3. In experiments in the laboratory streams, periphyton phosphorus increased hyperbolically with increasing DRP. Uptake then began levelling off around 50 μg L^?1. 4. The relationship between periphyton phosphorus and the light : phosphorus ratio was highly nonlinear in both the laboratory and natural streams, with phosphorus content declining sharply with initial increases in the light : phosphorus ratio, then leveling off at higher values of the ratio. 5. Although light and DRP both affected periphyton phosphorus content, the effects of DRP were much stronger than those of light in both the laboratory and natural streams. DRP explained substantially more of the overall variability in periphyton phosphorus than did light, and light effects were evident only at lower phosphorus concentrations (≤25 μg L^?1) in the laboratory streams. These results suggest that light has a significant negative effect on the food quality of grazers in streams only under a limited set of conditions.     

Effect of fire on benthic algal assemblage structure and recolonization in intermittent streams

Abstract: Dry biofilm on rocks and other substrata forms an important drought refuge for benthic algae in intermittent streams following the cessation of flow. This dry biofilm is potentially susceptible to disturbance from bushfires, including direct burning and/or scorching and damage from radiant heat, particularly when streams are dry. Therefore, damage to dry biofilms by fire has the potential to influence algal recolonization and assemblage structure in intermittent streams following commencement of flow. The influence of fire on benthic algal assemblages and recolonization was examined in intermittent streams of the Grampians National Park, Victoria, Australia, using a field survey and manipulative field experiment. The field survey compared assemblages in two intermittent streams within a recently burnt area (within 5 months of the fire) with two intermittent streams within an unburnt area. The two burnt streams were still flowing during the fire so most biofilms were not likely to be directly exposed to flames. Considerable site‐to‐site and stream‐to‐stream variation was detected during the field survey, which may have obscured potential differences attributable to indirect effects of the fire. The manipulative field experiment occurred in two intermittent streams and consisted of five treatments chosen to replicate various characteristics of bushfires that may influence dry biofilms: dry biofilm exposed directly to fire; dry biofilm exposed to radiant heat; dry biofilm exposed to ash; and two procedural controls. After exposure to the different treatments, rocks were replaced in the streams and algae were sampled 7 days after flow commenced. Differences occurred across treatments, but treatment differences were inconsistent across the two streams. For example, direct exposure to fire reduced the abundance of recolonizing algae and altered assemblage structure in both streams, while radiant heat had an effect on assemblage structure in one stream only. The manipulative field experiment is likely to have represented the intensity of a small bushfire only. Nonetheless, significant differences across treatments were detected, so these experimental results suggest that fire can damage dry biofilms, and hence, influence algal recolonization and assemblage structure in intermittent streams.     

Effects of nutrients and light on periphyton biomass and nitrogen uptake in Mediterranean streams with contrasting land uses

1. Nutrient diffusing substrata (NDS) were used to determine the relative importance of nutrients and light as potential limiting factors of periphyton biomass and nitrogen (N) uptake in Mediterranean streams subjected to different human impacts. The nutrients examined were phosphorus (P) and N, and we also further differentiated between the response of periphyton communities to N species (i.e. NO₃‐N and NH₄‐N). To examine the effect of light and nutrients on periphyton biomass, chlorophyll a accrual rates on NDS located at open and closed canopy sites were compared. The effect of nutrient availability on periphyton uptake was measured by ¹⁵N changes on the NDS after NO₃‐¹⁵N short‐term nutrient additions. 2. Results show that light was the main factor affecting algal biomass in the study streams. Algal biomass was in general higher at open than at closed canopy sites. Nutrient availability, as simulated with the NDS experiments, did not enhance algal biomass accrual in either of the 2 light conditions. 3. In the control treatments (i.e. ambient concentrations), periphyton NO₃‐N uptake rates increased and C : N molar ratios decreased consistently with increases in N availability across streams. NO₃‐N uptake rates were altered when ambient N concentrations were increased artificially in the N amended NDS. Periphyton assemblages growing on N enriched substrata seemed to preferentially take up N diffusing from the substratum rather than N from the water column. This response differed among streams, and depended on ambient N availability. 4. Periphyton biomass was not significantly different between substrata exposed to the two forms of available N sources. Nonetheless, we found differences in the effects of both N sources on the uptake of N from the water column. NH₄‐N seemed to be the preferred source of N for periphyton growing on NDS. 5. Results suggest that the effect of riparian zones on light availability, although seldom considered by water managers, may be more important than nutrients in controlling eutrophication effects derived from human activities. Finally, our results confirm that not only increases in concentration, but also stoichiometric imbalances should be considered when examining N retention in human altered streams.     

PERIPHYTIC DIATOM ASSEMBLAGES FROM ULTRA‐OLIGOTROPHIC AND UV TRANSPARENT LAKES AND PONDS ON VICTORIA ISLAND AND COMPARISONS WITH OTHER DIATOM SURVEYS IN THE CANADIAN ARCTIC1

Periphytic diatoms are potentially powerful indicators of environmental change in climatically‐sensitive high latitude regions. However, only a few studies have examined their taxonomic and ecological characteristics. We identified and enumerated diatom assemblages from sediment, rock, and moss habitats in 34 ultra‐oligotrophic and highly transparent lakes and ponds on Victoria Island, Arctic Canada. The similar limnological characteristics of the sites allowed us to examine the influence of habitat, independent of water chemistry, on the diatom assemblages. As is typical in shallow arctic water bodies, benthic taxa, including species of Achnanthes, Caloneis, Cymbella, Navicula, and Nitzschia, were most widely represented. Minor gradients in our measured environmental variables did not significantly explain any variance in diatom species, but there were marked differences in diatom assemblages among sites. Pond ephemerality seems to explain some diatom variation, because aerophilic taxa such as Achnanthes kryophila Petersen and A. marginulata Grunow were dominant in shallow sites that had undergone appreciable reductions in volume. We identified several taxa that exhibited strong habitat preferences to sediment, moss, or rock substrates and also found significant differences (P < 0.01) in diatom composition among the three habitats. In comparisons with three similar diatom surveys extending over 1200 km of latitude, we determined that surface sediment assemblages differed significantly (P < 0.001) among all regions examined. Diatom species diversity was inversely related to latitude, a result likely explained by differences in the lengths of growing seasons. These data contribute important ecological information on diatom assemblages in arctic regions and will aid in the interpretation of environmental changes in biomonitoring and paleolimnological studies.     

RESPONSE OF A LENTIC PERIPHYTON COMMUNITY TO NUTRIENT ENRICHMENT AT LOW N:P RATIOS1

We examined the effect of nitrogen:phosphorus (N:P) ratios and nutrient concentrations on periphyton when nutrients (N and P) are provided in excess. A gradient of seven N:P ratios ranging from 7.5:1 to 1:7.5 and each at three absolute concentrations, was established using nutrient‐releasing substrata placed in a meso‐oligotrophic lake. Differences in total algal biovolume among nutrient ratios were significant (analysis of covariance [ANCOVA]) when P concentration was entered as the co‐variate. In addition, total algal biovolume was significantly correlated with N concentration but not P. To further evaluate the relationship between nutrient ratios and biovolume, we analyzed (using four 1‐way analysis of variances [ANOVAs]) four subsets of data defined as a series of treatments where one nutrient concentration remained relatively constant as the other changed creating different N:P ratios. Ratios of data subsets ranged from 1:1 to 7.5:1 and 1:1 to 1:7.5 with low and high concentrations of both series. Only diatom biovolume varied with ratio but these differences are most likely related to increased green algal abundance. Species richness and diversity differed among N:P ratios (ANCOVA) when P concentration was used as the co‐variate. Stigeoclonium tenue (Ag.) Gomont, which generally accounted for the increase in green algal abundance, varied with nutrient ratio (ANCOVA) when P was the co‐variate. Based on the ANCOVAs, correlations, and one‐way ANOVAs, periphyton in this system appears to be affected by N concentration but not by N:P treatment ratios under nutrient‐rich conditions. When compared with previous studies, these data also suggest that the response of periphyton to in situ treatments constructed with nutrient‐releasing substrata vary between years.