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.     

Freshwater crabs (Decapoda: Pseudothelphusidae) increase rates of leaf breakdown in a neotropical headwater stream

1. Freshwater crabs are the largest macroconsumers in many neotropical headwater streams, but few studies have examined their roles in ecosystem processes such as leaf litter breakdown. As omnivorous macroconsumers, freshwater crabs affect multiple trophic levels. They may directly increase leaf breakdown through fragmentation and consumption or indirectly decrease breakdown by consuming other macroinvertebrates, including shredders and detritivores.
2. In a headwater stream in Monteverde, Costa Rica, we conducted an in-stream experiment with 40 enclosures to quantify the effects of pseudothelphusid crabs on both leaf breakdown and macroinvertebrate colonisation of leaves. Half of the enclosures were randomly selected to contain two crabs (mean carapace width = 30 mm) and half were controls without crabs. We sampled mixed leaf packs from the enclosures on days 11, 19, 28, 34, and 42. We found the leaves of one species (Koanophyllon pittieri) almost completely decomposed by day 28 in both treatments (crab versus no crab). The other two leaf species (Meliosma idiopoda, Quercus brenesii) composed the remaining leaf mass at the end of the experiment.
3. At 42 days, enclosures with crabs had faster rates of leaf breakdown than those without crabs (with crabs: k = −0.020; without crabs: k = −0.016; p = 0.034). This suggests that the magnitude of direct leaf breakdown by crabs, due to fragmentation, consumption, or manipulation of leaves, was greater than any indirect effects on leaf breakdown via crab consumption of other leaf-consuming species.
4. Macroinvertebrate composition based on taxa abundances or biomasses did not significantly differ between treatments (ANOSIM; p = 0.73 and p = 0.65, respectively). Shredder and detritivore abundances and biomasses increased significantly through time (ANOVA; p ≤ 0.001), but there was no evidence of an effect of crab presence (p > 0.2), nor were there significant interactions between crab presence and time (p > 0.3).
5. This is one of the first studies to quantify the effects of pseudothelphusid freshwater crabs on leaf breakdown rates. Our results suggest that these crabs can play a significant role in detrital processing in neotropical headwater streams. This study has also demonstrated that short-term enclosure experiments are useful in measuring in-stream effects of crab activity on leaf breakdown.

     

Green leaves enhance the growth and development of a stream macroinvertebrate shredder when senescent leaves are available

1. Freshly fallen green leaves and flowers of terrestrial plants enter temperate streams in spring and summer, when senescent leaf litter is often scarce. These resources appear to provide good supplementary food for macroinvertebrate shredders, but have some potential shortcomings as food or case material for caddisflies. 2. To compare suitability of green leaves or flowers and senescent leaves for the growth and development of stream shredders, we reared the caddisfly Lepidostoma complicatum in the laboratory with treatments that provided larvae with senescent (oak) and green (oak or maple) leaves separately, and also together, in case the combined use of both types of leaf may benefit the shredder. 3. Larvae supplied with green leaves alone grew at 65% of the rate of those provided with senescent leaves alone, due to their lower consumption rate. No individuals given green leaves alone developed into adults, whereas 70% of the individuals given senescent leaves alone did. Green leaves may inhibit larval consumption due to their high phenol content, or they may be unsuitable for case material because they are less tough than senescent leaves. 4. Larvae supplied with both senescent and green leaves (or flowers) had a higher growth rate and developed faster, than those given senescent leaves alone, whereas the proportions of successfully emerged individuals did not differ. Lepidostoma probably benefits from the higher nitrogen content of the green leaves when used together with senescent leaves. 5. These results suggest that green leaves (or flowers) cannot serve as an alternative food resource to senescent leaves, but that they can enhance the growth and development of a Lepidostoma stream shredder if senescent leaves are also available.     

Interactions between a stream caddisfly and the algae on its case: factors affecting algal quantity

1. Field experiments were conducted to examine interactions between larvae of the caddisfly Gumaga nigricula and the algae (primarily diatoms) located on their sand grain cases. Floating experimental enclosures were placed in sunlit pools of a California stream. Treatments applied to occupied cases, or to those from which caddisflies had been removed, included variations in grazer density, addition of nutrients, and modification of illumination using shade cloth. 2. Neither faecal-enriched agar in empty cases nor injections of faecal material into plugged cases affected case chlorophyll a concentration; injected nutrients (N and P), however, did increase chlorophyll a concentration. Faecal material may be nutrient poor, or nutrients may not be readily released. In addition, the relatively impermeable case wall may reduce the likelihood that nutrients from caddisflies reach the algae on the outer case surface. 3. Although a Gumaga larva may graze algae from its own case, grazing by conspecifics and the caddisfly Helicopsyche borealis causes a greater reduction in the chlorophyll a content of cases. 4. Chlorophyll a content of empty cases was decreased by experimental reduction of light. Because of shading associated with burrowing, chlorophyll a content of cases occupied by Gumaga was low and was unaffected by experimental light reduction. 5. Removal of Gumaga from its case results in rapid accrual of algae. Thus, the net effect of larval presence is to diminish the algal content of its case.     

Top-down and bottom-up influences on populations of a stream detritivore

1. Knowledge of the influence of predatory fish in detritus‐based stream food webs is poor. We tested whether larval abundance of the New Zealand leaf‐shredding caddisfly, Zelandopsyche ingens (family Oeconesidae), was affected by the presence of predatory brown trout, Salmo trutta and the abundance of their primary detrital resource (Nothofagus leaves). 2. The density of Z. ingens and the biomass of leaves were determined in seven fishless streams and four trout streams in the Cass region, central South Island, on four occasions spanning 5 years. 3. Physicochemical conditions were similar in trout and fishless streams, but ancova indicated that Z. ingens numbers were positively related to leaf biomass and that caddisfly numbers were significantly greater in fishless streams than trout streams for any given biomass of leaf. The cases of trout stream larvae were also heavier per unit length than those in fishless streams. 4. Our results provide evidence for both top‐down and bottom‐up influences on a detritus‐based stream food web. Although stream detritivores may benefit from a habitat that provides both food and a degree of protection from predators, top‐down effects of predators on detritivore population abundance were still important. Thus, detrital resource availability may determine maximum attainable population size, whereas predation is likely to reduce the population to a level below that.     

Factors affecting litter processing by Anisocentropus kirramus (Trichoptera: Calamoceratidae) from an Australian tropical rainforest stream

• 1 Larvae of the caddisfly Anisocentropus kirramus are common leaf shredders in rainforest streams in tropical Queensland. Laboratory experiments were undertaken to examine the effects of (a) differences in leaf species, age and degree of conditioning, (b) leaf preferences, (c) temperature and (d) larval stage on processing of leaf litter by A. kirramus.
• 2 Leaf species (five tested), age (fresh v senescent) and condition (conditioned in stream v unconditioned) each had a significant effect on the amount of leaf material that was processed. For most species, conditioned leaves were processed faster but for one species unconditioned leaves were processed faster. Senescent leaves were processed more rapidly than green leaves in three species but not in the other two. Given a choice of leaf types A. kirramus actively selected leaves that were processed faster; no preference was shown between two different ‘fast’ leaves.
• 3 Processing occurred at all temperatures tested (10, 18 and 25°C), with the processing rate increasing with temperature. Younger instar larvae processed leaves at a greater rate per unit body weight (up to 343% day^?1) than older instars. Final instar (5) larvae were capable of processing some tough leaves that younger instars did not process.
• 4 In rainforest streams, processing of leaves by A. kirramus takes place throughout the year. Its ability to process green leaves is important because of the high input of fresh green leaves into tropical streams, and because of the severe depletion of the supply of conditioned leaves and fine detritus after floods.

     

Biodiversity and ecosystem functioning in a species-poor guild: a test using tropical stream detritivores

The relationship between biodiversity and ecosystem functioning (B–EF) was investigated by examining top-down effects of aquatic detritivore diversity on the functional process of leaf-litter breakdown. This study was undertaken in tropical Hong Kong where the stream detritivore guild is depauperate and loss of one or a few species might have strong effects on processes. Effects of detritivore richness and composition were investigated by comparing feeding rates of three species of detritivores with their two- and three-species mixtures in laboratory trials. The detritivores were a caddisfly larva (Anisocentropus maculatus: Calamoceratidae), a snail (Brotia hainanensis: Pachychilidae) and a shrimp (Caridina cantonensis: Atyidae). Liquidambar formosana (Hamamelidaceae) litter was provided as food. All three detritivore species had positive non-additive effects on litter processing. Per capita and mass-specific feeding rates of each species were faster in mixtures than when they were alone, although the non-additive effects of shrimps and snails were larger than those attributable to caddisflies, and thus, litter processing was strongly influenced by the composition of detritivore mixtures. The compositional effect appears to be evidence of facilitation indicating a lack of functional redundancy amongst these detritivores, probably due to their evolutionary distinctness, implying that extinction consequences in this species-poor guild will depend on the identity of the species lost.     

Benthic macroinvertebrate response to sedimentation in a Typha angustifolia L. wetland

Fiberglass mesh enclosures (1 × 1 m²) in a Typha angustifoliaL. marsh were employed to examine the effects of clay additions on theresident macroinvertebrate communities. Total invertebrate density, insectdensity, and number of insect families decreased significantly by 33%,37%, and 17%, respectively, in enclosures receiving sediment. Morespecifically, incoming clay adversely affected densities of Coleoptera larvae,Diptera larvae, Megaloptera larvae, Odonata larvae, Pelecypoda, andGastropoda. Densities of specific families within the Diptera (larvae) andColeoptera were also affected; Dolichopodidae, Stratiomyidae,Hydrophilidae, Tabanidae, Dytiscidae adults, and Scirtidae larvae decreasedsignificantly in numbers in sedimented enclosures. In contrast, the effectof sedimentation on Carabidae (adults and larvae) and Dytiscidae larvaldensities varied significantly with time, whereby densities were higher in thesedimented treatment only for the initial two months of the study. Densities of predator-engulfer, collector-filterer, and scraper feeding groupswere reduced in sedimented plots by 28%, 44%, and 27%,respectively. Significant short- and long-term increases in turbidity andsuspended solids in enclosures treated with clay, as well as sedimentdeposition, were probably responsible for changes in the invertebratecommunities.     

Stream flow and predation effects on the spatial dynamics of benthic invertebrates

Field experiments were carried out to determine the influence of predation and prey movements on the accumulation of prey in enclosures. Experimental enclosures permitted exchange of prey with the benthos, but not of the large, predatory larvae of the caddisfly, Plectrocnemia conspersa (Curtis). Unseasonally heavy rainfalls during the experiment resulted in high flows and enabled us to examine the effects of a major, abiotic disturbance on invertebrate spatial dynamics. Prey colonization rates of cages without predators were determined in nine 24 h periods. Colonization rates increased exponentially with flow and were species-specific, depending on dispersal behaviour. Prey accumulation and predator impacts were measured in cages, with and without P. conspersa larvae, placed in the stream for 1, 2 or 3 weeks. Prey densities in cages increased with exposure time, but increases were not gradual and depended on flow regime. Flow was reduced within cages and they accumulated large numbers of invertebrates during high discharge. Analogous, naturally occurring refugia in the stream channel could be important for the recovery of lotic communities after major disturbances. Overall, prey densities were lowest in cages with predators. For fast colonizers, predation effects were detectable early in the experiment, but quickly obscured thereafter by continuous exchange of prey. For slow colonists, predation effects were detectable later, but persisted longer. Consumption rates for P. conspersa varied with prey density and flow regime. We suggest that the spatial dynamics of benthic invertebrates, especially as they are influenced by stochastic events, are important in understanding and detecting predation effects in stream communities.     

Effects of salmon carcasses on experimental stream ecosystems in Hokkaido,Japan

The effects of salmon carcasses on dissolved nutrients, epilithic production, leaf decomposition rates, and aquatic invertebrates were examined using 10-m-long artificial channels fed by an adjacent natural stream in Hokkaido, northern Japan. Bags containing maple leaf litter were placed in nine channels, each of which was assigned to one of three treatments with three replicates, for 6 weeks in fall 2002. The three treatments were: (1) salmon carcasses+invertebrates, (2) invertebrates only, and (3) control (no salmon carcasses or invertebrates added). Nutrient concentrations, biomass of epilithic algae (chlorophyll), leaf weight loss, abundance and biomass of invertebrates in the leaf packs were compared among the three treatments at 14, 27, and 40 days after the beginning of the experiment. The NH₄⁺ in stream water and chlorophyll concentrations of epilithic algae were higher in the salmon treatment than the other treatments, and the maple leaves decomposed faster in the salmon treatment than in the other treatments. Moreover, the N content of the leaves was highest and the C/N ratio was lowest in the salmon treatment, although not significantly so. The abundance and biomass of the dominant leaf-shredding invertebrate Goerodes satoi did not differ between the first two treatments. However, the stable N isotope ratio in G. satoi was nearly 3 higher in the salmon treatment, suggesting that around 20% of salmon-derived N was taken up by this shredder. Our results indicate that salmon carcasses affect stream ecosystems directly by enhancing primary production, indirectly by accelerating woody leaf decomposition, and finally by incorporating into the food web primary consumers that utilize fertilized woody leaves.     

Effects of atmospheric N deposition on coarse organic matter in a headwater stream

Input, storage, export potential, and system-level processing of coarse organic matter were investigated in the intermittent streams that drain the Bear Brook Watershed in Maine (BBWM). BBWM is a paired catchment study investigating ecosystem effects of atmospheric N and S deposition. We predicted that the increased N loading to the treatment catchment would elevate input of organic matter, result in higher levels of coarse organic matter biomass, and increase litter processing rates in the treatment stream relative to the reference stream. We found that the streams draining BBWM did not have statistically different coarse organic matter input, biomass, or processing rates and we found only modest differences in export potential. System-level processing rates for maple (Acer spp.) litter were similar to rates previously quantified using litterbag methods. However, system-level processing rates for American beech (Fagus grandifolia) litter were an order of magnitude faster than rates measured with litterbags. This difference was likely due to movements of these leaves from riffle/runs and pools into debris dams, rather than differences in measurements of leaf tissue processing rates between methods. Organic matter dynamics of the intermittent streams at BBWM were similar to other forested, headwater streams. Our results indicate that the long-term N manipulation experiment at BBWM has not altered input, storage or processing of coarse organic matter in the treatment stream. Physical characteristics of these stream ecosystems appear to regulate organic matter dynamics rather than differences in nutrient chemistry.     

Onshore–offshore variations of copepod community in northern South China Sea

Previous studies which have tested the feeding preferences of shredders for fungal species and the food quality of fungi used detritus uniformly colonized by a fungus, which is not the case for decaying leaves in streams. It is not known whether shredders in different development stages exhibit variations in feeding preference and larval performance. This study examined the feeding preferences and the growth of the third and the fifth instars of Pycnopsyche gentilis larvae using fungal-colonized patches and whole leaves, respectively, having different fungal species compositions (Alatospora acuminata, Anguillospora filiformis, Articulospora tetracladia, Tetrachaetum elegans, and all species combined). The aquatic hyphomycetes used were co-dominant on leaves in the stream inhabited by the caddisfly. During 14 d of feeding, the larvae of both instars did not show significant differences in feeding preferences for the patches growing on oak leaves, although the third instar larvae were slightly more selective than the fifth instar larvae. When fed with maple leaves for 18 d, larval growth rates, gross growth efficiencies, and survivorship were not significantly different among the fungal treatments. However, the larval growth of both instars fed with fungal-colonized leaves was always significantly greater than the growth of larvae fed with diets of uncolonized leaves. The third instar larvae grew faster than the fifth instar larvae, but the growth efficiencies of the two instars were similar. These results suggest that P. gentilis larvae exhibit less selectivity in their feeding than other caddisfly shredders that have been examined and that the dominant fungi colonizing leaves in their habitat are similar in palatability and food quality for this shredder. Handling editor: B. Oertli     

Do tadpoles affect leaf decomposition in neotropical streams?

1. Of the relatively few studies that have examined consequences of amphibian declines on stream ecosystems, virtually all have focused on changes in algae (or algal‐based food webs) and little is known about the potential effects of tadpoles on leaf decomposition. We compared leaf litter decomposition dynamics in two neotropical streams: one with an intact community of tadpoles (with frogs) and one where tadpoles were absent (frogless) as a result of a fungal pathogen that had driven amphibians locally extinct. The stream with tadpoles contained a diverse assemblage (23 species) of larval anurans, and we identified five species of glass frog (Centrolenidae) tadpoles that were patchily distributed but commonly associated with leaf detritus and organic sediments in pools. The latter reached total densities of 0–318 tadpoles m^?2. 2. We experimentally excluded tadpoles from single‐species leaf packs incubated over a 40‐day period in streams with and without frogs. We predicted that decomposition rates would be higher in control (allowing access of tadpoles) treatments in the study stream with frogs than in the frogless stream and, in the stream with frogs, in the control than in the tadpole exclusion treatment. 3. In the stream with frogs, Centrolene prosoblepon and Cochranella albomaculata tadpoles were patchily distributed in leaf packs (0.0–33.3 m^?2). In contrast to our predictions, leaf mass loss and temperature‐corrected leaf decomposition rates in control treatments were almost identical in our stream with frogs (41.01% AFDM lost, k_{degree day} = ?0.028 day^?1) and in the frogless stream (41.81% AFDM lost, k_{degree day} = ?0.027 day^?1) and between control and tadpole exclusion treatments within each stream. Similarly, there were no significant differences in leaf pack bacterial biomass, microbial respiration rates or macroinvertebrate abundance between treatments or streams. Invertebrate assemblages on leaf packs were similar between treatments (SIMI = 0.97) and streams (SIMI = 0.95) and were dominated by larval Chironomidae, Simuliidae (Diptera) and larval Anchytarsus spp. (Coleoptera). 4. In contrast to dramatic effects of grazing tadpoles on algal communities observed previously, tadpoles had no major effects on decomposition. While centrolenid tadpoles were common in the stream with frogs, their patchy distribution in both experimental and natural leaf packs suggests that their effects on detrital dynamics and microbes are probably more localised than those of grazing tadpoles on algae.     

Relative Importance of Bacteria and Fungi in a Tropical Headwater Stream: Leaf Decomposition and Invertebrate Feeding Preference

Bacteria and fungi provide critical links between leaf detritus and higher trophic levels in forested headwater food webs, but these links in tropical streams are not well understood. We compared the roles of bacteria and fungi in the leaf decomposition process and determining feeding preference for two species of freshwater shrimp found in the Luquillo Experimental Forest, Puerto Rico, using experimental microcosms. We first tested the effects of four treatments on decomposition rates for leaves from two common riparian species, Cecropia scheberiana (Moraceae) and Dacryodes excelsa (Burseraceae), in laboratory microcosms. Treatments were designed to alter the microbial community by minimizing the presence of bacteria or fungi. The fastest decay rate was the control treatment for D. excelsa where both bacteria and fungi were present (k = −0.0073 day⁻¹) compared to the next fastest rate of k = −0.0063 day⁻¹ for the bacterial-conditioned D. excelsa leaves. The fastest decay rate for C. scheberiana was also the control treatment (k = −0.0035 day⁻¹), while the next fastest rate was for fungal-conditioned leaves (k = −0.0029 day⁻¹). The nonadditive effect for leaf decomposition rates observed in the control treatments where both fungi and bacteria were present indicate that bacteria and fungi perform different functions in processing leaf litter. Additionally, leaf types differed in microbial colonization patterns. We next tested feeding preference for leaf type and microbe treatment in microcosms using two species of freshwater shrimp: Xiphocaris elongata, a shredder, and Atya lanipes, a scraper/filterer. To estimate feeding preferences of individual shrimp, we measured change in leaf surface area and the amount of particles generated during 5-day trials in 16 different two-choice combinations. X. elongata preferred D. excelsa over C. scheberiana, and leaves with microbial conditioning over leaves without conditioning. There was no clear preference for fungal-conditioned leaves over bacterial-conditioned leaves. This lack of preference for which microbes were responsible for the conditioning demonstrates the importance of both bacterial and fungal resources in these tropical stream food web studies.     

Effects of leaf type on the consumption rates of aquatic detritivores

Silver maple (Acer saccharinum) and cottonwood (Populus deltoides) leaves were incubated in cages excluding (controls) or containing (experimental) detritivores for periods of up to 123 days. Experimental cages contained either the cranefly larvae Tipula abdominalis, the amphipod Gammarus pseudolimnaeus or the caddisfly Pycnopsyche guttifer. Differences in daily consumption between leaf types and among species were compared.In control cages 20–23% of initial leaf weight was lost by leaching and 8–10% by microbial processes. T. abdominalis and P. guttifer consumed more silver maple than cottonwood during feeding intervals; no significant differences were observed for G. pseudolimnaeus. Invertebrate consumption of cottonwood leaves significantly increased with time; no significant differences in consumption of silver maple leaves with time were observed. Potential factors influencing the observed feeding pattern between leaf species are discussed.     

Nutrients stimulate leaf breakdown rates and detritivore biomass: bottom-up effects via heterotrophic pathways

Most nutrient enrichment studies in aquatic systems have focused on autotrophic food webs in systems where primary producers dominate the resource base. We tested the heterotrophic response to long-term nutrient enrichment in a forested, headwater stream. Our study design consisted of 2 years of pretreatment data in a reference and treatment stream and 2 years of continuous nitrogen (N) + phosphorus addition to the treatment stream. Studies were conducted with two leaf species that differed in initial C:N, Rhododendron maximum (rhododendron) and Acer rubrum (red maple). We determined the effects of nutrient addition on detrital resources (leaf breakdown rates, litter C:N and microbial activity) and tested whether nutrient enrichment affected macroinvertebrate consumers via increased biomass. Leaf breakdown rates were ca. 1.5 and 3× faster during the first and second years of enrichment, respectively, in the treatment stream for both leaf types. Microbial respiration rates of both leaf types were 3× higher with enrichment, and macroinvertebrate biomass associated with leaves increased ca. 2–3× with enrichment. The mass of N in macroinvertebrate biomass relative to leaves tended to increase with enrichment up to 6× for red maple and up to 44× for rhododendron leaves. Lower quality (higher C:N) rhododendron leaves exhibited greater changes in leaf nutrient content and macroinvertebrate response to nutrient enrichment than red maple leaves, suggesting a unique response by different leaf species to nutrient enrichment. Nutrient concentrations used in this study were moderate and equivalent to those in streams draining watersheds with altered land use. Thus, our results suggest that similarly moderate levels of enrichment may affect detrital resource quality and subsequently lead to altered energy and nutrient flow in detrital food webs. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Effect of carnivory on larvae and adults of a detritivorous caddisfly, Lepidostoma complicatum: a laboratory experiment

Lepidostomatid caddisfly larvae are typical detritivores, but they occasionally eat small dead animals. A laboratory feeding experiment was conducted with Lepidostoma complicatum (Kobayashi) larvae using two different feeding treatments: leaves or leaves and dead chironomids. L. complicatum larvae showed significantly higher growth rates and adult emergence weight and a significantly earlier emergence for the leaves and dead chironomids treatment than for the leaves alone treatment. However, the adult emergence rate was not different between the two feeding treatments. Thus, it is apparent that ingestion of dead chironomids by detritivorous L. complicatum larvae positively influences larval growth rates, adult emergence weight, and larval development time.     

Processing of leaf matter by lake-dwelling shredders at low oxygen concentrations

Organic sediments in freshwaters are regularly subject to low concentrations of oxygen. The ability of detritivores to sustain their feeding in such conditions should therefore be of importance for the decomposition process. In the present study, aquaria were used to determine processing rates of five lake-dwelling shredders at three different oxygen concentrations; normoxic (9 mg O₂ l^–1) and two levels of hypoxia (1 and 2 mg O₂ l^–1). Discs of alder leaves (Alnus glutinosa (L.)) were used as food. Four species of caddisfly larvae (Trichoptera Limnephilidae) and the isopod, Asellus aquaticus (L.) were compared in the experiments. Significant differences in processing rates per g animal biomass were found both at normoxia and 2 mg oxygen l^–1. At l mg O₂ l^–1 none of the invertebrates fed on leaf discs. The caddisfly larvae Halesus radiatus (Curtis), being one of the two most efficient shredders at normoxia, did not feed at 2 mg oxygen l^–1. The other species fed at rates 15–50 of that at normoxia. The least efficient shredder at normoxia, A. aquaticus was similar to two of the trichopterans at 2 mg O₂ l^–1. This study shows that the importance of specific shredder species may shift in case of hypoxia. Species-specific traits regarding oxygen sensitivity may also be influential for distribution patterns of shredder species both within and between lakes.     

Microbial Biomass,Respiration, and Decomposition of Hura crepitans L. (Euphorbiaceae) Leaves in a Tropical Stream1

The processing of leaves in temperate streams has been the subject of numerous studies but equivalent tropical ecosystems have received little attention. We investigated leaf breakdown of a tropical tree species (Hura crepitans, Euphorbiaceae), in a tropical stream using leaf bags (0.5 mm mesh) over a period of 24 days. We followed the loss of mass and the changes in adenosine triphosphate (ATP) concentrations and respiration rates associated with the decomposing leaves. The breakdown rate was fast (k=?0.0672/d, k_d=?0.0031/degree‐day), with 81 percent loss of the initial mass within 24 days. This high rate was probably related to the stable and high water temperature (22°C) favoring strong biological activity. Respiration rates increased until day 16 (1.1 mg O₂/h/g AFDM), but maximum ATP concentrations were attained at day 9 (725 nmol ATP/g AFDM) when leaf mass remaining was 52 percent. To determine the relative importance of fungi and bacteria during leaf decomposition, ATP concentrations, and respiration rates were determined in samples treated with antibiotics, after incubation in the stream. The results of the samples treated with the antifungal or the bacterial antibiotic suggest a higher contribution of the fungal community for total microbial biomass and a higher contribution of the bacterial community for microbial respiration rates, especially during the later stages of leaf decomposition. However, these results should be analyzed with caution since both antibacterial and antifungal agents did not totally eliminate microbial activity and biomass.     

Numerical and per capita responses to species loss: mechanisms maintaining ecosystem function in a community of stream insect detritivores

We experimentally reduced the diversity of detritivorous stream insects in field enclosures, and measured the effects on an ecosystem function, processing of leaf litter. Two dominant species were removed separately, the stonefly Pteronarcys californica and the caddisfly Lepidostoma unicolor . In principle, processing could be maintained after species loss in two ways: the remaining species could increase their rates of shredding (per capita response), or they could increase in abundance (numerical response). We imposed a numerical response in some treatments by experimentally increasing abundances of either all the remaining species or the other dominant species so that expected metabolic capacity of the assemblage returned to full-diversity levels. Numerical responses were generally effective in maintaining leaf breakdown when either Lepidostoma or Pteronarcys was removed, except that the treatment in which Lepidostoma was replaced by an equivalent metabolic capacity of all remaining species showed less leaf loss than the full-diversity treatment. Per capita responses by other species appeared effective in compensating for the removal of Pteronarcys (although there were other explanations) but were not effective in compensating for the removal of Lepidostoma . In summary, the consequences of reduced biodiversity varied with which species was lost and how the remainder responded. Thus there was no simple relationship between biodiversity and ecosystem functioning. However, when numerical or per capita compensation does occur, stability of function should rise with diversity in such "interactive" assemblages.