Alterations in the biomass-specific productivity of periphyton assemblages mediated by fish grazing

1. In an experimental flume, we examined the effects of a biomass reduction and alteration of taxonomic composition, because of grazing by the fish Plecoglossus altivelis, on the net biomass accumulation of periphyton. 2. Grazed and ungrazed assemblages with different biomass and taxonomic composition were first prepared in fish enclosures and exclosures, respectively. These assemblages were then set out in the flume and incubated for 2 days under grazing‐free conditions to examine (i) the relationship between biomass and biomass accumulation rate and (ii) the effect of taxonomic composition on the relationship between these two. 3. The grazed and ungrazed assemblages were dominated by upright filamentous cyanobacteria and diatoms, respectively. The rate of biomass accumulation decreased with increasing periphyton biomass in both the grazed and ungrazed assemblages, and was lower in the grazed than the ungrazed assemblages at any biomass level. 4. The results showed that the reduction in biomass and the alteration of taxonomic composition due to fish grazing have opposite effects on biomass‐specific productivity. Biomass accumulation rate increased in response to biomass reduction, although a shift in dominance from diatoms to upright filamentous cyanobacteria decreased the overall productivity of the periphyton.     

The seasonal biomass and productivity of the submerged macrophytes in a polluted Wisconsin stream

SUMMARY 1. We measured biomass and light/dark bottle productivity of macrophytes in a Wisconsin stream throughout one growing season. Except for a brief period in early spring when a Cladophora glomerata -filamentous algal community was dominant, Potamogeton pectinatus was the dominant macrophyte species in Badfish Creek.
2. Maximum community biomass was 710 g DW m⁻², with a maximum above ground biomass of 620 g DW m⁻² and a maximum below ground biomass of 120 g DW m⁻². Annual productivity was estimated at 1435 g DW m⁻² year⁻¹, with a calculated P/B of 2.01.
3. In situ net production averaged 2.83g C g AFDW⁻¹ h⁻¹ Net positive carbon gain by the P. pectinatus community occurred when water temperatures were above 15°C, and daylength at least 12h. This is correlated to the onset of tuber germination in spring, and the point of maximal biomass decline in autumn.     

Population dynamics and feeding of mayfly larvae in some acid and alkaline New Zealand streams

SUMMARY. 1. Population dynamics (density, biomass, annual production), gut contents and feeding rates of mayflies ( Deleatidium spp.; Leptophlebiidae) were compared in two naturally acid (mean pH≃4.8). brownwater streams and two alkaline (mean pH 7.5), clearwater streams in South Westland, New Zealand.
2. Mean densities of larvae (range 234–2318 m⁻²) were higher in alkaline streams on most of the six bimonthly sampling dates. Mean biomass (range 0.020–0.376 g larval dry weight (LDW) m⁻²) was always highest at the stable, spring-fed, alkaline site and was lower at the acid sites and another alkaline site where the population was always dominated by small larvae.
3. Annual production was high at the more stable, alkaline site (10.35 gLDW m⁻²) but much lower at the other sites (2.49–3.77 g m⁻²).
4. Gut contents of larvae were dominated by fine (45–75 μm widest diameter) paniculate matter (69–99%), diatoms (up to 21%) and. at one site, filamentous algae (8–13%).
5. Grazing rates of mayflies on epilithon were significantly higher on stones taken from acid than alkaline streams and material grazed from the former contained a higher proportion of inorganic material (87–93% and 61–83% inorganics, respectively).
6. Higher grazing rates may reflect lower quality of epilithic food in acid, brownwater streams, a factor that could contribute to the lower productivity of Deleatidium populations at these sites.     

Production and litter processing by crayfish in an Appalachian mountain stream

SUMMARY 1. Mean annual density and biomass () of Cambarus bartonii in an Appalachian mountain stream (U.S.A.) was 12 individuals m⁻² and 1669 mg (ash-free dry weight) m⁻².
2. Annual production ( P ) of C bartonii was 961 mg AFDW m⁻². Despite high biomass, low growth rates resulted in low production and a low P/ ratio of 0.58.
3. While C bartonii constituted 61% of the total macroinvertebrate biomass, it contributed only 13% of annual community secondary production.
4. Litter processing was positively related to temperature and crayfish size. Cambariis bartotnii was estimated to comminute 36 g m⁻² y⁻¹ of leaf litter (>1 mm²) to 24 g m−^>2 y⁻¹ fine particulate material (<1 mm²). The annual pattern of litter comminution by crayfish was regulated by temperature. As a result, >5()% of shredding activity by crayfish occurred from June to September which was also the period of lowest litter standing crops and activity of other shredding macroinvertebrates.
5. We speculate that during summer crayfish play an important role in temperate woodland streams by converting slowly processed leaf litter species (e.g. Rhododendron ) to fine particles which are then available to collector-gatherers (e.g. Chironomidae, Oligochaeta).     

Temporal patterns of grazer-periphyton interactions in laboratory streams

SUMMARY. 1. The snail Juga silicuta (500 m^?2) and the caddisfly Dicosmoecus gilvipes (50 m^?2) were introduced into separate laboratory streams on days 1, 9, 16 and 28 of algal development. The mayfly Baetis spp. (500 m^?2) was introduced on days 1 and 16, and two streams did not receive grazers. We assessed the interaction between succession in the pcriphyton, herbivore type and time of encounter in a 40-day experiment. 2. In ungrazed streams, the chlorophyte Scenedesmus obliquus was the most abundant early colonizer. The relative abundance of diatoms increased after day 9, and at day 40 the algal assemblage consisted of a thick mat of diatoms and S. obliquus with an overstorey of filaments of the chlorophyte Stigeoclonium tenue. In general, introductions of grazers at any stage altered this pattern by removing biomass, accelerating the replacement of S. obliquus by diatoms, and suppressing the growth of filaments. Grazing also reduced the relative abundance of the larger diatom Nitzschia oregona but increased the relative abundance of the smaller adnate diatoms Nitzschia frustulum var. perpusilla and Navicula minima. 3. Dicosmoecus decreased algal biomass and altered successional trajectories to a greater degree than either Juga or Baetis. Dicosmoecus rapidly grazed the entire substrate, whereas Juga and Baetis only cleared patches in the assemblages. Little alteration in algal development was observed in the Baetis streams after day 16, probably because (he periphyton assemblages attained a size and structure that prevented effective grazing by Baetis. 4. The patchy grazing patterns of Juga and Baetis resulted in more diverse algal assemblages than either the Dicosmoecus grazed or ungrazed streams. In natural streams, the temporal and spatial pattern of grazing relative to the developmental stage of the periphyton may contribute to maintaining a mosaic of algal patches in different serai stages.     

The Impact of Sheep Grazing on Net Nitrogen Mineralization Rate in Two Temperate Salt Marshes

Abstract: Nitrogen mineralization rate was studied in grazing trials with three different stocking rates (0, 3, 10 sheep ha^-1) in two man-made salt marshes, viz. a Puccinellia maritima -dominated low salt marsh and a high salt marsh dominated by Festuca rubra. Mineralization rates were derived from the amounts of mineral N which accumulated in situ during six-week incubation periods in tubes containing undisturbed soil cores from the upper 10 cm soil layer. The annual rates of net N mineralization were significantly higher in the better drained, high salt marsh (71 - 81 kg ha^-1 yr^-1) than in the low salt marsh (39 - 49 kg ha^-1 yr^-1). High amounts of belowground litter accumulated in the low salt marsh due to frequent water logging. Both N mineralization and nitrification rate were negatively correlated with soil water content. In the Puccinellia maritima salt marsh, grazing had neither an effect on N mineralization rates during any of the incubation periods nor on annual mineralization rates. In the Festuca rubra salt marsh, N mineralization rates increased earlier during spring at the intensively grazed site than at the moderately grazed and the ungrazed site. N mineralization and nitrification rates were significantly higher at the ungrazed site than at the intensively grazed site during the period of peak net N mineralization from the end of April until mid-June. Although sheep grazing affected the seasonal pattern of N mineralization in the high marsh, grazing did not affect the annual rate of net N mineralization.     

Periphyton as alternative food source for the filter-feeding cladoceran Daphnia magna

1.  Daphnia magna , a well-studied primary consumer, is mainly known as a filter feeder. In this study, we investigated the ability of D. magna to use periphyton as an alternative food source to phytoplankton. We examined the development of laboratory populations fed with different food sources ( Desmodesmus subspicatus and/or periphyton or neither) over a period of 42 days, and observed the behaviour of the daphnids.
2.  The addition of periphyton to phytoplankton food led to an increase of daphnid population biomass. When fed with periphyton as the only food source, a small but stable D. magna population developed.
3.  The behaviour of daphnids fed with both food sources revealed a preference for feeding on D. subspicatus . Only below a concentration of D. subspicatus of approximately 0.05 mg C L⁻¹ (0.4 × 10⁷ cells L⁻¹) did D. magna use periphyton as an alternative food source.
4.  Periphyton showed distinct reactions to grazing by D. magna . The thickness of the periphyton layer was reduced from about 4 to 1 mm and we observed a change in species composition due to grazing.
5.  The ability of D. magna to graze on periphyton could serve to stabilize its population density and reinforce its competitive advantage over other cladocerans. By switching between food sources, D. magna can act as a coupler between pelagic and benthic habitats and food webs.     

Grazing by oligochaetes and snails on epiphytes

SUMMARY. 1. The isotope ³³P was used to assess the effect of grazitig by oligochaetes (mainly Stylaria lacttstris L.) and the snail, Lymnaea peregra (Muller), on epiphytes within an Equisetum fluviatile L. stand.
2. Two 1 m² polystyrene enclosures were set up within the emergent macrophyte zone of the lake. At the start of the experiment ³³P-solution was mixed with the water in both tanks. Algal and animal samples for ³³P- analysis were collected during the peak occurrence of epiphytes in June.
3. Phosphorus release rates from the animals through defaecation and excretion were measured in the laboratory. The grazing rate of oligochaetes was 2.2–4.1 mg P m⁻² (of bottom) d⁻¹ of which about two - thirds was released and recycled. The oligochaete density ranged from 13,400 to 20,900 m⁻². The snails (25 ind. m⁻²) grazed 1.2-1.5 mg P m⁻²d⁻¹of whieh about a quarter was released through defaecation and excretion.
4. Daily consumption by the oligochaetes and snails corresponded to 22–45% of the average phosphorus uptake by the epiphytes.     

Invertebrate grazing during the regenerative phase affects the ultimate structure of macrophyte communities

1. Although the biomass of freshwater macrophytes consumed by invertebrate herbivores (excluding crayfish) is usually low, there is growing evidence that invertebrates do exert a structuring effect on macrophyte communities. To explain this, we postulated that the effect of invertebrates may be concentrated on macrophytes during their regenerative phase.
2. We tested this hypothesis by means of a mesocosm-based experiment, in which we investigated the effects of different densities of pond snails [ Lymnaea stagnalis (L.)] on macrophytes regenerating from the natural sediment propagule bank.
3. After 2 months, a diverse macrophyte community had established in the absence of snails, mainly from sexual propagules. Under moderate snail grazing (4 individuals m⁻²), the ultimate biomass of macrophytes was similar, but its species composition differed dramatically. Only a few unpalatable taxa, such as Ceratophyllum demersum and Nymphaeaceae, persisted. Moreover, the relative success of macrophytes regenerating from vegetative rather than sexual propagules improved. Under higher snail grazing (20 m⁻²), all macrophytes disappeared before the end of the experiment.
4. These results confirm that snails at natural densities can have a strong effect on the ultimate structure of macrophyte communities by selectively consuming some species at a juvenile stage. Therefore, the regenerative phase can be seen as a window of opportunity for invertebrate grazers, which can have a qualitative effect on communities that is disproportionate to the biomass consumed.     

The influence of drought on the fish and invertebrate populations of an upland stream in Wales

SUMMARY. 1. The exceptionally dry summer of 1976 presented an opportunity to assess the impact of drought on the fauna of an unregulated stream in Wales.
2. Total number of macroinvertebrates varied from 1808 m⁻² in September 1976 to 7412m⁻² in August 1977. Two main effects of drought were observed: an initial reduction in abundance during the drought and a change in community structure in the following year, probably through impaired reproduction.
3. Trout density ranged from 0.075±0.028 to 0.402±0.093 m⁻² and salmon parr density varied between 0.014±0.004 and 0.414±0.119 m⁻². The only detrimental effect of the drought on the fish fauna was the elimination of the 1976 year class of young salmon. probably due to prolonged high water temperature.     

Stream periphyton response to grazing and changes in phosphorus concentration

Grazing by the large caddisfly larva, Dicosmoecus gilvipes (Trichoptera; Limnephilidae), drastically reduced periphyton biomass in laboratory channels at a current velocity of 20 cm s^–1. Reduction in biomass as chl a and AFDW ranged from 88 to 93% and 82 to 85%, respectively. On average, grazing rate increased with in-channel SRP (soluble reactive phosphorus) content from 6 to 10 µg 1^–1. Grazing rates averaged 25.9–29.3 µg chl a m^–2 d^–1 and 10.8–12.2 µg chl a mg^–1 d^–1 based on area and grazer biomass, respectively, with most variability among treatments being due to the grazing effect. Grazing tended to shift the algal community increasingly to filamentous blue-green algae regardless of enrichment. After three weeks, Phormidium comprised over 61% of the community in grazed treatments but only 35% in ungrazed treatments. The stalked diatom Gomphonema comprised only 4% of the grazed community, but 11% in the three ungrazed channels with similar values for Scenedesmus. A model that includes grazing was calibrated to the data and produced a reasonable expectation of periphyton biomass over a range in SRP concentrations. While the model with constant grazer abundance predicts a gradually increasing grazed biomass as SRP increases, grazer production in natural streams may actually increase to accommodate the increased food production.     

Periphyton production in Fort River, Massachusetts

SUMMARY. The primary production and general ecology of a periphyton community of a New England, lowland stream were studied over a seventeen-month period. Temperature, light, periphyton chlorophyll-α, and community structure were monitored regularly. Seasonally distinct chlorophyll peaks coincided with the light maximum in early May, just prior to the appearance of leaves of riparian trees, and again in autumn after terrestrial leaf fall. During midwinter, despite low light and temperature levels and high stream discharge, mean chlorophyll concentrations remained similar to summer values.
A mathematical expression relating periphyton photosynthesis per unit chlorophyll-α to temperature, light and periphyton density was established with submersible light-dark chambers in situ . Survey data collected over the study period were employed in the empirical equation to estimate seasonal variations in periphyton primary production. Weekly mean daily estimates of periphyton gross production ranged from < 0.1 g O₂ m⁻², during midwinter, to 6.5 g O₂ m⁻² during early May. Estimated annual periphyton gross production and respiration were 0.58 and 1.27 kg O₂ m⁻², respectively. Factors influencing seasonal variations of Fort River periphyton standing crop are discussed.     

Vertical distribution and abundance of invertebrates within the sandy substrate of a low-gradient headwater stream

SUMMARY. 1 The vertical distribution of invertebrates (>0.053 mm) was studied in a sandy-bottomed, first-order stream on the Coastal Plain of Virginia, U.S.A. Invertebrate species composition, abundance and bio-mass were determined monthly over one year at sediment depth intervals of 0–1, 1–5, 5–15,15–30 and 30–40 cm.
2. The subsurface community was numerically dominated by species of Chironomidae, Nematoda and Crustacea, while much of the biomass was due to early instars of several species of Trichoptera.
3. Invertebrate density and biomass decreased significantly with depth in the substrate (ANOVA; P <0.05). Annual mean density decreased from 1,346,844 individuals m⁻³ at the surface to 13,578 individuals m⁻³ at 15–30 cm. Annual mean biomass decreased from 66.30 g m⁻³ at the surface to 0.44 g m⁻³ at 15–30 cm.
4. Dissolved oxygen decreased markedly from the surface to the 5 cm depth in the substrate, anaerobic conditions often occurring below 10 cm. Density and biomass both showed a significant positive relationship with dissolved oxygen concentration (Linear regression; P <0.05).
5. Physical forces were important in structuring the subsurface invertebrate community. Besides low dissolved oxygen concentration, sediment scouring resulting from storm discharge dramatically reduced density and biomass     

Modelling black fly production dynamics in blackwater streams

SUMMARY. 1. Two predictive models were employed along with intensive field sampling to estimate production of black flies ( Simulium spp.) on snags (submerged wood) in three blackwater streams on the Georgia Coastal Plain of the southeastern U.S.A. One model predicts daily growth rate from temperature and hydrograph pattern; the other predicts habitat abundance (of snags) from river height.
2. In the sixth order Ogeechee River, annual production was twice as high in 1982 (7.1 g dry mass [=DM] m⁻² of snag surface) as in 1983 (3.6 g DM m⁻²). When converted to production per m² of river bottom, values were 35–40% of the snag surface estimates. Annual production was much lower in fourth order Black Creek (1982, 1.3 g DM m⁻² of snag surface) and much higher in the sixth order Satilla River (1975, 15.6–40.0 g DM m⁻²).
3. There was a distinct bimodal pattern of black fly production in the Ogeechee River in both years, with peaks occurring in winter and summer. Similar bimodal patterns of production were found in Black Creek and in the Satilla River. Although there appears to be an intrinsic component to the bimodal pattern, production peaks (growth rate and biomass) appear to be associated with initial stages of flooding.
4. Annual production/biomass ratios (37–85) are the highest reported for black fly populations. The variation of annual P/B ratios among sites was more strongly dependent on the temporal distribution of standing stock biomass than on differences in growth rates. Variation in production among sites appears to be due to differences in current velocity, hydro-graph variability, and abundance of coexisting consumers.     

Facilitation of periphyton production by tadpole grazing: functional differences between species

1. This study examined how interactions between resources that vary in edibility, and herbivores that vary in ability to acquire resources, control primary productivity. In a northern California river, grazing on Cladophora glomerata , a relatively inedible filamentous green alga, and its more nutritious epiphytic diatoms, was manipulated by exposing cobbles to tadpoles ( Rana boylii or Hyla regilla ) or excluding tadpoles.
2. Rana indirectly facilitated Cladophora by removing diatoms, whereas Hyla did not significantly change biomass relative to controls. Algal ash-free dry mass on cobbles in Rana treatments was 65 and 72% greater than on controls in two years of investigation (1991 and 1993). Rana decreased epiphytic diatom biovolume by 56% and detritus by 87%.
3. Because nitrogen excretion rates of Hyla and Rana were similar, the differences in effect between the two species were probably due to their roles as consumers rather than as recyclers.
4. The net effect of Rana on periphyton was a 10% increase in areal specific primary productivity (mg O₂ h^–1 m^–2); Hyla caused an 18% decrease. Rana decreased biomass-specific productivity (mg O₂ h^–1 g^–1) 44%; Hyla had no effect.
5. In tadpole exclosures, grazers such as baetid mayfly larvae (mostly Centroptilum sp.) were 4.7 (1991) and 1.8 (1993) times more abundant, and midge larvae (Chironomidae) were 2.5 (1991) and 2 (1993) times more abundant than in Rana enclosures. Invertebrate assemblages in Hyla enclosures, however, were similar to exclosures. Few predatory insects and fish colonized Rana enclosures. Path analyses indicated that Rana affected macroinvertebrates via both interference and exploitation of epiphytic diatoms.     

Facilitation of periphyton production by tadpole grazing: functional differences between species

1. This study examined how interactions between resources that vary in edibility, and herbivores that vary in ability to acquire resources, control primary productivity. In a northern California river, grazing on Cladophora glomerata , a relatively inedible filamentous green alga, and its more nutritious epiphytic diatoms, was manipulated by exposing cobbles to tadpoles ( Rana boylii or Hyla regilla ) or excluding tadpoles.
2. Rana indirectly facilitated Cladophora by removing diatoms, whereas Hyla did not significantly change biomass relative to controls. Algal ash-free dry mass on cobbles in Rana treatments was 65 and 72% greater than on controls in two years of investigation (1991 and 1993). Rana decreased epiphytic diatom biovolume by 56% and detritus by 87%.
3. Because nitrogen excretion rates of Hyla and Rana were similar, the differences in effect between the two species were probably due to their roles as consumers rather than as recyclers.
4. The net effect of Rana on periphyton was a 10% increase in areal specific primary productivity (mg O₂ h^–1 m^–2); Hyla caused an 18% decrease. Rana decreased biomass-specific productivity (mg O₂ h^–1 g^–1) 44%; Hyla had no effect.
5. In tadpole exclosures, grazers such as baetid mayfly larvae (mostly Centroptilum sp.) were 4.7 (1991) and 1.8 (1993) times more abundant, and midge larvae (Chironomidae) were 2.5 (1991) and 2 (1993) times more abundant than in Rana enclosures. Invertebrate assemblages in Hyla enclosures, however, were similar to exclosures. Few predatory insects and fish colonized Rana enclosures. Path analyses indicated that Rana affected macroinvertebrates via both interference and exploitation of epiphytic diatoms.     

The invertebrate fauna of four macrophytes in a lotic system

SUMMARY. 1. Invertebrates associated with four species of aquatic plants and on the adjacent substrate were investigated in a riffle/run section of a hard-water stream.
2. Invertebrates were more numerous on stones than on any species of plant (> 17,000m⁻² versus 1000–4000 m⁻²). Gatherers were more abundant on Ranunculus longirostris , where proportionally more fine detritus was trapped in the leaf bundles than on other plants. Scrapers were most numerous on Charu vulgaris , where attached diatoms were also most prevalent.
3. Invertebrate community structure on three plant speeies was quite different from that on stones. This was not the case for Potamogeton amplifolius which appeared to be used by the organisms as a mere extension of the substrate.     

The effect of nutrient addition on the phytoplankton community of an oligotrophic lake

This investigation was performed during a 5-yr period (1974–1978) in the oligotrophic Lake Langvatn, Central Norway. In 1975 and 1976 the lake was enriched with a commercial fertilizer, In 1975 increase in phytoplankton biomass was first recorded more than three weeks after the fertilization started, despite a near fivefold increase in the primary production after fertilizer application. The mean seasonal biomass increased from c. 3500 mg wet weight m⁻² in 1974 to 4400 mg in 1975. In 1976 the biomass increased to near 9600 mg ⁻² and the seasonal primary production to 49.0 g C m⁻² (22.2 g C in 1975), despite a reduction in the nutrients added. Chrysophytes constituted the largest share of the seasonal algal biomass in all years, but in fertilization periods cryptophytes dominated in 1975 and diatoms, chlorophytes and cryptophytes in 1976. The highest biomass turnover rate was recorded during a period of cryptophyte dominance. The different biomass and production development in the fertilization years may be explained by a change in the consumer level.     

Population dynamics and production of Jesogammarus annandalei, an endemic amphipod, in Lake Biwa, Japan

SUMMARY 1. Population dynamics and production of Jesogammarus annandalei , an endemic amphipod in Lake Biwa, were examined from April 1997 to June 1998. The life cycle of this species was 1 year with the new generation beginning in early autumn. They preferred low temperature (<12 °C) and their spatial distribution varied seasonally and accordingly.
2. In deep water, the abundance of J. annandalei ranged from 200 to 63 000 m⁻² and decreased towards summer and the biomass (0.01∼3.6 g C m⁻²) was on average comparable that of zooplankton. The density was much higher than that recorded by a study conducted 35 years ago.
3. Individual growth rate of this amphipod was high in winter and spring but decreased in summer. Annual production of J. annandalei (6.2 g C m⁻² year⁻¹) was only 2% of primary production but was at the higher end of the range reported for amphipods in oligo- and mesotrophic lakes.
4. These results are consistent with the view that Lake Biwa is becoming more eutrophic, with a consequent decrease in the abundance of predatory fish in the profundal zone.     

Biomass and production estimates of a fish community in a small South African estuary

The fish community of the small (17·5 ha) intermittently open East Kleinemonde estuary was sampled between 1994 and 1997 to estimate population size, standing stock, growth and productivity. The estuarine-spawning species were numerically more abundant ( n c . 750 000) but due to their small size contributed only 11·7% to the total biomass. The total annual productivity of all fishes in the estuary ( n c . 890 000), with a standing stock of 28·44 g m ⁻², was calculated at 55·89 g m⁻² year⁻¹. The small sparid Rhabdosargus holubi with a production estimate of 41·35 g m⁻² year⁻¹ accounted for <74% of the total fish production in this estuary.