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     

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.     

Effects of consumers and nitrogen availability on heterotrophic microbial activity during leaf decomposition in headwater streams

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Land use change affects macroinvertebrate community size spectrum in streams: the case of Pinus radiata plantations

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Colonization and processing of leaf litter by macroinvertebrate shredders in streams of contrasting pH

• 1 Leaf litter processing rates and macroinvertebrate shredder assemblages in leaf packs were compared in four streams on the Allegheny plateau in the central Appalachian Mountains, U.S.A.; these streams were characterized by different bedrock geology and streamwater pH.
• 2 Leaf litter processing rates were fastest in the neutral streams, slowest in the acidic stream, and intermediate in the most alkaline stream.
• 3 Slower processing rates in the acidic stream were associated with lower total shredder biomass, made up predominantly by small leuctrid and nemourid stoneflies.
• 4 The differences in processing rates between the more alkaline stream and the neutral streams were not associated with differences in shredder biomass, but appeared to be related to taxonomic differences in the shredder assembiages. Insects were dominant in the neutral streams, and amphipods were dominant in the more alkaline stream.

     

An assessment of mesh bags and plastic leaf traps as tools for studying macroinvertebrate assemblages in natural leaf packs

Aggregation of leat litter formed against small mesh obstacles — placebo traps — were studied in four streams differing in natural retentiveness and pH. In three of the streams, natural benthic accumulations of leaf litter were available for comparison, and in these the fauna in the plastic traps and the natural accumulations was similar. In two of the streams comparisons were made, in terms of percent composition, between the fauna of the stony benthos and that colonizing plastic traps and leaf-filled mesh bags. In an acid, naturally retentive stream, the fauna of the three treatments was similar, although shredders were relatively more abundant in plastic traps and mesh bags. In a circumneutral, non-retentive stream diversity of taxa was reduced in plastic traps compared with the stony benthos, and in mesh bags compared with plastic traps. Numbers of animals per g of leaf litter were similar in plastic traps and mesh bags in the retentive stream. In the non-retentive stream, however, there were fewer animals in mesh bags than in the plastic traps. For many purposes, the plastic traps produce leaf packs which closely mimic natural packs, but the results from mesh bags depend on the background retentiveness of the streams in question.     

Stronger effects of litter origin on the processing of conifer than broadleaf leaves: A test of home‐field advantage of stream litter breakdown

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Aggregation among resource patches can promote coexistence in stream-living shredders

1. We conducted a new single‐site study and a meta‐analysis of pre‐existing studies from multiple streams to assess whether intraspecific aggregation among leaf packs promotes coexistence among leaf‐eating invertebrates (shredders) in Swedish streams. 2. In the single‐site study, 48 standardised leaf bags were exposed for 1 month for shredder colonisation in a homogeneous glide of a forested stream. Current velocity, water depth and substratum composition were additionally assessed to investigate how these factors affected shredder distributions. 3. The meta‐analysis included information on shredder colonisation of leaf packs from seven other studies of detritus decomposition to assess patterns of aggregation. Intra‐ and interspecific aggregation and their relative strength were assessed using indices (J, C and A respectively) originally developed for terrestrial insects also dependent on patchy and ephemeral resources. 4. In both parts of the study, intraspecific aggregation was much stronger than interspecific aggregation, which was weak overall, indicating that the conditions under which aggregation is expected to facilitate coexistence were fulfilled in our shredder assemblages. 5. In the single‐site study, shredder abundances were weakly associated with environmental variables suggesting that habitat heterogeneity only partly explains aggregation patterns. 6. Our results strongly suggest that shredder diversity in streams, particularly during periods of leaf limitation (such as might occur in spring), is promoted by the aggregation of individual species among patches of resource.     

Assessing the influence of wildfire on leaf decomposition and macroinvertebrate communities in boreal streams using mixed-species leaf packs

1. We investigated how compositional differences in riparian leaf litter derived from burned and undisturbed forests influenced leaf breakdown and macroinvertebrate communities using experimental mixed-species leaf packs in boreal headwater streams. Leaf pack mixtures simulating leaf litter from dominant riparian woody-stem species in burned and undisturbed riparian zones were incubated in two references and two fire-disturbed streams for 5 weeks prior to measuring temperature-corrected breakdown rates and macroinvertebrate community composition, richness, and functional metrics associated with decomposers such as shredder abundance and % shredders.
2. Leaf litter breakdown rates were higher and had greater variability in streams bordered by reference riparian forests than in streams where riparian forests had been burned during a wildfire. Streams bordered by fire disturbance showed significant effects of litter mixture on decomposition rates, observed as significantly higher decomposition rates of a fire-simulated leaf mixture compared to all other mixtures.
3. Variation among sites was higher than variation among litter mixtures, especially for macroinvertebrate community composition. In general, fire-simulated leaf mixtures had greater shredder abundances and proportions, but lower overall macroinvertebrate abundance; however, the shredder abundance trend was not consistent across all leaf mixtures at each stream.
4. These results show that disturbance-driven riparian forest condition and resulting composition of leaf subsidies to streams can influence aquatic invertebrate community composition and their function as decomposers. Therefore, if one of the primary goals of modern forest management is to emulate natural disturbance patterns, boreal forest managers should adapt silvicultural practices to promote leaf litter input that would arise post-fire, thereby supporting stream invertebrate communities and their function.

     

Detrital processing in streams exposed to acidic precipitation in the Central Appalachian Mountains

Continuing high rates of acidic deposition in the eastern United States may lead to long-term effects on stream communities, because sensitive catchments are continuing to lose anions and cations. We conducted a two-year study of the effects of pH and associated water chemistry variables on detrital processing in three streams with different bedrock geology in the Monongahela National Forest, West Virginia. We compared leaf pack processing rates and macroinvertebrate colonization and microbial biomass (ATP concentration) on the packs in the three streams. Breakdown rates of red maple and white oak leaf packs were significantly lower in the most acidic stream. The acidic stream also had significantly lower microbial and shredder biomass than two more circumneutral streams. Shredder composition differed among streams; large-particle detritivores dominated the shredder assemblages of the two circumneutral streams, and smaller shredders dominated in the acidic stream. Within streams, processing rates for three leaf species were not significantly different between the two years of the study even though invertebrate and microbial communities were different in the two years. Thus, macroinvertebrate and microbial communities differed both among streams that differed in their capacity to buffer the effects of acidic precipitation and among years in the same stream; these differences in biotic communities were not large enough to affect rates of leaf processing between the two years of the study, but they did significantly affect processing rates between acidic and circumneutral streams.The Unit is jointly sponsored by the National Biological Service, the West Virginian Division of Natural Resources, West Virginia University, and the Wildlife Management Institute.The Unit is jointly sponsored by the National Biological Service, the West Virginian Division of Natural Resources, West Virginia University, and the Wildlife Management Institute.     

Grass,mosses, algae,or leaves? Food preference among shredders from open-canopy streams

Shredder feeding is a vital process in making decomposition products available to biota in streams. To investigate which food sources shredders in open-canopy streams exploit, we conducted a feeding preference experiment with the invertebrate detritivores Limnephilus bipunctatus and Nemoura sp., which are commonly found in open-canopy streams on the Swedish island of Öland in the southern Baltic Sea. Leaves of birch, Swedish whitebeam, and shrubby cinquefoil; dead and fresh grass; water moss; and algae were offered to the shredders in multi- and single treatments. We hypothesized that food with high nutritional value would be preferred. Both taxa preferred leaves of shrubby cinquefoil, a bush common in the riparian zone of Öland streams; additionally Nemoura sp. also chose algae. Dead grass, the most abundant food source in the streams during the whole year, was the least consumed food type. The fresh food types had highest nutritional value, measured as carbon to nitrogen content. Therefore, food quality could not alone explain the preference of shrubby cinquefoil. However, among the detritus type offered, shrubby cinquefoil had the highest nutritional value. Shrubby cinquefoil may constitute one important energy source to these open-canopy stream ecosystems and may be essential in maintaining an abundant shredder community in these streams. Thus, the results of this study indicate that detrital resources are indeed important in open-canopy stream systems.     

Spatial and seasonal associations of benthic macroinvertebrates and detritus in an Alaskan subarctic stream

Summary Seasonal and spatial patterns of benthic invertebrate abundance were examined in relation to benthic detritus in Monument Creek, an Alaskan subarctic stream. The total macroinvertebrate fauna showed a mid-summer low in abundance, increasing to seasonal highs in winter/early spring (November/May). Shredders were a small portion of the benthic fauna or leaf pack fauna in summer but increased rapidly (in biovolume) following autumnal leaf fall to dominate the fauna by early winter (October/November). Abundance was strongly correlated with quantity of detritus in the sample. Comparison of benthic macroinvertebrate densities from Alaskan streams with comparable data from temperate zone streams shows that Alaskan streams are similar to temperate zone streams in range of abundance. Each unit of benthic detritus in Monument Creek is associated with a relatively large number of small (low individual biomass) shredders compared to streams in temperate regions. Detrital resources in this subarctic stream were meager, compared to temperate streams, and appeared to strongly influence the spatial and temporal patterns of detritivores.     

Leaf litter decomposition and macroinvertebrate communities in headwater streams draining pine and hardwood catchments

Benthic invertebrates, litter decomposition, andlitterbag invertebrates were examined in streamsdraining pine monoculture and undisturbed hardwoodcatchments at the Coweeta Hydrologic Laboratory in thesouthern Appalachian Mountains, USA. Bimonthlybenthic samples were collected from a stream draininga pine catchment at Coweeta during 1992, and comparedto previously collected (1989–1990) benthic data froma stream draining an adjacent hardwood catchment. Litter decomposition and litterbag invertebrates wereexamined by placing litterbags filled with pine ormaple litter in streams draining pine catchments andhardwood catchments during 1992–1993 and 1993–1994. Total benthic invertebrate abundance and biomass inthe pine stream was ca. 57% and 74% that of thehardwood stream, respectively. Shredder biomass wasalso lower in the pine stream but, as a result ofhigher Leuctra spp. abundance, shredderabundance was higher in the pine stream than thehardwood stream. Decomposition rates of both pine andred maple litter were significantly faster in pinestreams than adjacent hardwood streams (p<0.05). Total shredder abundance, biomass, and production weresimilar in maple bags from pine and hardwood streams. However, trichopteran shredder abundance and biomass,and production of some trichopteran taxa such asLepidostoma spp., were significantly higher in maplelitterbags from pine streams than hardwood streams(p<0.05). In contrast, plecopteran shredders(mainly Tallaperla sp.) were more important inmaple litterbags from hardwood streams. Shredderswere well represented in pine litterbags from pinestreams, but low shredder values were obtained frompine litterbags in hardwood streams. Resultssuggest conversion of hardwood forest to pinemonoculture influences taxonomic composition of streaminvertebrates and litter decomposition dynamics. Although the impact of this landscape-leveldisturbance on invertebrate shredder communitiesappeared somewhat subtle, significant differences indecomposition dynamics indicate vital ecosystem-levelprocesses are altered in streams draining pinecatchments.     

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.     

Responses of invertebrates and algae of a boreal coniferous forest stream to experimental manipulation of leaf litter inputs and shading

Forest harvesting alters leaf litter inputs and shading of small streams. Most of the previous studies of harvesting effects are limited to coastal or deciduous forests, so here we consider a sub-boreal forest stream. To test the hypothesis that changes in light and litter inputs would affect the benthic community in these streams, we experimentally manipulated these variables in stream mesocosms. We used a 2 × 2 factorial design with light (shaded or full light) and leaf litter inputs (equivalent to a forested stream or one quarter that rate) as factors. The high leaf litter treatment resulted in differences in macroinvertebrate community composition and higher densities of two shredders, Limnephilus sp. and Podmosta sp., suggesting food limitation. Algal filaments were longer in the high light treatments indicating a change in periphyton composition. There were no significant differences in chlorophyll a or ash-free dry mass, suggesting that light was not limiting to periphyton. The community structure clearly shifted in response to both resources, although primarily to detrital inputs. These results provide evidence that changes to shading and leaf inputs to small streams can affect the benthos and may limit secondary production.     

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.     

Flow intermittency negatively affects three phylogenetically related shredder stoneflies by reducing CPOM availability in recently intermittent Alpine streams in SW-Italian Alps

Several Alpine streams are currently facing recurrent summer drying events with detrimental consequences on stream detritivores, i.e., shredders, due to negative effects via changes the organic matter (CPOM) availability. We examined the ecological requirements of three phylogenetically related shredder genera belonging to the family of Nemouridae (Plecoptera), namely Nemoura, Protonemura and Amphinemura, in 14 Alpine streams recently facing recurrent summer flow intermittency events. We evaluated the overlap among their ecological niches measured in terms of hydraulic stress, substrate composition, changes in CPOM availability and competition with other shredder taxa (i.e., presence of individuals of other shredders) and we examined potential changes in their ecological niches between permanent and intermittent sites. The ecological niches of Protonemura and Amphinemura overlap broadly, but not with Nemoura, suggesting only partial potential competition. The reduced CPOM availability decreased the individual abundance of the three genera in intermittent sites, where they consistently preferred microhabitats with high CPOM availability and low competition with other shredder taxa, possibly due to food limitation. Overall, our results emphasize how the negative effect of flow intermittency on shredders in Alpine streams is mainly due to the decrease in CPOM availability, with consequent potential bottom up effects on stream ecosystem functionality.

     

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.     

Litter decomposition in a temperate and a tropical stream: the effects of species mixing,litter quality and shredders

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Fungal alteration of the elemental composition of leaf litter affects shredder feeding activity

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