The influence of periphyton, detritus and shelter on invertebrate colonization of aquatic bryophytes

1. Artificial bryophytes were placed in a shaded and an unshaded New Zealand alpine stream to investigate why invertebrates colonized these structures and, by inference, real plants. Three experiments were conducted to investigate the influence of (i) periphyton and detritus (ii) shelter, and (iii) time, on invertebrate colonization. 2. In the first experiment, seven taxa at the unshaded site displayed a preference for substrata with high detrital and periphyton biomass, presumably reflecting a food relationship. At the shaded, less stable site, only two taxa displayed such a relationship. 3. Reducing substratum ‘stem’ density (i.e. ‘shelter’) in the second experiment had little effect on the biomass of periphyton at each site, and only at the shaded site was detrital biomass reduced on low-density substrata. Abundances of most of the twenty-two invertebrate taxa analysed were unaffected by stem density reduction: densities of only four taxa at the unshaded site, and two at the shaded site were reduced. 4. Stepwise multiple regression showed that invertebrate abundance was little affected by stem density at either site. Indeed, shelter was the primary factor influencing abundance of only two of twenty-two taxa at the unshaded site, and none at the shaded site. Abundances of most taxa were related to periphyton or detrital biomass at each site. 5. The third experiment investigated temporal relationships between invertebrate density, periphyton and detrital biomass, and exposure time of artificial bryophytes. Regression analyses indicated that of twenty-two taxa at the stable, unshaded site, eight were influenced by periphyton biomass, three by detrital biomass, and two by exposure time. At the unstable shaded site, abundances of only eight of twenty-two taxa were significantly related to the measured variables, of which exposure time was most important (four taxa).     

The influence of riparian vegetation on macroinvertebrate community structure and functional organization in six new Guinea streams

Information on the ecology of New Guinea streams is meagre, and data are needed on the trophic basis of aquatic production in rivers such as the Sepik in Papua New Guinea which have low fish yields. This study investigates the relationship between riparian shading (from savanna grassland to primary rainforest), algal and detrital food, and macroinvertebrate abundance and community structure in 6 Sepik River tributary streams. A particular aim was to elucidate macroinvertebrate community responses to changes in riparian conditions. All streams supported diverse benthic communities, but morphospecies richness (overall total 64) was less than in streams on the tropical Asian mainland; population densities of benthic invertebrates, by contrast, were similar to those recorded elsewhere. Low diversity could reflect limited taxonomic penetration, but may result from the absence of major groups (Plecoptera, Heptageniidae, Ephemerellidae, Psephenidae, Megaloptera, etc.) which occur on the Asian mainland. Population densities of all 19 of the most abundant macroinvertebrate taxa varied significantly among the 6 study streams, but community composition in each was broadly similar with dominance by Baetidae and (in order of decreasing importance), Leptophlebiidae, Orthocladiinae, Elmidae and Hydropsychidae. Principal components analysis (PCA) undertaken on counts of abundant macroinvertebrate taxa clearly separated samples taken in two streams from the rest. Both streams contained high detrital standing stocks and one was completely shaded by rainforest. Stepwise multiple-regression analysis indicated that population densities of the majority of abundant taxa (11 out of 19) across streams (10 samples per stream; n = 60) were influenced by algae and/or detritus, although standing stocks of these variables were not clearly related to riparian conditions. When regression analysis was repeated on mean counts of taxa per stream (dependent variables) versus features of each stream as a whole (thus n = 6), % shading and detritus were the independent variables yielding significant regression models most frequently, but pH, total-nitrogen loads and algae were also significant predictors of faunal abundance. Further regression analysis, undertaken separately on samples (n = 10) from each stream, confirmed the ability of algae and detritus to account for significant portions of the variance in macroinvertebrate abundance, but the significance of these variables varied among streams with the consequence that responses of individual taxa to algae or detritus was site-specific.Community functional organization — revealed by investigation of macroinvertebrate functional feeding groups (FFGs) — was rather conservative, and streams were codominated by collector-gatherers (mean across 6 streams = 43%) and grazers (36%), followed by filter-feeders (15%) and predators (7%). The shredder FFG was species-poor and comprised only 0.4% of total macroinvertebrate populations; shredders did not exceed 2% of benthic populations in any stream. PCA of FFG abundance data was characterized by poor separation among streams, although there was some evidence of clustering of samples from unshaded sites. The first 2 PCA axes accounted for 84% of the variation in the data suggesting that the poor separation resulted from the general similarity of FFG representation among streams. Although stepwise multiple-regression analysis indicated that algae and detritus accounted for significant proportions of the variations in population density and relative abundance of some FFGs, the response of community functional organization to changes in riparian conditions and algal and detrital food base was weak — unlike the deterministic responses that may be typical of north-temperate streams.     

A manipulative study of macroinvertebrate grazers in Hong Kong streams: do snails compete with insects?

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Spatial and seasonal variations in benthic algal assemblages in streams in monsoonal Hong Kong

Samples from stone surfaces were collected in pools within four unpolluted hillstreams (two shaded and two unshaded) in monsoonal Hong Kong (lat. 23°N) to elucidate the extent of spatial (within and among streams) and temporal (seasonal) variations in algal biomass and assemblage composition. Sampling continued for over 12 months, incorporating the dry season when streams were at baseflow, and the wet season when spates were frequent. We anticipated that algal biomass would be lower in shaded streams and during the wet season, with associated seasonal differences in assemblage composition or relative abundance of different growth forms (e.g. erect versus prostrate). Benthic chlorophyll a (a proxy for algal biomass) varied among streams from an annual mean of 11.0–22.3 mg m⁻². Dry-season standing stocks were 18% higher than during the wet season when spate-induced disturbance reduced algal standing stocks. Algal biomass varied significantly at the stream scale, but not at the pool scale, and was lower in unshaded streams, where standing stocks may have been limited by high densities of algivorous balitorid loaches (mainly Pseudogastromyzon myersi). An overriding effect of grazers on algal biomass could also have reduced variations resulting from spate-induced disturbance. Significant differences in assemblage composition among streams, which were dominated by diatoms and cyanobacteria (totally 82 taxa) were not systematically related to shading conditions. Seasonal variations in algal assemblages were statistically significant but rather minor, and did not involve major shifts in composition or growth form caused by spate-induced disturbance. The abundance of filamentous cyanobacteria in all the streams may have been due to ‘gardening’ by balitorid loaches that removed erect or stalked diatoms and favoured cyanobacteria that persist through basal regeneration of filaments. This explanation requires validation through manipulative experiments. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Luis Mauricio Bini     

Bryophytes as invertebrate habitat in two New Zealand alpine streams

SUMMARY. 1. Quantitative samples of lotic benthic invertebrates and associated organic material were taken monthly for 18 months from bryophyte and gravel habitats in two New Zealand alpine streams in Arthur's Pass National Park, South Island; an unshaded site above the tree-line and a shaded stream flowing through mountain beech forest. This was to assess the ecological role of bryophytes in these habitats and to determine the relative effects on the fauna of algal and detrital resources. 2. Bryophytes were confined to stable bedrock substrata in shallow (mean depth of 4.5 and 10.5 cm), fast water (mean velocity of 0.21 ms^?1 and 0.30ms^?1) at the unshaded and shaded sites. Mean ash-free dry weight (AFDW) of organic material from bryophyte areas was similar at the unshaded and shaded sites (335.6 and 272.4 gm^?2). Low stream-bed stability of the gravel areas resulted in poor retention of organic matter, with average AFDW of 59.1 and 77.9 gm^?2 at the unshaded and shaded sites, respectively. Periphyton biomass, however, was much higher at the unshaded site. 3. Within bryophytes, water velocity is reduced and large quantities of periphyton and detritus accumulate. This explains the greatly enhanced invertebrate densities within bryophytes (218400 and 53500 individuals m^?2), compared with gravel areas (20900 and 7400 individuals m^?2) in the unshaded and shaded sites, respectively. The higher invertebrate densities at the unshaded site reflect greater periphyton biomass there. 4. Invertebrate densities followed clear seasonal patterns, attributable to changes in chironomid densities. These reflected larval chironomid hatching following adult oviposition. This illustrates the importance of these plants as oviposition sites, and as protective nurseries for small larvae where they can obtain shelter during floods and consume the abundant food sources available to them. 5. DECORANA ordination revealed taxonomically distinct invertebrate communities in each stream, and in bryophytes and gravels. Several taxa appeared restricted to either the unshaded site (e.g. Acroperla spiniger and Hydrobiosis silvicola) or the shaded site (e.g. Zelolessica     

Effects of overhead canopy on macroinvertebrate production in a Utah stream

SUMMARY. 1. Macroinvertebrate abundance and production were compared between an open and shaded site of a stream in the Wasatch Mountains, Utah. Mean biomass was significantly higher at the open site for midges (Chironomidae), 4.6×; Baetis bicaudatus , 5.7×; Baetis tricaudatus , 2.3×; Drunella coloradensis , 12× and Cinygmula sp., 1.6×, Abundance of most other macroinvertebrates (except black flies: Simuliidae) was also greater at the open site, but differences were not significant. Black fly biomass was 1.7× greater at the shaded site.
2. Seasonal production, estimated by the size-frequency and instantaneous growth rate methods, was greater at the open site than the shaded site for most taxa (except black flies) and reflected differences in standing crops between the sites rather than differences in rate of growth. Excluding black flies, production at the open site was twice as high as at the shaded site.
3. The greater abundance and production of most invertebrate taxa at the open site is probably associated with either higher quality food (algae and algal detritus), or a phototactic attraction to sunlit areas.
4. Sampling of large cobbles was an efficient method of sampling all taxa except Cinygmula sp. which was more abundant on smaller substrate particles.     

The influence of riparian vegetation on macroinvertebrate community structure in four Hong Kong streams

The benthic macroinvertebrate communities of four Hong Kong streams with similar water chemistry but differing riparian conditions were investigated. Differences in chlorophylls and detritus standing stocks reflected the degree of shading by riparian vegetation; Tai Po Kau Forest Stream (TPKFS) and Bride's Pool (BP) had an allochthonous food base while autochthonous energy sources were relatively more important in Lam Tsuen River (LTR) and Pui O Stream (POS). The macrobenthos was dominated by aquatic insects and morphospecies richness (excluding Chironomidac) ranged from 70 (POS) to 94 (TPKFS). Differences in total macroinvertebrate abundance across sites were not related to chlorophylls or detritus.
Highly significant differences in the morphospecies comprising the most numerous taxa at each site were observed, and TPKS and BP were more similar to each other than to the LTR-POS pair with respect to top-ranked taxa. Intraspecific comparisons of the abundance of these taxa revealed significant differences for 23 out of 30 morphospecies. Multiple regression of the abundance of key morphospecies against independent variables (chlorophylls a , b and c , total chlorophyll and detritus) yielded significant best-fit models for all taxa showing intersite differences in abundance. These data can be interpreted as reflecting the influence of riparian vegetation on stream community structure via an effect on food supply. The results were discussed with reference to the applicability of the River Continuum Concept in the tropics.     

Response of an algal assemblage to nutrient enrichment and shading in a Hawaiian stream

To investigate the effects of nitrate enrichment, phosphate enrichment, and light availability on benthic algae, nutrient-diffusing clay flowerpots were colonized with algae at two sites in a Hawaiian stream during spring and autumn 2002 using a randomized factorial design. The algal assemblage that developed under the experimental conditions was investigated by determining biomass (ash-free dry mass and chlorophyll a concentrations) and composition of the diatom assemblage. In situ pulse amplitude-modulated fluorometry was also used to model photosynthetic rate of the algal assemblage. Algal biomass and maximum photosynthetic rate were significantly higher at the unshaded site than at the shaded site. These parameters were higher at the unshaded site with either nitrate, or to a lesser degree, nitrate plus phosphate enrichment. Analysis of similarity of diatom assemblages showed significant differences between shaded and unshaded sites, as well as between spring and autumn experiments, but not between nutrient treatments. However, several individual species of diatoms responded significantly to nitrate enrichment. These results demonstrate that light availability (shaded vs. unshaded) is the primary limiting factor to algal growth in this stream, with nitrogen as a secondary limiting factor.     

Macroinvertebrate Responses to Algal and Bacterial Manipulations in Streams

Our study was designed to assess the relative importance of algae and bacteria as sources of energy for stream macroinvertebrates. In one experiment, we manipulated algae by artificially shading six sections in each of two streams, one stream with an open canopy (clear-cut drainage basin) and the other with a closed canopy (forested drainage basin); both streams were in Hubbard Brook Experimental Forest, New Hampshire, USA. Chlorophyll a concentrations were reduced from 0.2 to 0.05 μg/cm² in artificially shaded sections of both streams. However, macroinvertebrates showed no response to these algal manipulations in either the clear-cut or forested stream. Nutrient concentrations (N and P) were low and limiting to primary production in both the clear-cut and forested streams. Additionally, both streams had relatively low macroinvertebrate densities suggesting bottom-up controls were important in macroinvertebrate abundance. However, the forested stream did have higher macroinvertebrate densities presumably because of higher inputs of coarse particulate organic matter from the riparian vegetation. In a second experiment, in Augusta Creek, Michigan, we manipulated both algae and bacteria. To reduce algae, we artificially shaded experimental stream channels so that chlorophyll a was reduced from natural levels of 3.0–5.6 to 0.4–0.7 μg/cm². Half of the shaded channels had dissolved organic carbon (DOC – sucrose) dripped into them to raise DOC levels by 2–3 mg/l and thus stimulate bacterial abundance. Open channels, with higher algal abundance, had higher densities of Ephemerella, but only in November when nymphs were larger. Channels with increased DOC had higher bacterial abundances, higher densities of Chironomidae and lower densities of Heptageniidae. Several other macroinvertebrate taxa that were at relatively low abundance in our samples showed no significant response to these manipulations. Our results suggest that early instar Ephemerella may not rely as heavily on algae as later instars. Also, certain taxa were able to use the heterotrophic microbial community, especially chironomids which increased in numbers when bacterial density increased; thus, the bacterial carbon source may be more important to some stream macroinvertebrates than previous studies have suggested.     

Regulation of algal biomass in a small lowland stream: field experiments on the role of invertebrate grazing, phosphorus and irradiance

1. Field experiments were undertaken in a small Danish lowland stream to study the role of invertebrate grazing, phosphorus concentration and irradiance in the regulation of benthic algal biomass on stones. 2. Algal biomass was regulated by invertebrate grazing. The gastropod Ancylus fluviatilis prevented algal biomass build up in early spring at a density of about 900 ind. m^–2, and reduced algal biomass to very low levels during the algal growth period at a density of about 6000 ind. m^–2. Grazing pressure therefore might regulate the magnitude of peak algal biomass in Gelbæk stream, a finding in agreement with earlier field observations. 3. As phosphorus enrichment to around 152 μg l^–1 during the period when irradiance was not limiting did not result in any further increase in algal biomass, it can be concluded that algal growth in spring was not controlled by phosphorus limitation. 4. Algal biomass development differed significantly in a shaded and in a non-shaded reach of the stream. Only very low algal biomass accumulation was evident in the shaded reach, while in the non-shaded reach an algal biomass peak of about 800 mg chlorophyll m^–2 was evident after 6 weeks of colonization. The shaded reach represented the light condition in Gelbæk stream after leaf proliferation of the overhead canopy and high bank vegetation. The findings therefore suggest that irradiance available to the algal community after shading from riparian vegetation prevents further algal biomass increase and hence determines the timing of peak algal biomass in the stream. 5. The irradiance experiment also suggests that if not regulated by invertebrate grazing or shading by an overhead canopy in the summer, then the magnitude of peak biomass in Gelbæk stream will be regulated by self-shading in the algal community. 6. These field experiments support theories, derived from laboratory experiments and field studies, that regulation of algal biomass is a complex interaction of top-down and bottom-up mechanisms.     

Benthic invertebrate communities in six Danish forest streams: Impact of forest type on structure and function

I investigated the impact of riparian vegetation type on stream invertebrate communities in SIX Danish forest streams during the period 1992–1993 Two of the streams ran through beech forest Fagus sylvatica , two through mixed, mainly deciduous forest, and two through conifer plantations Algal biomass and detritus standing stock differed significantly depending on forest type In mixed deciduous forest streams algal biomass was higher and detritus standing stock lower than in beech and conifer forest streams Benthic invertebrate community composition was functionally similar in all streams, with detritivores comprising >80% by numbers of the community Grazer abundance was low in all streams Shredder abundance in the four deciduous streams was significantly correlated to coarse detritus standing stock When taking into account both quantitative and qualitative POM parameters for all six streams combined, shredder abundance correlated significantly to the amount of CPOM There were significantly more invertebrates with a ≥ 2 yr life cycle in the conifer and beech forest streams than m the mixed forest streams, probably reflecting the larger and more stable food resource For all streams combined there was a significant correlation between predator abundance and the abundance of potential prey My findings suggest that Danish forest streams are regulated by "bottom-up control" at all trophic levels within the invertebrate community, and hence that forest type can structure benthic communities in forest streams     

Experimental nutrient enrichment of forest streams increases energy flow to predators along greener food‐web pathways

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Response of grazing impacts of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) to seasonal disturbance in Hong Kong streams

1. Manipulative experiments were carried out in four Hong Kong streams (two shaded, two unshaded) to investigate the impact of grazing by an algivorous fish, Pseudogastromyzon myersi, on benthic algal biomass and assemblage composition. Experiments were conducted and repeated during both the dry and wet seasons to determine whether spate‐induced disturbance modified any grazing effect. Treatments comprised fish exclusion and inclusion via closed and open cages, with a no‐cage treatment used as a control for the cage effect. Treatments were maintained for 4 weeks in each experimental run. 2. Grazing by P. myersi reduced benthic algal biomass and the organic matter content of periphyton in open cages and the no‐cage treatment relative to closed cages. The similarity between open‐cage and no‐cage treatments was evidence that the overall difference among treatments was caused by limiting fish access to closed cages and not merely an artifact of caging. Grazing effects were broadly similar in all streams, but there was a significant statistical interaction between treatments and seasons. 3. Analysis of dry‐season data matched the overall trend in inter‐treatment differences, confirming the effects of grazing by P. myersi on algal biomass and periphyton organic matter. Significant differences in algal assemblage composition between closed‐cage and no‐cage treatments during the dry season reflected reductions in the abundance of erect, stalked diatoms (Gomphonema) and filamentous cyanobacteria (Homeothrix). Removal of these vulnerable overstorey algae by P. myersi resulted in greater abundance of understorey diatoms (Achnanthes and Cocconeis) in the no‐cage treatment in all streams during the dry season. The composition of algal assemblages in open cages was intermediate between the other two treatments. 4. Although fish densities were greater in all streams during the wet season, spate‐induced disturbance obscured grazing effects and there were no significant differences among treatments attributable to fish grazing. Seasonal variation in impacts of P. myersi grazing provides support for the harsh‐benign hypothesis, and confirms that biotic factors are less important controls of stream algal biomass and assemblage structure during periods (i.e. the wet season in Hong Kong) when abiotic disturbances are frequent or intense.     

The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.     

Production and population dynamics of the prosobranch snail Sulcospira hainanensis (Pachychilidae), a major secondary consumer in Hong Kong streams

Pachychilid snails are common and widespread in tropical Asian streams, and given their abundance, these heavy-shelled prosobranchs may serve as a ‘trophic dead-end’ preventing energy transfer to higher trophic levels. Sulcospira hainanensis (Bröt, 1872) is an abundant secondary consumer in Hong Kong streams, and tends to have higher standing biomass in less shaded sites. We predicted that S. hainanensis would be more productive in unshaded streams, where snails have a higher assimilation of more nutritious algal carbon. The production and population dynamics of S. hainanensis were studied along a gradient of shading in four Hong Kong streams. The increment-summation method was considered to most realistically represent snail production, which ranged from 1,612.8 to 6,123.9 mg ash-free dry mass m^?2 year^?1; these estimates were among the upper range of values reported for stream prosobranchs. Production, turnover, and growth were higher in unshaded streams, where the combined effects of higher light input and nutrient concentrations likely enhanced the supply of algal food. Despite high standing biomass (26% of total benthic fauna), S. hainanensis had a relatively low (~16%) contribution to total production in one site, suggesting that it is not a major trophic dead-end in the food webs of tropical Hong Kong streams.     

Benthic algal biomass in an unshaded first-order lowland stream: distribution and regulation

The vertical distribution of algal biomass in the bed sediment and the seasonal development of benthic algae on stones and fine-grained sediments were studied in a small unshaded stream. In addition, field experiments were conducted on the role of irradiance and phosphorus in regulating algal biomass. We found that algal biomass was high at a sediment depth of ten centimetres. Comparison of studies on algal biomass where different depths of the sediment are used should therefore be made with caution. Substrata-dependent differences in algal biomass development were substantial. While algal biomass development on stones was controlled by macroinvertebrate grazing, that on the fine-grained sediment followed the dynamics of incident irradiance, but was attenuated by sediment rebedding. Because of the high grazing pressure on algal biomass on stony substrata, no significant response to phosphorus enrichment was attained. In contrast, algal biomass development on fine-grained sediments was phosphorus-limited. Heavy shading of the fine-grained sediments did not significantly affect algal biomass development, thus suggesting that phosphorus limitation prevents algae from fully utilizing the light resource in this stream. This revised version was published online in July 2006 with corrections to the Cover Date.     

NUTRIENT AND LIGHT LIMITATION OF ALGAE IN TWO NORTHERN CALIFORNIA STREAMS1

Nutrient-diffusing subsrates were used to investigate nutrient limitation of attached algal assemblages in a shaded stream and an unshaded stream in northern California. Water from both streams contained low levels of nitrogen (< 14 μg.L^?1) and very low N:P ratios (< 2). After 31 days of colonization and growth, attached algal biomass on nitrate-diffusing substrates was significantly greater than on control substrates in the unshaded stream. Nitrate-diffusing substrates also supported larger numbers of grazing insects in the unshaded stream. The prostrate diatoms Achnanthes lanceolata Bréb. and Coconeis placentula Ehr. displayed the most consistent positive responses to nitrate enrichment. Nutrient enrichment did not increase the accrual of algal biomass in the shaded stream, but algal biomass was significantly greater at sites located under openings in the tree canopy, implicating light as a limiting factor in this stream. Several Navicula and Nitzschia species, and one unidentified Gomphonema species, were positively associated with higher light levels in the shaded stream. Shade appears to be the primary factor limiting algal growth in small northern California streams, but when its effect is reduced by logging, the inherently low levels of nitrogen in these streams can become limiting.     

Exotic crayfish in a brown water stream: effects on juvenile trout, invertebrates and algae

SUMMARY 1. The impact of the introduced omnivorous signal crayfish (Pacifastacus leniusculus) on trout fry, macroinvertebrates and algae was evaluated in a brown water stream in southern Sweden using in situ enclosures. We also examined the gut content of all surviving crayfish in the enclosures. Two crayfish densities in addition to a control without crayfish were used in replicate enclosures (1.26 m²) in a 1‐month experiment. Additionally, 20 trout fry (Salmo trutta) were stocked in each enclosure to assess the effects of crayfish on trout survival and growth. 2. Detritus was the most common food item in crayfish guts. Animal fragments were also frequent while algae and macrophytes were scarcer. Crayfish exuviae were found in crayfish guts, but the frequency of cannibalism was low. 3. Trout survival in enclosures was positively related to water velocity but was unaffected by crayfish. 4. Total invertebrate biomass and taxon richness were lower in crayfish treatments. The biomass of all predatory invertebrate taxa was reduced but only three of six non‐predatory taxa were reduced in the crayfish treatments. 5. Epiphytic algal biomass (measured as chlorophyll a, on plastic strips) was not related to crayfish density, whereas the biomass of epilithic algae (measured as chlorophyll a) was enhanced by high water velocity and high crayfish density. The latter was possibly mediated via improved light and nutrient conditions, as active crayfish re‐suspend and/or remove detritus and senescent algal cells during periods of low water velocity. 6. We conclude that the introduced signal crayfish may affect stream communities directly and indirectly. Invaded communities will have reduced macroinvertebrate taxon richness and the signal crayfish will replace vulnerable invertebrate predators such as leeches. In streams that transport large amounts of sediment or organic matter, a high density of crayfish is likely to enhance benthic algal production through physical activity rather than via trophic effects.     

Effects of omnivorous shrimp in a montane tropical stream: sediment removal,disturbance of sessile invertebrates and enhancement of understory algal biomass

Freshwater shrimp dominate the faunal biomass of many headwater tropical streams: however, their role in community organization is unclear. Enclosure/exclosure experiments in a montane Puerto Rican stream examined direct and indirect effects of two dominant taxa of atyid (Atyidae) shrimp, Atya lanipes Holthuis and Xiphocaris elongata Guerin-Meneville. Both shrimp taxa caused significant reductions in sediment cover on rock substrata, reducing sedimentation and enhancing algal biovolume on clay tiles in cages. When tiles incubated in shrimp exclosures for 2 wks were placed outside of cages, atyid shrimp removed 100% of the sediment cover within a 30 min observation period. Atyid shrimp appear to play an important role in stream recovery after high discharge events by rapidly removing sediments and detritus deposited on benthic substrata in pools. We evaluated the mechanism by which A. lanipes influences algae and benthic insects by comparing patterns of algal biomass, taxonomic composition, and insect abundance between shrimp-exclusion and shrimp-presence treatments both with and without manual sediment removal. The shrimp exclusion treatment without manual sediment removal bad significantly lower algal biomass and greater sedimentation than all other treatments. The treatment in which shrimp were excluded but sediment was manually removed, however, accrued almost the same algal biovolume as the shrimp enclosure treatment, supporting the hypothesis that sediment removal enhances the biovolume of understory algal taxa. Algal community composition was similar between stream bottom bedrock exposed to natural densities of shrimp and all experimental treatments for both Atya and Xiphocaris: a diatom community strongly dominated (78–95%) by the adnate taxon, Achnanthes lanceolata Breb ex. Kutz. Atyid shrimp are important in determining the distribution and abundance of benthic insects through both direct and indirect effects. Sessile, retreat-building chironomid larvae (Chironomidae: Diptera) are negatively affected by both A. lanipes and X. elongata, through direct removal by foraging activities and/or indirectly through depression of sediment resources available to larvae for the construction of retreats. In constrast, the mobile grazer, Cloeodes maculipes (Baetidae: Ephemeroptera) was not adversely affected and atyid shrimp have the potential to exert positive indirect effects on this taxon by facilitating its exploitation of algal resources and/or through enhancement of understory algal food resources through sediment removal.     

Food resources of shredders and other benthic macroinvertebrates in relation to shading conditions in tropical Hong Kong streams

1. Gut content analyses (GCA) of benthic macroinvertebrates, supplemented by carbon and nitrogen stable isotope analyses (SIA), were used to determine the relative contribution of leaf litter and autochthonous food sources to consumer biomass in five shaded and five unshaded streams in tropical Hong Kong. 2. Only four obligate shredders and two facultative shredders were identified out of 58 morphospecies dissected. Non‐shredder taxa consumed little (<23% food eaten) coarse particulate organic matter (CPOM) in spite of its abundance in streams, and GCA revealed that fine particulate organic matter was the major food (25–99%) of most primary consumers. 3. Stable isotope analysis results were in general agreement with the findings of GCA, and confirmed that three of the four obligate shredders had a high dependence (55–78% of assimilated carbon) on CPOM. 4. Autochthonous energy sources were important in all streams: non‐shredding primary consumers examined, which accounted for 72% of total macroinvertebrate abundance in shaded streams, derived (on average) 61% of their biomass from autochthonous foods; the equivalent values for unshaded streams were 72% (abundance) and 71% (biomass).