Top‐Down Control of Reed Detritus Processing in a Lake Littoral Zone: Experimental Evidence of a Seasonal Compensation between Fish and Invertebrate Predation

We investigated whether predatory fish exert a top‐down control on reed leaf packs processing in a lake littoral zone through a trophic cascade. Exclosure experiments were repeated in summer and winter, under high and low natural fish abundance, respectively. Fish exclusion effects on detritus processing and fungal conditioning were consistent with trophic cascade predictions only in summer. In winter, however, results indicated that a trophic cascade was induced by predatory invertebrates. In both seasons, variations in detritivores abundance generally supported a cascade scenario, whereas several taxon‐specific departures occurred during the experimental periods. We conclude suggesting that predators may continuously regulate leaf detritus processing in lake littoral zones, through a seasonal shift in the relative contribution of fish and invertebrate predation. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Functional significance of crayfish in stream food webs: roles of omnivory,substrate heterogeneity and sex

Heterogeneity of species interactions in food webs can result from characteristics of substrates as well as attributes of top consumers. We performed a streamside channel experiment to evaluate the impact of crayfish on lower trophic levels in detritus‐based (leaf packs) and algal‐based food webs (hard‐bottoms). After 43 days, both male and female crayfish had dramatically promoted leaf decomposition, with males processing material at a faster rate. However, the difference in leaf processing rates was not related to a greater level of male activity. Despite the sex‐related difference in residual leaf dry mass, densities of invertebrates in leaf packs were similarly low in the presence of crayfish of either sex, due to resource consumption, physical dislodgment (bioturbation) and/or predation. No trophic cascade was evident in the leaf pack assemblage. In the hard‐bottom assemblage, the results confirmed circumstantial field evidence that crayfish reduce predatory Tanypodinae and indirectly increase collector‐gatherer Chironominae following the prediction of a trophic cascade. However, no other taxa were indirectly facilitated, because of strong direct effects of crayfish on algal abundance (through direct consumption and bioturbation). Overall, impacts of crayfish on lower trophic levels were more pronounced in the structurally complex, detritus‐based assemblages than in its hard‐bottom, algal‐based counterpart. This conflicts with the expectation that net predation effects should be weaker where structural complexity is greater but is mainly a consequence of the profound engineering effects of crayfish in reducing colonisable substrate when they shred and disturb detrital material. Effects of crayfish may therefore propagate differently and with varying strength depending on substrate. Moreover, engineering activities and predation by crayfish appear to have been of overwhelming significance with subtle sex differences in leaf processing rates failing to lead to differences in invertebrate densities.     

Body size and substrate association of littoral insects in relation to vegetation structure

Variation in substrate association types and maximum size of aquatic insects were studied in a vegetated littoral zone of three lake basins. The basins differed from each other in trophic status, biomass of benthivorous fish, and abundance of macrophytes. Four types of substrate association – swimmers, crawlers, semisessiles and burrowers, respectively – were assumed to represent decreasing vulnerability to fish predators. Large-sized species were also hypothesised to be more vulnerable to fish predators. The distributions of species traits were examined in relation to vegetation density. Inferring from ``predation hypothesis' opposite selection pressures on the species traits were expected along the vegetation density. Dense macrophyte beds were thought to be dominated by invertebrate predators and open water by fish predators, since the predation efficiency of fish decreases in complex environments. In the case of invertebrate predator domination, large size and higher activity should be favoured traits among the prey species. Distribution patterns of modes of the two studied traits were explored separately for predatory and non-predatory insects. As expected, swimmers and large-sized crawlers were characteristic of the insect assemblages of dense macrophyte beds. The densities of Odonata, Corixidae, Dytiscidae, Ephemeroptera and Sialidae were higher among macrophytes than in open water, where these insect taxa were possibly depleted by fish. On the other hand, the small-sized and fairly immobile Chironomidae were the most abundant group in open water. These results support the existence of a predator transition zone among littoral vegetation, ranging from domination of invertebrate predation among the dense beds to that of fish predation in open water.     

Small stonefly predators affect microbenthic and meiobenthic communities in stream leaf packs

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Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

Stable isotopes indicate that zebra mussels (Dreissena polymorpha) increase dependence of lake food webs on littoral energy sources

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Contrasting cascades: insectivorous birds increase pine but not parasitic mistletoe growth

1. Intraguild predation occurs when top predators feed upon both intermediate predators and herbivores. Intraguild predators may thus have little net impact on herbivore abundance. Variation among communities in the strength of trophic cascades (the indirect effects of predators on plants) may be due to differing frequencies of intraguild predation. Less is known about the influence of variation within communities in predator-predator interactions upon trophic cascade strength. 2. We compared the effects of a single predator community between two sympatric plants and two herbivore guilds. We excluded insectivorous birds with cages from ponderosa pine Pinus ponderosa trees parasitized by dwarf mistletoe Arceuthobium vaginatum. For 3 years we monitored caged and control trees for predatory arthropods that moved between the two plants, foliage-feeding caterpillars and sap-feeding hemipterans that were host-specific, and plant damage and growth. 3. Excluding birds increased the abundance of ant-tended aphids on pine and resulted in an 11% reduction in pine woody growth. Mutualist ants protected pine-feeding aphids from predatory arthropods, allowing aphid populations to burgeon in cages even though predatory arthropods also increased in cages. By protecting pine-feeding aphids from predatory arthropods but not birds, mutualist ants created a three-tiered linear food chain where bird effects cascaded to pine growth via aphids. 4. In contrast to the results for tended aphids on pine, bird exclusion had no net effects on untended pine herbivores, the proportion of pine foliage damaged by pine-feeding caterpillars, or the proportion of mistletoe plants damaged by mistletoe-feeding caterpillars. These results suggest that arthropod predators, which were more abundant in cages as compared with control trees, compensated for bird predation of untended pine and mistletoe herbivores. 5. These contrasting effects of bird exclusion support food web theory: where birds were connected to pine by a linear food chain, a trophic cascade occurred. Where birds fed as intraguild predators, the reticulate food webs linking birds to pine and mistletoe resulted in no net effects on herbivores or plant biomass. Our study shows that this variation in food web structure occurred between sympatric plants and within plants between differing herbivore guilds.     

Direct and indirect effects of predatory young‐of‐year fishes in a dryland river food web

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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.     

Understanding of colonization and breakdown of leaves by invertebrates in a tropical stream is enhanced by using biomass as well as count data

We hypothesized that (i) the importance of shredders for leaf breakdown is more evident in terms of their biomass than their abundance, due to the large bodies and high-feeding efficiencies of some typical shredders; (ii) non-shredder invertebrates select more refractory leaves because these are a more stable substrate for colonization or to obtain other forms of food. To test these hypotheses, we performed a decomposition experiment with leaves of contrasting chemical composition in a tropical stream, and determined the changes in the ash-free dry mass (AFDM) of the litter, and the invertebrate abundance and biomass during a 44-day period. The biomass of shredders showed a positive relationship with AFDM remaining, whereas their abundance was unrelated to AFDM. While shredder abundance represented only 4–12% of total invertebrate abundance, shredder biomass constituted 19–36% of total invertebrate biomass. We conclude that (i) shredder biomass expresses better than abundance the role of this guild in the decomposition of leaf detritus, demonstrating that they are important for the functioning of tropical streams; (ii) incubation time rather than stability of leaf litter as a substrate influences colonization by non-shredder invertebrates.     

Riparian plant species loss alters trophic dynamics in detritus-based stream ecosystems

Riparian vegetation is closely connected to stream food webs through input of leaf detritus as a primary energy supply, and therefore, any alteration of plant diversity may influence aquatic ecosystem functioning. We measured leaf litter breakdown rate and associated biological parameters in mesh bags in eight headwater streams bordered either with mixed deciduous forest or with beech forest. The variety of leaf litter types in mixed forest results in higher food quality for large-particle invertebrate detritivores (‘shredders’) than in beech forest, which is dominated by a single leaf species of low quality. Breakdown rate of low quality (oak) leaf litter in coarse mesh bags was lower in beech forest streams than in mixed forest streams, a consequence of lower shredder biomass. In contrast, high quality (alder) leaf litter broke down at similar rates in both stream categories as a result of similar shredder biomass in coarse mesh bags. Microbial breakdown rate of oak and alder leaves, determined in fine mesh bags, did not differ between the stream categories. We found however aquatic hyphomycete species richness on leaf litter to positively co-vary with riparian plant species richness. Fungal species richness may enhance leaf litter breakdown rate through positive effects on resource quality for shredders. A feeding experiment established a positive relationship between fungal species richness per se and leaf litter consumption rate by an amphipod shredder (Gammarus fossarum). Our results show therefore that plant species richness may indirectly govern ecosystem functioning through complex trophic interactions. Integrating microbial diversity and trophic dynamics would considerably improve the prediction of the consequences of species loss.     

Leaf breakdown and associated macroinvertebrates in alpine glacial streams

1. The relationship between macroinvertebrate assemblages and the breakdown of alder [Alnus viridis (Chaix), Dc.] leaves was examined by exposing leaf packs in four streams in an alpine glacial floodplain over 8 months. Although glacially fed, the four sites (pro-glacial, glacial lake outlet, main channel, and a side-channel with a mix of water sources) differed physically and contained different benthic communities.
2. Leaf breakdown and associated fungal properties differed widely among sites. Leaf decay rate varied by an order of magnitude ( k ranged from 0.0029 to 0.0305 day^–1), and was fastest at the lake outlet (< 20% leaf mass remaining by day 45) and slowest at the pro-glacial site (> 75% remaining on day 45). Rapid processing at the lake outlet was because of the presence of Acrophylax zerberus Brauer, a shredding caddisfly.
3. There were few macroinvertebrate taxa at the pro-glacial site (two to four taxa present in packs) and leaf breakdown was attributed primarily to micro-organisms. Leuctra abundance in leaf packs was strongly correlated with fungal biomass but not with the sporulation activity of any specific aquatic hyphomycete. Other taxa, such as Baetis and chironomids, showed no relationship with any leaf characteristic, suggesting that leaf packs were used mainly as a habitat and not as a food resource.
4. The predatory stonefly Isoperla was significantly associated with the abundance of macroinvertebrate prey ( Baetis , Chironomidae and Leuctra ) in leaf packs at the main and side-channel sites. The results indicate that leaf breakdown can vary widely in alpine lotic environments, reflecting site-specific differences in habitat characteristics, and in macroinvertebrate and fungal composition.     

Fish trophic structure in estuaries,with particular emphasis on estuarine typology and zoogeography

A comparative analysis of the fish trophic structure was undertaken on some 190 South African estuaries spanning three zoogeographic regions and incorporating three broad estuarine types. Fish biomass trophic guild compositions and biomass trophic spectrum profiles were analysed using multivariate statistical techniques and included both inter‐regional (zoogeographic) and intra‐regional (estuarine typology) comparisons. Differences in the fish trophic structure of the various estuary types within each zoogeographic region were observed; these were linked to the relative biomass contribution of the various trophic guilds and also to differences in biomass trophic spectrum profiles of the fishes in each estuary type within each region. In spite of these differences in trophic structure, all estuaries were dominated by detritivores, which suggests that the main food source (detritus) is similar in all biogeographic regions. Preliminary indications are that a similar dependence by estuary‐associated fishes on detritus food sources exists on a global basis but that detailed studies are required in order to confirm this assertion.     

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.     

Feeding of fish young from littoral of the gulf Arguin (Mauritania)

The results of ichthyologic investigations on littoral of the gulf Arguin in 2000–2004 are represented in this article. All main trophic fish groups—phytophagans, planktonphagans, benthophagans and predators—were discovered in the zone of littoral. The transfer to another trophic group is marked for the young of the most species while growth. Detritus and detritus-algal units have a special place in fish young diet. Sea grasses with covering area in the gulf of 390–490 km² are the origin of detritus. Favourable environmental conditions and high food provision make the gulf Arguin vitally important place for existense of neritic ichthyocene of northern Africa, the place for young fish reproduction and pasture.     

APPLIED ISSUES Increasing litter retention in moorland streams: ecological and management aspects of a field experiment

1. The retention characteristics of two moorland streams in mid-Wales were manipulated for 2 years by the addition of small traps which accumulated detritus. Leaf litter was also added to these essentially treeless streams at regular intervals to simulate natural inputs to a woodland stream. 2. Leaf traps retained a significantly higher biomass of detritus - both naturally occurring and added - than either the surrounding stream bed or unmanipulated reference sections. They also supported significantly higher numbers of nemourid, leuctrid, tipulid and elminthid insect larvae. 3. Among functional feeding groups, detritivores and, to a lesser extent, predators responded to increased detritus availability. Numbers of mayfly larvae were low in leaf packs, suggesting a negative effect of detritus aggregations on their numbers. 4. The taxa which responded positively to increased detrital biomass, particularly stonefly larvae, are known to be tolerant of low pH conditions, whereas those affected detrimentally are generally absent from acid waters. It is proposed therefore that increasing detrital inputs and litter retention in culturally acidified upland streams may serve to increase their invertebrate productivity.     

Trophic interactions among native and introduced fishes in a northern Patagonian oligotrophic lake

In summer the littoral zone of Lake Rivadavia contained almost all the juveniles of the native fish species present in the lake together with juveniles of the introduced salmonid Oncorhynchus mykiss . In particular, a high degree of foraging interaction was found among juveniles of the native species Aplochiton zebra and Percichthys trucha and juveniles O. mykiss. Differences in the juvenile assemblage composition, both in the presence and in the relative proportion of the species were associated with areas with different densities of vegetation. The adults of nearly all species were generalized benthic invertebrate predators. Only adults of A. zebra were absent in the littoral. Juvenile and adult O. mykiss were the only fish that preyed on planktonic organisms. Oncorhynchus mykiss was the only species that performed an ontogenetic diet shift between juveniles and adults, changing main prey categories from benthic to planktonic species.     

The diets and parasites of larval and 0+ juvenile twaite shad in the lower reaches and estuaries of the rivers Wye,Usk and Towy,UK

Aquatic predators and habitat permanence can jointly affect benthic invertebrate biomass and community composition. In 2006 I sampled fish and invertebrates in ten ponds embedded in a seasonal wetland before and after a natural drought. Drought reduced fish biomass and density leaving some ponds in a fishless condition when rains returned in July. In July, large aquatic insects and crayfish colonized and reproduced in the ponds, but did not colonize all of the ponds equally. Using measurements of fish abundance and other environmental parameters of the ponds, I conducted linear regression analyses to explore potential drivers of variable invertebrate biomass in July. Fish biomass had a negative effect on invertebrate biomass and it explained more of the variation in total invertebrate biomass and total non-shrimp biomass than fish abundance (number of fish caught). Dissolved oxygen and pond depth were both correlated with fish biomass, but were poorer predictors of invertebrate biomass. Ponds with few or no fish had 20× greater total biomass and 200× more non-shrimp biomass than ponds with high fish biomass. Shrimp dominated the invertebrate composition, and were only found in the two deepest ponds with the highest fish biomass; predatory insects and crayfish dominated the other eight ponds. When taxa were analyzed separately, fish biomass explained a large portion of the variation for predatory insects (Coleoptera, Hemiptera, and Odonata) and crayfish (Procambarus alleni), but dissolved oxygen was the best predictor of larval stratiomyid (order Diptera) biomass. These results are generally consistent with studies demonstrating negative effects of fish on large predatory invertebrates, but also suggest that more severe local droughts can seasonally enhance insect and crayfish populations by generating fishless or nearly fishless conditions. Handling editor: J. Trexler     

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

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