In situ nutrient assays of periphyton growth in a lowland Costa Rican stream

Nutrient limitation of primary production was experimentally assessed using an in situ bioassay technique in the Quebrada Salto, a third-order tropical stream draining the northern foothills of the Cordillera Central in Costa Rica. Bioassays employed artificial substrata enriched with nutrients that slowly diffuse through an agar-sand matrix (Pringle & Bowers, 1984). Multiple comparisons of regression coefficients, describing chlorophyll-a accrual through time for different nutrient treatments, revealed positive micronutrient effect(s). Micronutrient treatment combinations (Fe, B, Mn, Zn, Co, Mo, EDTA), supplemented with and without nitrate and phosphate, exhibited significantly greater chlorophyll-a accrual over all other treatments (P < 0.05), supporting over three times that of the control after 14-d of substratum colonization. Neither of the major nutrients (N or P) produced a significant stimulation, although the N treatment displayed 50% more chlorophyll-a than the control after 14-d. Similarly, Si, EDTA, and Si + N + P treatments did not exhibit chlorophyll-a response curves that were significantly different from the control. During the experiment, mean NH₄-N and (NO₂ + NO₃)-N concentrations in the Salto were 2.0 µM (28.6 µg · l^–1) and 7.2 µM (100.2 µg · l ^–1), respectively. High concentrations of PO₄-P ( = 2.0 µM; 60.9 µg · l^–1) and TP ( = 3.0 µM; 94.0 µg · l^–1) were also found, and consequently low molar N:P ratios = 4.7). Despite the potential for N limitation in the system, both N and P appear to be at growth saturating levels. This may be due to micronutrient limitation and/or light limitation of periphyton growth in densely shaded upstream portions of the stream.     

Alteration of trophic interactions between periphyton and invertebrates in an acidified stream: a stable carbon isotope study

The hypothesis according to which proliferation of periphytic algae under acid conditions results from a release of grazing pressure is tested. Stable carbon isotope analysis is used to investigate the autochthonous/allochthonous balance of invertebrate feeding in streamside artificial channels that were experimentally acidified. We find that the relative contribution of autochthonous food sources (epilithon) to total invertebrate biomass was slightly lower (after 1 mo of acidification) or not altered (after 2 mo) under acidified conditions when compared with a control. Feeding shifts were exhibited by some invertebrate taxa and provided evidence that acidification modifies trophic interactions between attached algae and primary consumers. Cross-treatment calculations showed that reduction of grazing pressure after the first month of acidification was an effect rather than the cause of periphyton proliferation. Our approach using stable carbon isotope analysis and biomass measurements of macroinvertebrates allows the quantification of the trophic base of lotic secondary producer communities under both experimental and natural conditions.     

The effects of shrimps on benthic material in a Brazilian island stream

1. Shrimps are abundant in many tropical coastal streams. Studies in Central America and the Caribbean have demonstrated the importance of shrimps in removing sediments and altering the composition of the benthos. Previous work in our study area showed that ephemeropterans and not shrimps were important in removing benthic material. 2. Here we used an experimental exclusion to test the hypothesis that shrimps exert strong influence on sediment dynamics with direct and indirect effects on the benthic algal and faunal community at a site where they are the predominant element of macrofauna. We used electricity to exclude Macrobrachium olfersi and Potimirim glabra from small quadrats (0.135 m²) for 34 days in a stream located at Ilha Grande, Rio de Janeiro state, Brazil. We analysed benthic sediment dynamics and community colonisation during this period on artificial substrates in electrified and not‐electrified quadrats. 3. Significantly higher sediment accrual had occurred in the electrified treatment after 14 days and persisted for the 34‐day course of the experiment; substrates protected from shrimps contained about four times as much ash‐free dry mass (AFDM) as those of the controls. After 34 days, significantly more pennate diatoms were present in the exclusion treatment, but chlorophyll was not significantly different between treatments. Densities of baetid ephemeropteran nymphs were significantly higher in the presence of shrimps. 4. We attribute all these effects to the atyid P. glabra, the more abundant and active shrimp observed in the control treatment. Our results suggest that atyid shrimps played an important role related to sediment removal on hard substrate by direct ingestion and ‘bioturbation’. They reduced certain components of the periphyton (pennate diatoms) without affecting primary production (chlorophyll a). The activity of these shrimps on periphyton affected also faunal components such as baetid ephemeropterans and seems to determine the composition of the benthic community.     

Energy flow and the trophic basis of macroinvertebrate and amphibian production in a neotropical stream food web

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The ecological effect of acid conditions and precipitation of hydrous metal oxides in a Rocky Mountain stream

Periphyton and benthic invertebrates assemblages were studied at the confluence of two Rocky Mountain streams, Deer Creek and the Snake River near Montezuma, Colorado. Upstream from the confluence the Snake River is acidic and enriched in dissolved trace metals, while Deer Creek is a typical Rocky Mountain stream. In the Snake River, downstream from the confluence, the pH increases and hydrous metal oxides precipitate and cover the streambed. The algal and benthic invertebrate communities in the upstream reaches of the Snake River and in Deer Creek were very different. A liverwort, Scapania undulata var. undulata, was abundant in the Snake River, and although periphyton were very sparse, there were as many benthic invertebrates as in Deer Creek. Downstream from the confleunce, the precipitation of hydrous metal oxides greatly decreased the abundance of periphyton and benthic invertebrates. This study shows that in streams metal precipitates covering the streambed may have a more deleterious effect on stream communities than high metal-ion activities.     

Changes in atrazine toxicity throughout succession of stream periphyton communities

A study was made to describe atrazine toxicity and its changes throughout succession of periphyton communities of an undisturbed Mediterranean stream. Toxicity was assessed by short-term physiological tests (concentration-effect curves of photosynthesis to atrazine) in the laboratory using artificial substrates colonized in one stream site during winter, and two stream sites (one open and the other shaded) during summer. In the winter experiment, when environmental conditions were relatively steady and chlorophyll content was low, toxicity increased according to the increases in cell density and chlorophyll content throughout colonization. EC₅₀ (concentration inhibiting photosynthesis by 50%) was above 0.8 μM atrazine until day 16 and below 0.4 μM atrazine after three weeks. In the summer experiment, under more variable environmental conditions, the differences between the EC₅₀ at the beginning and the end of the colonization experiments were not significant (one factor ANOVA) at the two sites. EC₅₀ was on average 0.89 μM atrazine in the shaded site and 0.29 μM atrazine in the open site. A significant negative correlation between irradiance and EC₅₀ was observed all the experiments were considered together (r = 0.464, n = 20, p<0.05), suggesting that light history may have an important role in the response to atrazine. This investigation reveals that the response of stream periphyton to atrazine is likely to be influenced by colonization time and the corresponding changes in algal density and community composition as well as by environmental conditions (e.g. light regime) throughout succession. This revised version was published online in September 2006 with corrections to the Cover Date.     

The response of stream periphyton to Pacific salmon: using a model to understand the role of environmental context

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Stable isotope analysis indicates microalgae as the predominant food source of fauna in a coastal forest stream,south‐east Brazil

Abstract: Stable isotope studies of food webs in floodplains, large rivers, mangroves, and seagrasses have shown that, although a large proportion of the biomass may come from higher plants, microalgae provide a disproportionate amount of carbon assimilated by metazoan consumers. Evidence is building that this may also be the case for streams, especially those in the tropics. At the level of individual consumer species we also see that the apparent diet may not be reflected in the carbon assimilated. Tropical streams commonly have omnivore‐detritivore species that potentially show this phenomenon. We tested these concepts in four moderately shaded sites in a stream in well‐preserved Atlantic rainforest at Ilha Grande, Rio de Janeiro. We sampled aquatic insects, shrimps and fish as well as potential terrestrial and aquatic primary food sources. Carbon stocks from terrestrial sources predominated over carbon of algal origin (>99% of total). The primary sources of carbon showed distinctly different isotopic signatures: terrestrial sources had δ¹³C values close to ?30‰, microalgae were ?20‰ and macroalgae were ?25‰. All fauna had δ¹³C values consistent with a carbon source derived from microalgae. Baetid mayflies and atyid shrimps exert a strong grazing pressure on periphyton and organic sediments but appear to assimilate predominantly microalgae. The palaemonid shrimp Macrobrachium olfersi also ingests large amounts of detritus of terrestrial origin, but apparently assimilates animal prey with algal δ¹³C signatures. These results support the growing view that tropical stream food chains are primarily algal based.     

Inconsistency in the experimentally derived relationship between epilithon abundance and the micro-distribution of Agapetus monticolus (Trichoptera)

Abstract Studies of the micro-distribution of herbivorous lotic macroinvertebrates have tended to rely on short-term experiments. While these experiments have provided information on the factors that determine the distributions of herbivorous invertebrates, they ignore the complexity of the epilithon as a food source, any interaction between physical and biological factors and the possibility that the factors that determine an invertebrate species’ distribution may vary through time. Five colonization experiments were performed in an Australian upland stream to determine the role of epilithon abundance in the distribution of an abundant invertebrate (Agapetus monticolus: Trichoptera). Artificial cobbles with different types and amounts of epilithon were placed in the Taggerty River at two sites and the response of A. monticolus recorded. The experiments revealed that, although A. monticolus expressed a preference for patches of epilithon with abundant periphyton, the response varied among experiments performed at different times and was always secondary to the flow preferences of A. monticolus. Studies that fail to acknowledge the presence of temporal variation in the response of animals to their environment run the risk that they will over- or underestimate the importance of particular factors because of the timing of their experiment.     

The influence of periphyton biomass and density on grazing in Physella virgata

We studied how differences in periphyton colonization interval and snail density affected grazing rates in Physella virgata, and whether snails controlled periphyton biomass. Both egestion rates and incorporation rates of ¹⁴C labeled periphyton were estimated in laboratory experiments. Periphyton biomass increased with field colonization interval in all experiments, but did not consistently influence estimates of grazing rate. However, increased periphyton abundance in one of the experiments could still explain higher grazer rates in that year, although larger snail body size is a confounding explanation. Increased snail density also resulted in decreased grazing rates, as observed in earlier studies with this snail species, as well as in studies with other snail grazers. Our results suggest grazing rates and resulting impacts may change seasonally with variation in either periphyton biomass, grazer life-history stage or population density.     

Light,nutrients and the fatty acid composition of stream periphyton

1. While the balance of light and nutrients is known to influence the food quality of herbivores by altering algal phosphorus and nitrogen content, the combined effects of light and nutrients on fatty acid synthesis in freshwater periphyton are relatively unknown. In this study, we manipulated light and phosphorus concentration in large, flow‐through experimental streams to examine their effects on both elemental stoichiometry and fatty acid content in periphyton. 2. Two levels of phosphorus (4 and 80 μg L^?1) and three of light (17, 40, 110 μmol photons m^?2 s^?1) were applied in a factorial design in two separate experiments. Diatoms dominated periphyton communities in both experiments, comprising >95% of algal biovolume. Periphyton growth in the streams was simultaneously affected by both resources, even at low rates of supply. 3. Periphyton C/P and C/N ratios increased with light augmentation and decreased with phosphorus enrichment, and consistent with the light : nutrient hypothesis (LNH). Light effects were strongest in streams with low phosphorus concentrations. 4. Periphyton fatty acids reflected the dominance of diatoms : palmitic (16 : 0), palmitoleic (16 : 1ω7) and eicosapentanoic (20 : 5ω3) were the principal saturated (SAFA), monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA), respectively. Linoleic (18 : 2ω6) and linolenic (18 : 3ω3) acids, characteristic of chlorophytes and cyanophytes, were rare, comprising <2% of total fatty acids. 5. Periphyton fatty acid profiles were highly sensitive to light and phosphorus. The proportion of fatty acids comprised by SAFA and MUFA increased with light augmentation and decreased with phosphorus enrichment, whereas PUFA decreased with light and increased with phosphorus. Light effects on fatty acid composition were strongest in phosphorus‐poor streams. PUFA declined with increasing light/phosphorus ratios in the streams, whereas ‘energy’ fatty acids (16 : 0 and 16 : 1) increased. The ratio of SAFA/PUFA was strongly and positively correlated with C/P and C/N ratios. SAFA and MUFA, normalised to dry mass, increased two‐ to threefold with increasing light, while PUFA normalised to dry mass was not significantly affected by light. 6. Similarities in the responses of fatty acids and elemental stoichiometry to light and phosphorus treatments suggested that they were influenced by a common mechanism. Both components of food quality appeared to be sensitive to light‐regulated rates of carbon fixation which, when coupled with insufficient supplies of phosphorus, caused diatom cells to store surplus carbon in SAFA, MUFA and other carbon‐rich compounds that diluted both essential fatty acids and mineral nutrients.     

Complexity in conservation: lessons from the global decline of amphibian populations

As part of an overall "biodiversity crisis" many amphibian populations are in decline throughout the world. Numerous causes have been invoked to explain these declines. These include habitat destruction, climate change, increasing levels of ultraviolet radiation, environmental contamination, disease, and the introduction of non-native species. In this paper, we argue that amphibian population declines are caused by different abiotic and biotic factors acting together in a context-dependent fashion. Moreover, different species and different populations of the same species may react in different ways to the same environmental insult. Thus, the causes of amphibian population declines will vary spatially and temporally. Although some generalizations (e.g. those concerning environmental stress and disease outbreaks) can be made about amphibian population declines, we suggest that these generalizations take into account the context-dependent dynamics of ecological systems.     

The effects of disturbance on a herbivore-epilithon interaction in an upland stream

Two experiments were performed in an upland stream to determine the effects of the frequency of physical disturbance on the relationship between an abundant glossosomatid trichopteran (Agapetus monticolus; Banks) and the epilithon upon which it feeds. Artificial cobbles with an established epilithic community were tumbled either every 1, 2 or 4 weeks. The first experiment failed to detect any significant effects of rock tumbling on the abundance of A. monticolus or the epilithon: a result due to several spates. The first experiment did reveal that disturbances may disrupt the ability of A. monticolus to locate patches of abundant food. The second experiment found that although the abundance of A. monticolus was not affected by the disturbances, periphyton abundance was significantly reduced. Increasing the frequency of disturbance did not magnify this effect. Comparisons of these results with other studies of disturbance in streams indicate that the effects of disturbance on herbivory may be highly variable. A variety of factors, such as the relative resistances of the herbivores and the epilithon, need to be examined before the effects of disturbances on lotic herbivorous interactions can be completely understood.     

Diel changes in the benthos of stones and of drift in a southern Australian upland stream

Diel changes in the density of the fauna of stones and of drift were investigated in the Toorongo River, an upland river of southern Australia. The densities of the ten most common taxa and of the total fauna in the drift and on the stones were negatively correlated, with 16 out of 33 cases being significant (p < 0.05). Five of the ten most common taxa displayed a general trend of reaching day-time peaks in the benthos (11 out of 15 cases) and night-time peaks in the drift (10 out of 15 cases). The total density on the stones reached a significant peak in the day-time while the total drift density peaked at night.     

Context-dependent aggregation in Common Frog Rana temporaria tadpoles: influence of developmental stage, predation risk and social environment

1. This study examines the aggregation behaviour and activity of larvae of the Common Frog Rana temporaria in relation to the early social environment, ontogeny and the presence of chemical cues from a predatory fish to address three main questions: (i) Does previous social interaction influence aggregation behaviour in later developmental stages? (ii) To what extent does aggregation behaviour depend upon the risk level perceived by the individual? (iii) Does aggregation behaviour change through ontogeny?
2. Tadpoles were reared from eggs either in small groups or in complete isolation. In relatively early stages of development, tadpoles showed no preference for the side of the test container containing siblings over the side containing no larvae regardless of the social context experienced (isolation or contact with siblings).
3. The presence of chemical cues from a potential predator did not trigger the aggregation behaviour of tadpoles, but it had a remarkable effect on their activity and spatial distribution. Tadpoles exposed to water preconditioned by a predator spent significantly less time swimming and avoided the central area of the test container more frequently than did controls exposed to unconditioned water.
4. Tadpoles were more active, avoided the central section and associated preferentially with conspecifics (siblings) at later stages of development regardless of the social conditioning they had previously experienced.
5. Individuals reared in groups were twice as active as individuals reared in isolation. This suggests that the early social environment experienced by larvae can influence future behaviour and thereby growth and development rates.
6. The expression of conspecific attraction is probably linked to an ontogenetic shift in larval behaviour. However, reduced activity, rather than aggregation, appears to be the basic antipredator mechanism in larval Common Frog.     

All creatures great and small: patterns in the stream benthos across a wide range of metazoan body size

SUMMARY 1. The whole metazoan community (i.e. including the meiofauna) of an acidic, fishless stream in south-east England was surveyed over 14 months between March 1999 and April 2000. Invertebrate density, biomass and taxonomic richness were assessed on each sampling occasion in relation to physico-chemical variables.
2. The meiofauna were more numerous and diverse than the macrofauna, while their total biomass occasionally equalled that of the macrofauna.
3. The meiofaunal and macrofaunal assemblages appeared to respond to different environmental factors. The meiofauna showed genuine species turnover through the year, while the macrofauna varied less in taxonomic composition though there were substantial variations in density.
4. These data suggest that the meiofauna and macrofauna exist at different temporal and spatial scales and perceive their environment with a different 'grain'.     

NEOSQUIRREL: a data set of ecological knowledge on Neotropical squirrels

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Environmental variability and the ecological effects of spawning Pacific salmon on stream biofilm

1. Variation in resource subsidies can create or reinforce heterogeneity in recipient ecosystems. Related activities of organisms delivering resource subsidies, such as ecosystem engineering by Pacific salmon spawners (Oncorhynchus spp.), also alter heterogeneity. We studied whether heterogeneity in stream environmental conditions and spawner abundances were reflected in the net ecological effects of salmon (i.e. enrichment by resource subsidies and disturbance by ecosystem engineering) on benthic biofilm. 2. We sampled seven Southeast Alaska streams over 3 years, both before and during the salmon run. In each stream and year, stream environmental characteristics and their influence on responses of benthic biofilm [mean and coefficient of variation of chlorophyll a (chl a), ash‐free dry mass (AFDM) and autotrophic index (AFDM:chl a)] to spawners were assessed. 3. Streams and periods before and during the salmon run were distinct based on their environmental characteristics. The responses of most biofilm metrics to spawners were stream‐ and year‐specific, suggesting that the ecological effect of spawners ranged from net enrichment to net disturbance depending on the stream or year studied. The environmental context, especially temperature, large wood, and sediment size, explained >50% of biofilm variability during the run, but <30% over the entire study, suggesting that salmon can alter environmental constraints. 4. Precision of biofilm estimates improved by increasing either the number of streams or the number of years sampled (i.e. spatial or temporal replication). However, combining data from different North Pacific Rim ecoregions inflated the confidence interval as compared with a single ecoregion, indicating the importance of regional environmental contexts for net salmon effects. 5. Our results suggest that biofilm responses to salmon can vary greatly, even within a single ecoregion, and that environmental conditions can modify net salmon effects. Consequently, generalisations about biofilm responses across the native range of salmon may be challenging.