Indirect effects of predatory trout on organic matter processing in detritus-based stream food webs

Indirect effects of predators on basal resources in allochthonous-based food webs are poorly understood. We investigated indirect effects of predatory brown trout ( Salmo trutta ) on detritus dynamics in southern beech ( Nothofagus spp.) forest streams in New Zealand through predation on the obligate detritivore, Zelandopsyche ingens (Trichoptera, Oeconesidae). Trout presence/absence and Z. ingens density were manipulated in flow-through tanks to investigate the lethal and sub-lethal effects of trout on litter processing by Z. ingens . An experiment that allowed trout access to Z. ingens showed trout predation reduced densities of Z. ingens resulting in slower breakdown of coarse particulate organic matter (CPOM) and reduced production of fine particulate organic matter (FPOM). A second experiment that prevented trout access to Z. ingens , but allowed the transmission of trout cues, resulted in no change in litter processing rates in the presence of trout. Litter processing rates were higher in high Z. ingens density treatments compared to low density treatments. Thus, trout effects on litter processing were due to reduced Z. ingens densities, not trout-induced modifications to Z. ingens feeding behaviour. Field assays of litter processing rates using artificial leaf packs in natural streams showed significant reductions in CPOM loss in trout streams compared to fishless streams. Z. ingens dominated biomass in fishless stream leaf packs, but a facultative shredder, Olinga feredayi , dominated trout stream leaf packs. Thus, the absence of Z. ingens drove differences in processing rates between trout and fishless streams and the indirect effects of trout on litter processing observed in mesocosms were evident in complex, natural food webs. Overall our study provides evidence that predators can influence resource dynamics in donor-controlled food webs through their effects on consumers.     

Density reductions by predatory trout increase adult size and fecundity of surviving caddisfly larvae in a detritus-based stream food web

1. In some situations, individuals surviving in environments where predation is intense can grow faster because the benefits of release from intraspecific competition outweigh costs associated with anti-predator responses. Whether these 'thinning' effects of predation occur in detritus-based food webs where resource renewal occurs independently of consumption by consumers was studied. We investigated how effects of predatory brown trout ( Salmo trutta ) on the larvae of the detritivorous stream caddisfly, Zelandopsyche ingens , influenced the size and fecundity of the caddisfly adults.
2. Trout substantially reduced the abundance of Z. ingens larvae, but adult male and female Z. ingens were significantly larger in trout streams compared to fishless streams. Females in trout streams had 33% more eggs than fishless stream females, and egg sizes were not significantly different. In mesocosms, Z. ingens larvae in low density treatments reflecting trout stream abundances grew significantly faster than larvae in high density treatments that were characteristic of fishless stream abundances. Non-lethal trout presence did not influence case building behaviour, feeding rates or growth or Z. ingens larvae, indicating non-lethal effects of predators were negligible.
3. Increased adult size and fecundity associated with trout stream individuals were probably a result of predator thinning of larval density indirectly releasing surviving Z. ingens from intraspecific competition. Thus, predator thinning did influence interactions between larvae in this detritus-based food web as larval growth was strongly density-dependent. However, extrapolating the total number of eggs potentially produced indicates the increased fecundity of females in trout streams would not compensate for losses of larvae to trout predation.     

Cascading effects of predatory fish on the composition of benthic algae in high‐altitude streams

Cascading effects of predators can affect ecosystem properties by changing plant biomass, distribution and assemblage composition. Using data from field surveys and whole‐stream experiments we tested the hypothesis that predatory trout change assemblage composition of benthic algae in high‐elevation streams mediated by grazer behavior. Field surveys revealed that the taxonomic composition of algal assemblages differed significantly between streams that contained trout and those that were fishless; but comparisons of palatable versus unpalatable algal taxa between fish and fishless streams were equivocal because of high natural variability. Therefore, we tested for a behavioral (non‐consumptive) trophic cascade experimentally by adding brook trout chemical cues to six naturally fishless streams for 25 days and compared responses of grazers and algae to six reference streams without fish cues added. Algal response variables included rates of change in the abundance of three physiognomic categories, from most palatable (attached erect and prostrate diatoms) to least palatable (non‐diatoms), as determined from food selectivity analyses of the most common grazers (mayflies and caddisflies). Fish cues did not affect the mean densities or changes in densities of total grazers or any individual grazer species. However, in streams where fish cues were added, rates of accrual of attached erect diatoms, which was the preferred algal type for the grazer most vulnerable to trout predation (Baetis), were higher and their densities increased significantly faster with increasing densities of this grazer species than in reference streams. Results of his experiment support the hypothesis that predator induced suppression of grazer foraging behavior, rather than cascading effects of top predators on grazer density, may contribute to variation in the composition of algal assemblages among streams by allowing proliferation of most palatable algal species.     

Bioturbation by a dominant detritivore in a headwater stream: litter excavation and effects on community structure

Bioturbation can affect community structure by influencing resource distribution and habitat heterogeneity. Bioturbation by detritivores in small headwater streams could affect community structure by reintroducing buried detrital resources into the food web and could also affect the distribution of various taxa on detritus. We evaluated the ability of the caddisfly Pycnopsyche gentilis to uncover experimentally buried leaves in a headwater stream. Packs of leaves were placed in enclosures and covered with a known volume of sediment. We added 0, 3 or 6 large Pycnopsyche to the enclosures which were permeable to most other invertebrate taxa. Leaf packs were sampled after 23 days and leaf pack mass, the amount of sediment covering the leaf packs, and macro‐ and microinvertebrate densities on leaf packs were quantified. There was a significant negative relationship between Pycnopsyche density and leaf pack mass. Pycnopsyche also reduced the volume of sediment covering leaf packs. Pycnopsyche had complex effects on the abundance of invertebrate taxa associated with the leaves. Some taxa exhibited their highest abundance in the 3 Pycnopsyche treatment while others exhibited non‐significant increases as Pycnopsyche density increased. These results suggest that the beneficial effects of Pycnopsyche (e.g. uncovering leaves which increases the availability of habitat and food) outweigh any negative effects (e.g. disturbance, encounter competition) of the caddisfly when it is present at lower densities. However, the negative impacts of Pycnopsyche appear to outweigh the positive effects via sediment removal at higher caddisfly densities for some taxa. Our results suggest that bioturbating organisms in streams have the potential to reintroduce organic matter to detrital food webs and affect the distribution and abundance of benthic taxa associated with organic matter.     

The influence of introduced trout on the benthic communities of paired headwater streams in the Sierra Nevada of California

1. Non‐native trout have been stocked in streams and lakes worldwide largely without knowledge of the consequences for native ecosystems. Although trout have been introduced widely throughout the Sierra Nevada of California, U.S.A., fishless streams and their communities of native invertebrates persist in some high elevation areas, providing an opportunity to study the effects of trout introductions on natural fishless stream communities. 2. We compared algal biomass and cover, organic matter levels and invertebrate assemblages in 21 natural fishless headwater streams with 21 paired nearby streams containing stocked trout in Yosemite National Park. 3. Although environmental conditions and particulate organic matter levels did not differ between the fishless and trout streams, algal biomass (as chlorophyll a concentration) and macroalgal cover were, on average, approximately two times and five times higher, respectively, in streams containing trout. 4. There were no differences in the overall densities of invertebrates in fishless versus paired trout streams; however, invertebrate richness (after rarefaction), evenness, and Simpson and Shannon diversities were 10–20% higher in fishless than in trout streams. 5. The densities of invertebrates belonging to the scraper‐algivore and predator functional feeding guilds were higher, and those for the collector‐gatherer guild lower, in fishless than trout streams, but there was considerable variation in the effects of trout on specific taxa within functional feeding groups. 6. We found that the densities of 10 of 50 common native invertebrate taxa (found in more than half of the stream pairs) were reduced in trout compared to fishless streams. A similar number of rarer taxa also were absent or less abundant in the presence of trout. Many of the taxa that declined with trout were conspicuous forms (by size and behaviour) whose native habitats are primarily high elevation montane streams above the original range of trout. 7. Only a few taxa increased in the presence of trout, possibly benefiting from reductions in their competitors and predators by trout predation. 8. These field studies provide catchment‐scale evidence showing the selective influence of introduced trout on stream invertebrate and algal communities. Removal of trout from targeted headwater streams may promote the recovery of native taxa, community structure and trophic organisation.     

Cascading effects of predatory fish exclusion on the detritus-based food web of a lake littoral zone (Lake Vico, central Italy)

An exclosure experiment was carried out in the reed-dominated littoral zone of a volcanic lake (Lake Vico, central Italy) to test whether the impact of predatory fish on benthic invertebrates cascades on fungal colonisation and breakdown of leaf detritus. The abundance, biomass, and Shannon diversity index of the invertebrate assemblage colonising Phragmites australis leaf packs placed inside: (1) full-exclosure cages, (2) cages allowing access only to small-sized fish predators, and (3) cageless controls, were monitored over a 45-day period together with the mass loss and associated fungal biomass of leaf packs. The species composition of the fungal assemblage was further assessed at the end of the manipulation. In general, invertebrate predators did not show any significant response to fish exclusion, either on a trophic guild or on a single taxon level. In contrast, the exclusion of large predatory fish induced a diverse spectrum of changes in the abundance and population size-structure of dominant detritivore taxa, ultimately increasing the biomass and Shannon diversity index of the whole detritivorous guild. These changes corresponded with significant variations in leaf detritus decay rates as well as in the biomass and assemblage structure of associated fungal colonisers. Our experimental findings provide evidence that in Lake Vico effects of fish predators on invertebrate detritivores influence the fungal conditioning and breakdown of the detrital substrate. We conclude that in lacustrine littoral zones predator-driven constraints may structure lower trophic levels of detritus-based food webs and affect the decomposition of leaf detritus originated from the riparian vegetation.     

Stream inflow and predation risk affect littoral habitat selection by benthic fish

1. We examined small, fishless headwater streams to determine whether transport of macroinvertebrates into the littoral zone of an oligotrophic lake augmented food availability for Cottus asper, an abundant predatory fish in our study system. We sampled fish and macroinvertebrates during the recruitment and growth season of 2 years, either monthly (2004) or bi‐monthly (2005), to observe whether stream inputs increased prey availability and whether variation in total macroinvertebrate biomass was tracked by fish. 2. Observations from eight headwater streams indicated that streams did not increase the total macroinvertebrate biomass in the shallow littoral zone at stream inflows, relative to adjacent plots without stream inputs (controls). The taxonomic composition of stream macroinvertebrates drifting toward the lake differed from that in the littoral lake benthos itself, although there was no evidence of any species change in the composition of the littoral benthos brought about by stream inputs. 3. Although streams made no measurable contribution to the biomass or taxonomic composition of the littoral macroinvertebrate benthos, there was substantial temporal variation in biomass among the eight sites for each of the (n = 7) sample periods during which observations were made. Variation in total biomass was primarily a function of bottom slope and benthic substrata in the lake habitats. Dominant taxonomic groups were Baetidae, Ephemerellidae (two genera), Leptophlebiidae, Chironomidae (three subfamilies) and Perlodidae, although we did not determine the specific substratum affinities of each taxon. 4. Mixed effects linear models identified a significant interaction between macroinvertebrate biomass and plot type (stream inflow vs. control) associated with fish abundance. Across the observed range of macroinvertebrate biomass, fish showed a significant preference for stream inflows, but more closely tracked food availability in the controls. For young‐of‐the‐year (YOY), a negative effect of temperature was also included in the model, and we observed lower temperatures at stream inflows. However, abundance of predatory adults affected habitat selection for YOY. Lake‐bottom slope also accounted for variation in abundance in both fish models. 5. Our results suggest that the effect of fishless headwater streams on downstream fish may not always be through direct delivery of food. In this study system, fish preferred stream inflow plots, but this preference interacted with macroinvertebrate biomass in a manner that was difficult to explain. For YOY, predation risk was related to the preference for stream inflows, although the specific factor that mitigates predation risk remains poorly understood.     

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

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Crayfish in lakes and streams: individual and population responses to predation, productivity and substratum availability

1. In a correlative study, we investigated the relative importance of fish predation, refuge availability and resource supply in determining the abundance and size distributions of the introduced and omnivorous signal crayfish (Pacifastacus leniusculus) in lakes and streams. Moreover, the biomass and food selection of predatory fish was estimated in each habitat type and stable isotopes of carbon and nitrogen were measured in perch (Perca fluviatilis), the dominant predator in the lakes, and in its potential food sources (crayfish, juvenile roach and isopods). 2. In lakes, crayfish were the most frequent prey in large perch (46%), followed by other macroinvertebrates (26%, including the isopod Asellus aquaticus) and small fish (25%). Crayfish and fish dominated the gut contents of large perch with respect to biomass. Nitrogen signatures showed that perch were one trophic level above crayfish (approx. 3.4‰) and a two‐source mixing model using nitrogen isotope values indicated that crayfish (81%) contributed significantly more to perch isotope values than did juvenile roach (19%). A positive correlation was found between the abundance of crayfish and the biomass of large perch. Crayfish abundance in lakes was also positively correlated with the proportion of cobbles in the littoral zone. Lake productivity (chlorophyll a) was positively correlated with crayfish size, but not with crayfish abundance. 3. In streams, brown trout (Salmo trutta) were the most abundant predatory fish. Gut contents of large trout in a forested stream showed that terrestrial insects were the most frequently found prey (60%), followed by small crayfish (27%) and isopods (27%). In contrast to lakes, the relative abundance of crayfish was negatively correlated with the total biomass of predatory fish and with total biomass of trout. However, abundance of crayfish at sites with a low biomass of predatory fish varied considerably and was related to substratum grain size, with fewer crayfish being caught when the substratum was sandy or dominated by large boulders. The mean size of crayfish was greater at stream sites with a high standing stock of periphyton, but neither predator biomass nor substratum grain size was correlated with crayfish size. 4. Our results suggest that bottom‐up processes influence crayfish size in lakes and streams independent of predator biomass and substratum availability. However, bottom‐up processes do not influence crayfish abundance. Instead, substratum availability (lakes) and interactions between predation and substratum grain size (streams) need to be considered in order to predict crayfish abundance.     

Spatial and Temporal Pattern of Caddisfly Distribution at a Mesohabitat Scale in two Patagonian Mountain Streams Subjected to Pastoral Use

Substrate, flow type, nutrients, aquatic vegetation, organic matter, and caddisfly community structure were studied at two low order streams (Glyn and Nant y Fall) subjected to pastoral use in the Patagonian mountains. At both sites, we examined the effect of habitat type (boulder‐pebble with and without filamentous algae, cobble‐pebble, gravel‐sand, leaf‐pack, the submerged macrophytes Myriophyllum quitense and Isoetes savatieri) and season (high and low water period) on caddisfly assemblages. Benthic particulate organic matter (BPOM) ranged between 4.6 and 472 g m^–2, all allochtonous detrital fractions were significantly higher at leaf‐packs at Glyn, whereas M. quitense habitats supported more BPOM and macrophytes biomass at Nant y Fall. As expected, boulder‐pebble sustained higher Trichoptera richness than M. quitense and gravel‐sand, moreover all habitats showed higher density than M. quitense at Nant y Fall. According to our results at least nine caddisfly species exhibited some habitat preference with boulder‐pebble and cobble‐pebble the most selected habitat. These particular habitats sustained more than 68% of the total caddisfly species. Multidimensional scaling ordination highlighted differences in composition per habitat for both sites showing a clear distinction among depositional and erosional habitats. Substrate, flow type, detritus biomass were important predictors defining assemblages. Based upon our findings, those anthropogenic actions or stressors that change hydraulic as well substrate attributes in mountain streams such as stock trampling, dredging, clearing of riparian areas, will reduce caddisfly richness. These results are relevant for outline management and conservation biomonitoring and schemes in headwater Patagonian streams as well as other similar environments worldwide (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

The influence of predatory fish on mayfly drift: extrapolating from experiments to nature

1. A knowledge of how individual behaviour affects populations in nature is needed to understand many ecologically important processes, such as the dispersal of larval insects in streams. The influence of chemical cues from drift‐feeding fish on the drift dispersal of mayflies has been documented in small experimental channels (i.e. < 3 m), but their influence on dispersal in natural systems (e.g. 30 m stream reaches) is unclear. 2. Using surveys in 10 Rocky Mountain streams in Western Colorado we examined whether the effects of predatory brook trout (Salvelinus fontinalis) on mayfly drift, that were apparent in stream‐side channels, could also be detected in natural streams. 3. In channel experiments, the drift of Baetis bicaudatus (Baetidae) was more responsive to variation in the concentration of chemical cues from brook trout than that of another mayfly, Epeorus deceptivus (Heptageniidae). The rate of brook trout predation on drifting mayflies of both species in a 2‐m long observation tank was higher during the day (60–75%) but still measurable at night (5–10%). Epeorus individuals released into the water column were more vulnerable to trout predation by both day and night than were Baetis larvae treated similarly. 4. Drift of all mayfly taxa in five fishless streams was aperiodic, whereas their drift was nocturnal in five trout streams. The propensity of mayflies to drift was decreased during the day and increased during the night in trout streams compared with fishless streams. In contrast to the channel experiments, fish biomass and density did not alter the nocturnal nature nor magnitude of mayfly drift in natural streams. 5. In combination, these results indicate that mayflies respond to subtle differences in concentration of fish cues in experimental channels. However, temporal and spatial variation in fish cues available to mayflies in natural streams may have obscured our ability to detect responses at larger scales.     

Seasonal dynamics of macroinvertebrate assemblages in the benthos and associated with detritus packs in two low-order streams with different riparian vegetation

• 1 The seasonal dynamics of the benthic macroinvertebrate assemblage, and the subset of this assemblage colonising naturally formed detritus accumulations, was investigated in two streams in south‐west Ireland, one draining a conifer plantation (Streamhill West) and the other with deciduous riparian vegetation (Glenfinish). The streams differed in the quantity, quality and diversity of allochthonous detritus and in hydrochemistry, the conifer stream being more acid at high discharge. We expected the macroinvertebrate assemblage colonising detritus to differ in the two streams, due to differences in the diversity and quantity of detrital inputs.
• 2 Benthic density and taxon richness did not differ between the two streams, but the density of shredders was greater in the conifer stream, where there was a greater mass of benthic detritus. There was a significant positive correlation between shredder density and detritus biomass in both streams over the study period.
• 3 Detritus packs in the deciduous stream were colonised by a greater number of macroinvertebrates and taxa than in the conifer stream, but packs in both streams had a similar abundance of shredders. The relative abundance of taxa colonising detritus packs was almost always significantly different to that found in the source pool of the benthos.
• 4 Correspondence analysis illustrated that there were distinct faunal differences between the two streams overall and seasonally within each stream. Differences between the streams were related to species tolerances to acid episodes in the conifer stream. Canonical correspondence analysis demonstrated a distinct seasonal pattern in the detrital composition of the packs and a corresponding seasonal pattern in the structure of the detritus pack macroinvertebrate assemblage.
• 5 Within‐stream seasonal variation both in benthic and detritus pack assemblages and in detrital inputs was of similar magnitude to the between‐stream variation. The conifer stream received less and poorer quality detritus than the deciduous stream, yet it retained more detritus and had more shredders in the benthos. This apparent contradiction may be explained by the influence of hydrochemistry (during spate events) on the shredder assemblage, by differences in riparian vegetation between the two streams, and possibly by the ability of some taxa to exhibit more generalist feeding habits and thus supplement their diets in the absence of high quality detritus.

     

Trophic‐level dependent effects on CO2 emissions from experimental stream ecosystems

Concern over accelerating rates of species invasions and losses have initiated investigations into how local and global changes to predator abundance mediate trophic cascades that influence CO₂ fluxes of aquatic ecosystems. However, to date, no studies have investigated how species additions or losses at other consumer trophic levels influence the CO₂ flux of aquatic ecosystems. In this study, we added a large predatory stonefly, detritivorous stonefly, or grazer tadpole to experimental stream food webs and over a 70‐day period quantified their effects on community composition, leaf litter decomposition, chlorophyll‐a concentrations, and stream CO₂ emissions. In general, streams where the large grazer or large detritivore were added showed no change in total invertebrate biomass, leaf litter loss, chlorophyll‐a concentrations, or stream CO₂ emissions compared with controls; although we did observe a spike in CO₂ emissions in the large grazer treatment following a substantial reduction in chlorophyll‐a concentrations on day 28. However, the large grazer and large detritivore altered the community composition of streams by reducing the densities of other grazer and detritivore taxa, respectively, compared with controls. Conversely, the addition of the large predator created trophic cascades that reduced total invertebrate biomass and increased primary producer biomass. The cascading effects of the predator additions on the food web ultimately led to decreased CO₂ emissions from stream channels by up to 95%. Our results suggest that stream ecosystem processes were more influenced by changes in large predator abundance than large grazer or detritivore abundance, because of a lack of functionally similar large predators. Our study demonstrates that the presence/absence of species with unique functional roles may have consequences for the exchange of CO₂ between the ecosystem and the atmosphere.     

Fish determine macroinvertebrate food webs and assemblage structure in Greenland subarctic streams

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Predator effects on prey population dynamics in open systems

Animal population dynamics in open systems are affected not only by agents of mortality and the influence of species interactions on behavior and life histories, but also by dispersal and recruitment. We used an extensive data set to compare natural loss rates of two mayfly species that co-occur in high-elevation streams varying in predation risk, and experience different abiotic conditions during larval development. Our goals were to generate hypotheses relating predation to variation in prey population dynamics and to evaluate alternative mechanisms to explain such variation. While neither loss rates nor abundance of the species that develops during snowmelt (Baetis bicaudatus) varied systematically with fish, loss rates of the species that develops during baseflow (Baetis B) were higher in streams containing brook trout than streams without fish; and surprisingly, larvae of this species were most abundant in trout streams. This counter-intuitive pattern could not be explained by a trophic cascade, because densities of intermediate predators (stoneflies) did not differ between fish and fishless streams and predation by trout on stoneflies was negligible. A statistical model estimated that higher recruitment and accelerated development enables Baetis B to maintain larger populations in trout streams despite higher mortality from predation. Experimental estimates suggested that predation by trout potentially accounts for natural losses of Baetis B, but not Baetis bicaudatus. Predation by stoneflies on Baetis is negligible in fish streams, but could make an important contribution to observed losses of both species in fishless streams. Non-predatory sources of loss were higher for B. bicaudatus in trout streams, and for Baetis B in fishless streams. We conclude that predation alone cannot explain variation in population dynamics of either species; and the relative importance of predation is species- and environment-specific compared to non-predatory losses, such as other agents of mortality and non-consumptive effects of predators. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Detrital Fruit Processing in a Hawaiian Stream Ecosystem1

In temperate forested streams, fruit from riparian trees is generally a minor and seasonal component of the allo‐chthonous detritus. In contrast, riparian fruit input to tropical streams is often high and continuous. Detrital fruit is abundant in some forested Hawaiian streams compared to other forms of riparian detritus, and rates of leaf litter processing by macroscopic invertebrates are very low. These observations suggested that fruit is an important food resource for detritivores. A microcosm system was used to measure the rates at which two common detrital fruits, guava and mango, were processed by two common detritivores, the prawn Macrobrachium lar and the gastropod Tarebia granifera. Comparisons of fruit weight loss rates normalized by detritivore weight indicated that M, lar processed guavas at significantly higher rates than T. granifera, differences in rates of mango processing by M. lar and T. granifera were not significant. Microcosms containing both M. lar and T. granifera were used to test for interactions between the invertebrates that affected rates of mango processing. No interspecific interactions were detected. A field study was conducted in Kaiwiki Stream, Island of Hawaii, to determine rates of detrital fruit input and export. Detrital fruit was supplied to the study area year‐round, with peaks corresponding to summer and autumn fruiting seasons. Guavas and mangos accounted for 85 percent of the fruit biomass entering the stream and 92 percent of the fruit exported from the stream. Mean daily export rates of guava were 7 percent of input, and export rates of mango were 5 percent of input. These measurements suggested that most of the fruit entering the stream is retained and comprises a substantial food resource for detritivores. Comparisons of the biomass‐specific rates at which M. lar and T. granifera processed mangos and guavas with the rates at which mangos and guavas entered Kaiwiki Stream suggested that these invertebrates can process most of the detrital fruit in the stream.     

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.     

Assessing the functional importance of large‐bodied invertebrates in experimental headwater streams

Recent theoretical advances in food web ecology emphasize the importance of body size disparities among species for the structure, stability and functions of ecosystems. Experimental confirmations of the functional importance of large species, independent of their trophic position, are scarce. We specifically examine the multiple ecological roles of large invertebrates from two distinct trophic levels in headwater streams. We experimentally manipulated the presence of large predatory invertebrates (two Perlid stoneflies) or detritivores (a limnephilid caddisfly and a Pteronarcys stonefly) in a two‐by‐two design in stream channels open to immigration/emigration of smaller biota. We assessed treatment effects on the trophic structure of the benthic invertebrate community, dynamics of basal resources (benthic algae and leaf litter of cedar and alder), and stability of litter decomposition rates against an experimental pulse perturbation (fine sediment input). The presence of the large invertebrates was associated with a ten‐fold decrease in the biomass of invertebrate filterers whereas other trophic groups were unaffected by the large species. The biomass of benthic algae was lower and the rate of mass loss of alder litter was higher in channels lacking the large predators, thus revealing trophic cascades operating along both algal‐based and detritus‐based food chains. The large predators had no detectable effect on the decomposition of cedar whereas both cedar and alder disappeared faster in the presence of the large detritivores. Furthermore, the large predators and large detritivores interactively influenced the decomposition of the cedar–alder mixture through a litter diversity effect and the variability of the rate of alder decomposition after a pulse of fine sediment. Because the large invertebrates affected multiple ecosystem properties, and as their absence was not rapidly compensated for by small immigrant species, our findings support the notion that large species could be critically important in controlling ecosystem structure and functioning.     

The influence of moss on grazers in high‐altitude streams: food,refuge or both?

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