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.     

Diversity patterns of leaf-associated aquatic hyphomycetes along a broad latitudinal gradient

Information about the global distribution of aquatic hyphomycetes is scarce, despite the primary importance of these fungi in stream ecosystem functioning. In particular, the relationship between their diversity and latitude remains unclear, due to a lack of coordinated surveys across broad latitudinal ranges. This study is a first report on latitudinal patterns of aquatic hyphomycete diversity associated with native leaf-litter species in five streams located along a gradient extending from the subarctic to the tropics. Exposure of leaf litter in mesh bags of three different mesh sizes facilitated assessing the effects of including or excluding different size-classes of litter-consuming invertebrates. Aquatic hyphomycete evenness was notably constant across all sites, whereas species richness and diversity, expressed as the Hill number, reached a maximum at mid-latitudes (Mediterranean and temperate streams). These latitudinal patterns were consistent across litter species, despite a notable influence of litter identity on fungal communities at the local scale. As a result, the bell-shaped distribution of species richness and Hill diversity deviated markedly from the latitudinal patterns of most other groups of organisms. Differences in the body-size distribution of invertebrate communities colonizing the leaves had no effect on aquatic hyphomycete species richness, Hill diversity or evenness, but invertebrates could still influence fungal communities by depleting litter, an effect that was not captured by the design of our experiment.     

Exotic riparian vegetation lowers fungal diversity but not leaf decomposition in Portuguese streams

1. We characterised the fungal communities of eight streams in Portugal, four bordered by native deciduous forest and four bordered by pure stands of Eucalyptus globulus .
2. Aquatic hyphomycete species richness and evenness, but not numbers of water-borne conidia, of aquatic hyphomycetes were significantly lower in eucalypt bordered streams.
3. Multivariate analyses subdivided the fungal communities into two distinct groups corresponding to riparian vegetation.
4. Despite these differences in the dominant decomposer community, decay rates of eucalypt leaves (accounting for ≥98% of naturally occurring leaves in eucalypt bordered streams, absent in native forest) and chestnut leaves (occurring naturally in native forests) did not differ between the two groups of streams.     

Microscopy- or DNA-based analyses: Which methodology gives a truer picture of stream-dwelling decomposer fungal diversity?

We assessed aquatic hyphomycete diversity in autumn and spring on oak leaves decomposing in five streams along a gradient of eutrophication in the Northwest of Portugal. Diversity was assessed through microscopy-based (identification by spore morphology) and DNA-based techniques (Denaturing Gradient Gel Electrophoresis and 454 pyrosequencing). Pyrosequencing revealed five times greater diversity than DGGE. About 21% of all aquatic hyphomycete species were exclusively detected by pyrosequencing and 26% exclusively by spore identification. In some streams, more than half of the recorded species would have remained undetected if we had relied only on spore identification. Nevertheless, in spring aquatic hyphomycete diversity was higher based on spore identification, probably because many species occurring in this season are not yet connected to ITS barcodes in genetic databases. Pyrosequencing was a powerful tool for revealing aquatic hyphomycete diversity on decomposing plant litter in streams and we strongly encourage researchers to continue the effort in barcoding fungal species.     

Copper and zinc mixtures induce shifts in microbial communities and reduce leaf litter decomposition in streams

1. To assess the impact of metal mixtures on microbial decomposition of leaf litter, we exposed leaves previously immersed in a stream to environmentally realistic concentrations of copper (Cu) and zinc (Zn) (three levels), alone and in all possible combinations. The response of the microbial community was monitored after 10, 25 and 40 days of metal exposure by examining leaf mass loss, fungal and bacterial biomass, fungal reproduction and fungal and bacterial diversity.
2. Analysis of microbial diversity, assessed by denaturing gradient gel electrophoresis and identification of fungal spores, indicated that metal exposure altered the structure of fungal and bacterial communities on decomposing leaves.
3. Exposure to metal mixtures or to the highest Cu concentration significantly reduced leaf decomposition rates and fungal reproduction, but not fungal biomass. Bacterial biomass was strongly inhibited by all metal treatments.
4. The effects of Cu and Zn mixtures on microbial decomposition of leaf litter were mostly additive, because observed effects did not differ from those expected as the sum of single metal effects. However, antagonistic effects on bacterial biomass were found in all metal combinations and on fungal reproduction in metal combinations with the highest Cu concentrations, particularly at longer exposure times.     

Microbial growth and detritus transformations during decomposition of leaf litter in a stream

SUMMARY. 1. Despite the widely accepted importance of bacteria and fungi in degrading detritus in aquatic ecosystems there is still very little quantitative information on the abundance and dynamics of these microorganisms. Using epifluorescent microscopy, we measured the biomass of bacteria and fungi during decomposition of three types of leaf detritus. Bacterial production was determined from the rate of incorporation of ³H-thymidine into DNA.
2. The transformation of leaf carbon into dissolved organic carbon and fine particulate organic carbon was followed in order to compare the amounts of leaf material that were converted into these 'end-products' of decomposition versus the amount converted into microbial biomass.
3. The amount of microbial carbon in the leaf-detritus complex never exceeded 5.2% of the total carbon, and fungal biomass was always much greater than bacterial biomass. Despite the greater standing stock of fungi, the rapid turnover of bacteria (doubling about once per day) implies that their role in degrading leaf litter or as a food source for detritivores might be as great as for fungi.
4. Removal of microbial biomass from leaf litter may occur as release of fungal spores and consumption or shedding of bacterial biomass. Fungal spores can be a significant part of the fine particulate organic carbon released from leaf detritus and potentially represent an important food resource for filter-feeding organisms.     

Detritivores in Kenyan highland streams: more evidence for the paucity of shredders in the tropics?

1. The relationship between coarse particulate organic matter (CPOM) standing stock and benthic macroinvertebrate assemblages in Kenyan highland streams was determined by sampling seven sites on three rivers (2000–2700 m a.s.l.). Taxa recorded were allocated to functional feeding groups using published literature, mouthpart analysis and examination of gut contents. Patterns were compared with five structurally similar streams in three areas of Europe (south-west France, south-east England, north-east England).
2. Number of individuals and proportion of detritivores in Kenyan streams were equivalent to, or greater than, those in European sites. Shredders were, however, almost completely absent from Kenyan sites, despite high standing stocks of CPOM. Shredders were abundant in all European sites.
3. The phenomenon of low shredder abundance has been observed in other tropical streams in south-east Asia and Central and South America but, in contrast to these regions, the African rivers studied were devoid of shrimps or fish which may occupy the shredding niche elsewhere.
4. These preliminary data suggest that shredder-mediated detritus processing, which is a key functional component of streams in the North Temperate Zone, does not operate in East African streams. There are three possible reasons for this. The first is that tropical African rivers are functionally different to those in temperate regions. This could be because of enhanced microbial activity replacing shredder activity at high temperatures. Alternatively, it could be a result of low palatability of detrital inputs from dominant riparian trees in the region. The second and third are methodological: that our allocation to functional feeding groups is incorrect, and that our sampling methods missed a potentially key shredding taxon – the freshwater crab Potamonautes sp.     

Fungal communities on decaying leaves in streams: a comparison of two leaf species

Decomposition of leaf litter is a microbial mediated process that helps to transfer energy and nutrients from leaves to higher trophic levels in woodland streams. Generally, aquatic hyphomycetes are viewed as the major fungal group responsible for leaf litter decomposition. In this study, traditional microscopic examination (based on identification of released conidia) and phylogenetic analysis of 18S rRNA genes from cultivated fungi were used to compare fungal community composition on decomposing leaves of two species (sugar maple and white oak) from a NE Ohio stream. No significant differences were found in sporulation rates between maple and oak leaves and both had similar species diversity. From the 18S rRNA gene sequence data, identification was achieved for 12 isolates and taxonomic affiliation of 12 of the remaining 14 isolates could be obtained. A neighbor-joining tree (with bootstrap values) was constructed to examine the taxonomic distribution of the isolates relative to sequences of known operational taxonomic units (OTUs). Surprisingly, only 2 of the isolates obtained were aquatic hyphomycetes based on phylogenetic analysis. Overall, there were no differences between the two leaf types and a higher diversity was observed via culturing and subsequent 18S rRNA gene sequencing than by conidia staining. These differences resulted from the fact that traditional microscopy provides estimates of aquatic hyphomycete diversity while the other approach revealed the presence of both aquatic hyphomycete and non-aquatic hyphomycete taxa. The presence of this broad array of taxa suggests that the role of aquatic hyphomycetes relative to other fungi be re-evaluated. Even though the functional role of these non-aquatic hyphomycetes taxa is unknown, their presence and diversity demonstrates the need to delve further into fungal community structure on decomposing leaves.     

Aquatic Hyphomycete Species Are Screened by the Hyporheic Zone of Woodland Streams

Aquatic hyphomycetes strongly contribute to organic matter dynamics in streams, but their abilities to colonize leaf litter buried in streambed sediments remain unexplored. Here, we conducted field and laboratory experiments (slow-filtration columns and stream-simulating microcosms) to test the following hypotheses: (i) that the hyporheic habitat acting as a physical sieve for spores filters out unsuccessful strategists from a potential species pool, (ii) that decreased pore size in sediments reduces species dispersal efficiency in the interstitial water, and (iii) that the physicochemical conditions prevailing in the hyporheic habitat will influence fungal community structure. Our field study showed that spore abundance and species diversity were consistently reduced in the interstitial water compared with surface water within three differing streams. Significant differences occurred among aquatic hyphomycetes, with dispersal efficiency of filiform-spore species being much higher than those with compact or branched/tetraradiate spores. This pattern was remarkably consistent with those found in laboratory experiments that tested the influence of sediment pore size on spore dispersal in microcosms. Furthermore, leaves inoculated in a stream and incubated in slow-filtration columns exhibited a fungal assemblage dominated by only two species, while five species were codominant on leaves from the stream-simulating microcosms. Results of this study highlight that the hyporheic zone exerts two types of selection pressure on the aquatic hyphomycete community, a physiological stress and a physical screening of the benthic spore pool, both leading to drastic changes in the structure of fungal community.     

Contribution of fungal biomass to the growth of the shredder, Pycnopsyche gentilis (Trichoptera: Limnephilidae)

1.  1. It has been accepted that aquatic hyphomycetes colonising submerged leaves increase the nutritional value of leaf detritus and suggested that fungal biomass plays a greater role in the growth of shredders than leaf tissue itself. However, it is not clear what proportion of the nutritional needs of shredders is met by fungal biomass.
2.  We fed Pycnopsyche gentilis larvae with tulip poplar ( Liriodendron tulipifera ) leaf discs colonised by the aquatic hyphomycete, Anguillospora filiformis , which had been radiolabelled to quantify the contribution of fungal carbon to the growth of the shredder at different larval developmental stages. Instantaneous growth rates of larvae on this diet were also estimated.
3.  When provided with fungal-colonised leaves (14–16% fungal biomass), the third and the fifth instar larvae of P. gentilis grew at the rates of 0.061 and 0.034 day⁻¹, respectively, but on a diet of sterile leaves, both larval instars lost weight. The incorporation rates of fungal carbon were 31.6 μg C mg⁻¹ AFDM day⁻¹, accounting for 100% of the daily growth rate of the third instar larvae and 8.6 μg C mg⁻¹ AFDM day⁻¹, accounting for 50% of the daily growth rate of the fifth instar larvae.
4.  These results suggest that leaf material colonised by A. filiformis is a high quality food resource for P. gentilis larvae, and that fungal biomass can contribute significantly to the growth of these larvae. Differences in feeding behaviour and digestive physiology may explain the significantly greater assimilation of fungal biomass by the earlier instar than the final instar. To satisfy their nutritional needs the fifth instar larvae would have to assimilate detrital mass that may have been modified by fungal exoenzymes.     

Physico-chemistry and aquatic insects of a glacier-fed and a spring-fed alpine stream

1. Physico-chemical conditions and benthic macroinvertebrates were studied in two adjacent alpine streams in the Tyrolean Alps, Austria, for 2 years, and aquatic insect emergence was recorded for 1 year.
2. In the spring-fed system, maximum discharge and increased concentrations of suspended solids, nitrate and particulate phosphorus occurred during snowmelt in June. In the glacier-fed stream, high discharge and strong diel fluctuations in flow and concentrations of suspended solids created a harsh and unstable environment during summer. Glacial ablation, variation in groundwater inflow, and water inputs from tributaries draining calcareous rocks caused water chemistry to vary both seasonally and longitudinally in glacier-fed Rotmoosache.
3. A total of 126 aquatic or semi-aquatic invertebrate taxa were collected, 94 of which were found in the glacier-fed stream and 120 in the spring-fed stream. Chironomid abundance was 2–8 times and taxa richness 2–3 times lower in the glacier-fed stream than in the spring-fed stream, as was the number of chironomid taxa (72 versus 93 total).
4. These results broadly support the conceptual model by Milner & Petts (1994) concerning glacier-fed stream systems. However, single samples and seasonal means showed relatively high invertebrate abundance and richness, especially during winter, indicating a considerable degree of spatial and temporal variability.
5. We suggest that the seasonal shifts from harsh environmental conditions in summer to less severe conditions in autumn and a rather constant environment in winter are an important factor affecting larval development, life-history patterns and the maintenance of relatively high levels of diversity and productivity in glacier-fed streams.     

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     

Macroinvertebrate diversity in headwater streams: a review

1. Headwater streams are ubiquitous in the landscape and are important sources of water, sediments and biota for downstream reaches. They are critical sites for organic matter processing and nutrient cycling, and may be vital for maintaining the 'health' of whole river networks.
2. Macroinvertebrates are an important component of biodiversity in stream ecosystems and studies of macroinvertebrate diversity in headwater streams have mostly viewed stream systems as linear reaches rather than as networks, although the latter may be more appropriate to the study of diversity patterns in headwater systems.
3. Studies of macroinvertebrate diversity in headwater streams from around the world illustrated that taxonomic richness is highly variable among continents and regions, and studies addressing longitudinal changes in taxonomic richness of macroinvertebrates generally found highest richness in mid-order streams.
4. When stream systems are viewed as networks at the landscape-scale, α-diversity may be low in individual headwater streams but high β-diversity among headwater streams within catchments and among catchments may generate high γ-diversity.
5. Differing ability and opportunity for dispersal of macroinvertebrates, great physical habitat heterogeneity in headwater streams, and a wide range in local environmental conditions may all contribute to high β-diversity among headwater streams both within and among catchments.
6. Moving beyond linear conceptual models of stream ecosystems to consider the role that spatial structure of river networks might play in determining diversity patterns at the landscape scale is a promising avenue for future research.     

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.

     

Benthic detritus dynamics in a mountain stream

The dynamics of sedimentary detritus was examined over a 17-month period in a third-order mountain stream in Colorado. Autumnal and vernal peaks in total detritus standing crop (197 g and 165 g AFDW m⁻², respectively) were associated with leaf abscission and ice out. The flushing action of snowmelt runoff rapidly reduced benthic detritus to 14 g AFDW m⁻². Detrital size classes differentially contributed to temporal fluctuations in total detritus. The autumnal maximum was largely comprised of coarse particulate organic matter (CPOM), primarily leaf litter. The large increase following ice out was mainly due to fine detritus (FPOM, 0.45–75 μm) which accounted for > 50% of benthic detritus during 9 months of the year. Evidence suggests that changes in standing crop and particle size distribution during the extended period of ice cover (November-March) resulted from in situ processing rather than transport. Despite major differences in climate, geochemistry and hydrology, the Colorado stream exhibited patterns of detrital dynamics remarkably similar to an eastern deciduous forest stream.     

Filters and templates: stonefly (Plecoptera) richness in Ouachita Mountains streams, U.S.A.

1.  We collected adult stoneflies periodically over a 1-year period at 38 sites in two headwater catchments in the Ouachita Mountains, Arkansas, U.S.A. The 43 species collected were a subset of the Ozark-Ouachita fauna and the much larger fauna of the eastern U.S.A. We estimated 78–91% species coverage in the two catchments using jackknife extrapolation of species richness from our survey.
2.  Many streams, especially small ones, lacked surface water for months, but others, both small and large, flowed permanently.
3.  Using published regional presence–absence and coarse ecological data in a discriminant function analysis (DFA), we identified stream size (negative) and regional frequency of occurrence (positive) as predictors of presence in these headwater catchments. For the combined catchments, the extrapolated richness (51 spp.) was similar to an estimate (48 spp.) based on predicted absences from DFA and the Ouachita provincial total of known stonefly species (57 spp.).
4.  Local species richness (1–27 spp. per site) was correlated strongly with stream size (catchment area) but was independent of stream drying. Generic richness was correlated negatively with stream drying and positively, but less strongly, with stream size.
5.  Regionally endemic stoneflies dominated in drying streams, and widely distributed species dominated in more permanent streams. The composition of stonefly assemblages was associated with regional factors (species pools, regional abundance, evolution of tolerant endemic species, regional climate) and local factors (drying, stream size).     

Growth of an invertebrate shredder on native (Populus) and non-native (Tamarix, Elaeagnus) leaf litter

1. Large-scale invasions of riparian trees can alter the quantity and quality of allochthonous inputs of leaf litter to streams and thus have the potential to alter stream organic matter dynamics. Non-native saltcedar ( Tamarix sp.) and Russian olive ( Elaeagnus angustifolia ) are now among the most common trees in riparian zones in western North America, yet their impacts on energy flow in streams are virtually unknown.
2. We conducted a laboratory feeding experiment to compare the growth of the aquatic crane fly Tipula (Diptera: Tipulidae) on leaf litter from native cottonwood ( Populus ) and non-native Tamarix and Elaeagnus . Tipula showed positive growth on leaf litter of all three species; however, after 7 weeks, larvae fed Tamarix leaves averaged 1.7 and 2.5 times the mass of those fed Elaeagnus and Populus , respectively. Tipula survival was highest on Populus , intermediate on Tamarix and lowest on Elaeagnus .
3. High Tipula growth on Tamarix probably reflects a combination of leaf chemistry and morphology. Conditioned Tamarix leaf litter had intermediate carbon : nitrogen values (33 : 1) compared to Populus (40 : 1) and Elaeagnus (26 : 1), and it had intermediate proportions of structural carbon (42%) compared to Elaeagnus (57%) and Populus (35%). Tamarix leaves are also relatively small and possibly more easily ingested by Tipula than either Elaeagnus or Populus .
4. Field surveys of streams in the western U.S.A. revealed that Tamarix and Elaeagnus leaf packs were rare compared to native Populus , probably due to the elongate shape and small size of the non-native leaves. Thus we conclude that, in general, the impact of non-native riparian invasion on aquatic shredders will depend not only on leaf decomposition rate and palatability but also on rates of leaf litter input to the stream coupled with streambed retention and subsequent availability to consumers.     

Intraspecific diversity affects stress response and the ecological performance of a cosmopolitan aquatic fungus

Fungi play an important role in organic matter turnover and ensure key ecosystem services in freshwaters. The relationships between intraspecific fungal diversity and key ecological processes remain largely unknown. We examined the effects of intraspecific diversity of Articulospora tetracladia, a cosmopolitan fungal decomposer thriving on plant detritus in streams. Alder leaves were inoculated with 1 or mixtures of 2–8 fungal strains for 35 d, and leaf litter decomposition and fungal reproduction were quantified in the presence and absence of 2 mg L⁻¹ of cadmium (Cd), a common stressor in polluted streams. Intraspecific diversity and identity affected fungal reproduction, but not leaf decomposition. Under metal stress, leaf decomposition slightly increased with intraspecific diversity. Fungal reproduction increased with intraspecific diversity and was greater in mixed assemblages, either in the absence or presence of Cd. Effect size of intraspecific diversity was higher under Cd stress for fungal reproduction, but no differences were found for leaf mass loss, with or without metal. The impacts of intraspecific diversity loss may jeopardize fungal survival and fungal functions, namely microbial leaf decomposition and leaf litter condition for invertebrate shredders in streams, particularly under metal stress.     

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.     

Limited evidence that mixing leaf litter accelerates decomposition or increases diversity of decomposers in streams of eastern Canada

Most studies of terrestrial litter decomposition in streams and rivers have used leaves from a single tree species, but leaf packs in streams in eastern North America are usually mixtures of two or more species. Litter mixtures may decay more quickly than either of the component species. If so, estimates of stream energy and nutrient budgets may be inaccurate. In northern Nova Scotia, Canada, we measured mass loss from binary mixtures (1:1 mass ratio) of leaf litter in mesh bags, using freshly fallen or air-dried litter from five species of canopy trees. We repeated the experiment eight times, in summer and fall, in two streams and a small river, over 3 years. In some trials we enumerated benthic invertebrate and fungal colonization of decaying litter. Although there were marked differences in mass loss rates among litter types, decomposition was accelerated in mixtures relative to the mean of the component species in only three of eight trials, and only in mixtures containing N-rich speckled alder leaves. Mixing yellow birch and red maple leaves inhibited decomposition. Diversity (Shannon–Weaver Index), species richness, and abundance of aquatic hyphomycete fungi, as indexed by conidial production, were never greater (and sometimes less) on litter mixtures than on the component species. Total numbers, taxonomic richness and diversity of benthic invertebrates generally, and litter-feeding species in particular, were not augmented by mixing litter types. Litter mixtures appear to dilute a preferred substrate with patches of a less preferred substrate. Our results provide only weak support for the contention that combining two litter types leads to acceleration of decomposition rates. Handling editor: K. Martens