THE BENTHIC MACROINVERTEBRATE FAUNA OF A SOUTH AFRICAN MOUNTAIN STREAM AND ITS RESPONSE TO FIRE

Summary

The effects of a late-summer prescribed burn on the temperature and benthic macroinvertebrate fauna of a south-western Cape mountain stream were investigated over a period of 12 months. Temperature and discharge regimes appear well-defined and relatively predictable from year to year. As in other mediterranean-type ecosystems, seasonal changes in the structure of the invertebrate community and the relative abundance of different feeding groups appear to be associated primarily with changes in the physical environment. Distinctive summer and winter communities were identified, with chironomids dominating the fauna in summer and simuliids dominant in winter. Although the riparian vegetation was only slightly damaged by the fire, a heavy, aseasonal leaf-fall occurred shortly afterwards. The canopy remained sparse for approximately four months. Stream temperature in the post-burn year was not demonstrably affected by increased exposure to solar radiation, however, probably because the canopy remained open during the winter months. The fire appeared to have little effect on the invertebrate fauna. Apart from five rare elements of the biota, all species recorded in the pre-burn year were present in the post-burn year and in similar densities. It is concluded that the riparian vegetation is of major importance in maintaining the integrity of the stream environment.     

Tracing carbon sources in small urbanising streams: catchment‐scale stormwater drainage overwhelms the effects of reach‐scale riparian vegetation

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Particulate organic matter in a mountain stream in the south-western Cape,South Africa

The quality and quantity of allochthonous inputs and of benthic organic matter were investigated in a second-order, perennial mountain stream in the south-west Cape, South Africa, between April 1983 and January 1986. Although the endemic, riparian vegetation is sclerophyllous, low and evergreen, inputs of allochthonous detritus to the stream (434 to 500 g m^–2y^–1) were similar to those recorded for riparian communities worldwide, as were calorific values of these inputs (9548 to 10 032 KJ m^–2y^–1). Leaf fall of the riparian vegetation is seasonal, occurring in spring (November) as discharge decreases, resulting in retention of benthic organic matter (BOM) on the stream bed during summer and early autumn (maximum 224 g m^–2). Early winter rains (May) scoured the stream almost clean of benthic detritus (winter minimum 8 g m^–2). Therefore, BOM was predictably plentiful for about half of each year and predictably scarce for the other half. Coarse BOM (CBOM) and fine BOM (FBOM) constituted 46–64% of BOM standing stock, ultra-fine BOM (UBOM) 16–33% and leaf packs 13–24%. The mean annual calorific value of total BOM standing stock was 1709 KJ m^–2. Both standing stocks and total calorific values of BOM were lower than those reported for streams in other biogeographical regions. Values of C:N ratios decreased with decrease in BOM particle size (CBOM 27–100; FBOM 25–27; UBOM 13–19) with no seasonal trends. The stream is erosive with a poor ability to retain organic detritus. Its character appears to be dictated by abiotic factors, the most important of which is winter spates.     

Allochthonous input and retention in a small mountain stream,South Africa

Allochthonous input and benthic coarse particulate organic matter (CPOM) standing stocks were investigated in a first-order stream in South Africa between May 1984 and April 1985. Monthly falls into the stream of all litter types (total) ranged from 11 (September) to 79 g m^–2 (March). Total annual litter fall was 426 g dry weight, which corresponds to 1.2 g m^–2 d^–1. Flowers, fruits and seeds contributed 37 g m^–2, woody debris, 122 g m^–2, and leaves 267 g m^–2 to this total. Leaf fall from native trees, which accounted for approximately 57% of total litter input (244 g m^–2 a^–1), was significantly higher in summer than in winter. The summer peak in leaf fall recorded is far smaller and more protracted than the autumnal peak recorded for many Northern Hemisphere streams.Monthly total standing stocks of CPOM ranged from 14 g dry weight m^–2 in January to 69 g m^–2 in August, and a mean total CPOM standing stock of 41 g m^–2 mth^–1 was estimated. This comprised 18 g m^–2 mth^–1 soft litter, and 23 g m^–2 mth^–1 hard litter. CPOM standing stocks showed no seasonal trends, and with the exception of two species, standing stocks of endemic leaf species reflected their contributions to the total litter fall. Contrary to earlier reports for streams in the Fynbos Biome, Window Stream has CPOM standing stocks well within the ranges reported for low-order streams worldwide.     

Urban catchment hydrology overwhelms reach scale effects of riparian vegetation on organic matter dynamics

1. Urbanisation severely affects stream hydrology, biotic integrity and water quality, but relatively little is known about effects on organic matter dynamics. Coarse particulate organic matter (CPOM) is a source of energy and nutrients in aquatic systems, and its availability has implications for ecosystem productivity and aquatic communities. In undisturbed environments, allochthonous inputs from riparian zones provide critical energy subsidies, but the extent to which this occurs in urbanised streams is poorly understood. 2. We investigated CPOM inputs, standing stocks, retention rates and retention mechanisms in urban and peri‐urban streams in Melbourne, Australia. Six streams were chosen along a gradient of catchment urbanisation, with the presence of reach scale riparian canopy cover as a second factor. CPOM retention was assessed at baseflow via replicate releases of marked Eucalyptus leaves where the retention distance and mechanism were recorded. CPOM and small wood (>1 cm diameter) storage were measured via cores and direct counts, respectively, while lateral and horizontal CPOM inputs were assessed using riparian litter traps. Stream discharge, velocity, depth and width were also measured. 3. CPOM inputs were not correlated with urbanisation, but were significantly higher in ‘closed’ canopy reaches. Urbanisation and riparian cover altered CPOM retention mechanisms, but not retention distances. Urban streams showed greater retention by rocks; while in less urban streams, retention by small wood was considerably higher. CPOM and small wood storage were significantly lower in more urban streams, but we found only a weak effect of riparian cover. 4. These findings suggest that while riparian vegetation increases CPOM inputs and has modest/weak effects on storage, catchment scale urbanisation decreases organic matter availability. Using an organic matter budget approach, it appears likely that the increased frequency and magnitude of high flows associated with catchment urbanisation exerts an overriding influence on organic matter availability. 5. We conclude that to maintain both organic matter inputs and storage, the restoration and protection of streams in urban or rapidly urbanising environments relies on the management of both riparian vegetation and catchment hydrology.     

Benthic organic matter and detritivorous macroinvertebrates in two intermittent streams in south-eastern Australia

Temporal changes in coarse (> 1 mm), fine (< 1 mm, > 250 m), and woody benthic organic matter (BOM), and densities of detritivores in pools and riffles were monitored at three sites on two intermittent streams (Werribee and Lerderderg Rivers) in Victoria, Australia during a drought year followed by a wetter year. Standing stocks of BOM peaked in both habitats during summer when discharge ceased and eucalypt leaf fall was greatest. During high winter and spring discharges, concentrations of BOM were low. Floods did not always scour BOM from the pools and riffles; after floods in October, standing stocks rose when BOM was imported from upstream or mobilized from the riparian zone. Densities of benthic detritivores, collector-gatherers, and shredders also varied seasonally, usually peaking in summer. Correlations between detritivore feeding group densities and amounts of putative food resource were habitat-specific. There were few significant correlations in depositional habitats, even after incorporating lag-times of two and four weeks into the analysis. However, detritivore densities in riffles, especially in the Lerderderg River, were strongly positively correlated with the amounts of BOM. Possibly, physico-chemical conditions in riffles are more conducive to litter conditioning and invertebrate colonization and breakdown of leaf material than are those in pools.     

Small reductions in forest cover weaken terrestrial-aquatic linkages in headwater streams

1. We assessed the impacts of deforestation on the energy base of headwater food webs in seven headwater streams in the Upper Chattahoochee basin, GA, U.S.A where percentage forest in catchments ranged from 82 to 96%. We measured terrestrial organic matter standing crop and determined consumer (crayfish and insectivorous fish) dependence on terrestrial versus aquatic energy sources via gut content and stable isotope analyses. 2. Standing crop of coarse particulate organic matter (CPOM) declined with deforestation at large scales (i.e. catchment deforestation and riparian deforestation at the entire stream network scale). Terrestrial plant matter, the dominant component of crayfish guts, declined in crayfish guts with reductions in CPOM standing crop and with deforestation. 3. Crayfish and insectivorous fish δ¹³C showed enrichment trends with deforestation, indicating isotopic divergence from CPOM, the most ¹³C‐depleted basal resource, with reductions in catchment and riparian forest cover. Crayfish δ¹³C also exhibited enrichment with decreased instream CPOM standing crop. 4. A concentration‐dependent mixing model was used to calculate the relative dependence of crayfish and fish on terrestrial versus aquatic basal resources. Results suggested that both allochthonous CPOM and autochthonous production were important basal resources. Consumer dependence on CPOM decreased with reductions in canopy cover. 5. Our data suggest the importance of forest cover to headwater food webs at multiple scales, and that relatively low levels of riparian deforestation along headwater streams can lead to reductions in stream food web dependence on terrestrial subsidies.     

Revealing the pathways by which agricultural land‐use affects stream fish communities in South Brazilian grasslands

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Seasonality and composition of benthic coarse particulate organic matter in two coastal tropical streams with different land uses

This work studies benthic CPOM in two streams of Ecuador: the Atacames stream, located in a developed watershed, and the Súa stream, located in a rural watershed and used as a reference. It is tested whether the amount, composition and timing of benthic CPOM will differ between them as a function of watershed and riparian land uses. Benthic CPOM was collected at five study sites on each stream with a Surber net and classified into four categories: leaves, twigs and bark, flowers and fruits and debris. Leaves were further identified to genus or species. There were no significant differences in the amount, composition and timing of benthic CPOM between the streams. CPOM storage showed strong seasonality linked to seasonal rainfall and a weak relation with land uses, channel width and stream order. Diversity of the benthic CPOM was high and 30 species contributed to the benthic leaf pool. Presence or absence of Ficus species with heavy leaves that are easily retained in the streambed explained the spatial distribution of benthic CPOM, so spatial differences in the composition of the riparian vegetation in these tropical streams seem to be more important to explain CPOM distribution than in their temperate counterparts.

     

Temporal and Spatial Patterns in Inputs and Stock of Organic Matter in Savannah Streams of Central Brazil

Coarse particulate organic matter (CPOM) inputs from riparian vegetation into streams and CPOM benthic stock vary naturally in space and time, but most studies in the tropical savannah (Cerrado) have been done over a small temporal scale (<1 year), which does not allow for the determination of inter-annual patterns. We found that CPOM collected over two years differed temporally and spatially, whereas there was no significant variation between years for the benthic stock, which indicates high stability in the energy balance of streams. The largest monthly inputs occurred between August and October, at the end of the dry season and the onset of the rainy season, which was partially explained by precipitation. Other factors such as photoperiod, which was not studied, could also have important roles in this pattern. Spatial differences in CPOM between streams were attributed to topography and channel morphology. The plant density was lowest in the stream with a more irregular topography and a deeper channel, which results in drier riparian soil. The benthic stock was highest in the stream with a flat channel, where the lower water speed facilitates the accumulation of CPOM in the stream bed. Inter-annual differences in CPOM were attributed more to the differences in the beginning of the dry and wet periods between years than to the average values of precipitation. Longer-term studies are needed to clarify this temporal pattern.     

Seasonal Variability of Particulate Organic Matter in a Mountain Stream in Central Spain

The aim of this paper was to study the influence of environmental characteristics of the Mediterranean climate on seasonal variability of particulate organic matter abundance in a mountain stream. Coarse and fine fractions of both suspended and benthic particulate organic matter were determined on 14 occasions between February 1998 and November 1999 in a second‐order Mediterranean stream in Central Spain (Arroyo Mediano). Temporal variability of suspended organic matter followed a seasonal pattern, attributed to litter‐fall inputs, instream processing, and the hydrological regime. Suspended organic matter (SOM) and its seasonal variability fall well within the range reported for streams in temperate non‐Mediterranean deciduous forest. However, we found no seasonal trend in benthic organic matter (BOM) storage, and it seems that the amount of BOM remained fairly constant throughout the year. Reach retention (evaluated as the ratio between BOM and SOM per m²) was higher in summer during reduced stream flow, mainly due to coarse particulate organic matter storage. These observations do not differ from those reported for other headwater streams in temperate forested biomes, from which we conclude that there was no evidence of a Mediterranean influence on particulate organic matter dynamics in the Mediano stream, nor probably in other headwater Mediterranean streams. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Lateral input of particulate organic matter from bank slopes surpasses direct litter fall in the uppermost reaches of a headwater stream in Hokkaido, Japan

In forested streams, surrounding riparian forests provide essential supplies of organic matter to aquatic ecosystems. We focused on two pathways of particulate organic matter inputs: direct input from upper riparian forests and indirect lateral input from bank slopes, for which there are limited quantitative data. We investigated the inputs of coarse particulate organic matter (CPOM) and carbon and nitrogen in the CPOM into the uppermost reaches of a headwater stream with steep bank slopes in Hokkaido, Japan. CPOM collected by litter traps was divided into categories (e.g., leaves, twigs) and weighed. Monthly nitrogen and carbon inputs were also estimated. The annual direct input of CPOM (ash-free dry mass) was 472 g m⁻², a common value for temperate riparian forests. The annual lateral CPOM input was 353 g m⁻¹ and 941 g m⁻² when they were converted to area base. This value surpassed the direct input. Organic matter that we could not separate from inorganic sediments contributed to the total lateral input from the bank slopes (124 g m⁻¹); this organic matter contained relatively high amounts of nitrogen and carbon. At uppermost stream reaches, the bank slope would be a key factor to understanding the carbon and nitrogen pathways from the surrounding terrestrial ecosystem to the aquatic ecosystem.     

Impact of a eucalyptus (Eucalyptus globulus Labill.) plantation on the nutrient content and dynamics of coarse particulate organic matter (CPOM) in a small stream

Litterfall inputs, benthic storage and the transport of coarse particulate organic matter (CPOM) were studied in two headwater streams, one flowing through a mixed deciduous forest and one through a plantation of Eucalyptus globulus. Vertical and lateral traps, transported CPOM and benthic CPOM were sampled monthly to biweekly and sorted into four categories: leaves, twigs and bark, fruits and flowers and debris. The litterfall inputs were about 20% lower at the eucalyptus site but this reduction was unevenly distributed among the litter categories. The reduction of the nitrogen and phosphorus inputs was larger (50%) than that of CPOM because of the low nutrient concentration of the CPOM at the eucalyptus site. Transported CPOM was also lower at the eucalyptus site. Although total CPOM inputs to the stream were reduced in the eucalyptus plantation, benthic storage of CPOM was 50% higher due to (1) high inputs of CPOM and low discharge during summer, (2) more twig and bark inputs, (3) eucalyptus leaves being retained more efficiently in the stream than deciduous leaves (4) a lower discharge, which may in part be attributable to eucalyptus-induced changes in the hydrological cycle. Increased retention balanced lower nitrogen and phosphorus content of CPOM, so benthic storage of nitrogen and phosphorus was similar at both sites. This work demonstrates that the timing, quality and quantity of inputs and benthic storage of CPOM in streams changes substantially because of the substitution of natural deciduous forest with eucalyptus plantation. Maintenance of buffer strips of natural vegetation may be the best way to protect ecological functioning of small, forested streams.     

Effects of livestock grazing on benthic invertebrates from a native grassland ecosystem

SUMMARY 1. The effects of cattle grazing on stream bank stability, biomass of riparian vegetation, instream vegetation cover, biomass of coarse particulate organic matter (CPOM) and epilithon and benthic invertebrate community structure were investigated over a 2‐year period using: (i) enclosures containing different cattle grazing treatments and (ii) by comparing streams with different grazing intensities in the Cypress Hills Provincial Park, Alberta, Canada. 2. Livestock enclosure experiments comprised four treatments of: (1) early season cattle grazing (June–August), (2) late season cattle grazing (August–September), (3) all season cattle grazing (June–September) and (4) cattle‐absent controls. All four treatments were replicated in two streams while two treatments (i.e. cattle‐absent controls, early season cattle grazing) were established in a third stream. 3. Bank stability, estimated visually based on sediment inputs to the stream channel, increased significantly in cattle‐absent treatments compared with cattle‐present enclosures over the 2‐year study period. 4. Epilithic chlorophyll a was significantly affected by time, but neither cattle nor the interaction of time and treatment were significant. 5. At the end of the experiment, total invertebrate biomass in the late and all‐season treatment exceeded that in the early and cattle‐absent treatments. However, excluding cattle from the streams, at any of the different treatments, had little clear impact on the total benthic invertebrate abundance or the abundance of the predominant functional feeding groups over the 2‐year study period. 6. In contrast, comparisons of benthic assemblages from streams with different grazing intensities showed that the non‐grazed reach of Storm Creek contained significantly higher biomass of CPOM and shredders compared with the cattle‐absent enclosures in Battle, Graburn and Nine Mile creeks. Redundancy analysis showed that benthic communities from all enclosures and Storm Creek in summer and autumn 2000 were affected primarily by CPOM biomass, pH, nitrate, turbidity and benthic chlorophyll a. Construction of a 99% probability ellipse from enclosure sites showed that invertebrate communities from livestock enclosures differed from that in the non‐grazed Storm Creek. 7. Results from stream‐scale comparisons indicate that current livestock grazing practices in the Cypress Hills significantly impact riparian zones, stream channels and benthic invertebrate community structure and that alternative practices, such as rotational grazing, need to be developed.     

Changes in stream benthic organic matter following watershed disturbance

Benthic organic matter was collected quarterly from streams draining a 9-yr-old clearcut, an 18-yr-old "old-field", a 25-yr-old successional forest, and two reference watersheds at Coweeta Hydrologic Laboratory in the Appalachian Mountains of North Carolina. USA. Samples were separated into large benthic organic matter (LBOM >1 mm) and fine benthic organic matter (FBOM <1 mm). An additional survey of large (>5 em diam.) and small (1–5 cm diam.) wood was conducted. Standing stocks of LBOM ranged from 124 to 235 g AFDM m⁻² (ash tree dry mass) and were significantly higher in streams draining reference watersheds and the successional forest than in either the recent clearcut or old-field. Reference sites exhibited LBOM peaks in late autumn and spring. No seasonal patterns were observed in disturbed streams. Standing stocks of FBOM averaged 113 to 387 g AFDM m⁻², and the stream draining the successional forest had significantly higher FBOM levels than the other sites. In reference streams, FBOM abundance peaked in spring. In disturbed streams, FBOM standing stocks were highest in summer or late autumn. Standing stocks of large wood ranged from 0 to 3956 g AFDM m⁻² and were significantly higher in the reference streams than in streams draining the old-field or successional forest. Small wood averaged 11 to 342 g AFDM m⁻² and was significantly lower in the stream draining the old-field than at the other sites. Comparisons of organic matter inputs with standing stocks indicated that disturbed streams at Coweeta receive less material and process it faster than reference streams. Disturbed streams also appear to be less retentive than reference streams and exhibit a gradual decline in FBOM during the winter when large, long-duration storms combined with low particle generation rates deplete accumulated FBOM.     

Storage and decomposition of particulate organic matter along the longitudinal gradient of an agriculturally-impacted stream

An agriculturally-impacted stream in northern Idaho was examined over a two-year period to determine seasonal and longitudinal patterns of the storage and decomposition of particulate organic matter. Biomass of benthic organic matter (BOM) was considerably less than values reported in the literature for comparable, undisturbed streams. Coarse, fine, and total benthic particulate organic matter were not correlated with parameters pertaining to stream size (e.g., stream order), but were correlated with sample site and amount of litterfall. The association of BOM with site and litterfall suggests that storage of particulate organic matter is a function of local characteristics rather than stream size. Low biomass of stored organic matter is a response to the low input of terrestrially-derived organic matter resulting from removal of climax vegetation.Leaf packs of alder, Alnus sp., were placed in the stream seasonally for 30 and 60 d. While there were significant differences for months, there was no significant difference among sites for leaf packs exposed for 30 d. Significant differences were observed among both sites and months for leaf packs exposed for 60 d; however, differences among sites accounted for only 5% of the variance. The absence of differences in decomposition of organic matter along the gradient of Lapwai Creek, despite heterogeneity of the drainage basin and availability of organic matter, may be in response to the overall low biomass of stored benthic organic matter. This study demonstrates that agricultural activity can substantially influence instream heterotrophic processes through reduced availability of organic matter and can shape community structure and ecosystem dynamics of streams flowing through agricultural drainage basins.     

Benthic organic matter dynamics in Texas prairie streams

Concentrations of benthic particulate organic matter (POM) in six Texas prairie streams (2nd–4th order, intermittent and perennial) were monitored over a 20 month period to determine temporal and spatial dynamics. Benthic POM mass was highly variable, having coefficients of variation (CV) in excess of 300%. Benthic POM mass in all streams was similar with the exception of the 4th order intermittent stream which had significantly higher concentrations. Benthic POM at all sites was dominated by coarse POM (CPOM), followed by fine POM (FPOM), ultrafine POM (UPOM), and medium POM (MPOM). The dominance by CPOM is especially noteworthy in the 4th order intermittent stream where it accounted for 83% of the annual POM mass. Seasonally, benthic POM was highest in summer and lowest in the fall.     

The role of aquatic fungi in transformations of organic matter mediated by nutrients

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A novel method reveals how channel retentiveness and stocks of detritus (CPOM) vary among streams differing in bed roughness

1. Coarse particulate organic matter (CPOM) is a fundamental resource in freshwater streams, providing food, shelter and habitat for diverse invertebrate taxa and playing a key role in metabolism in low-order streams. Benthic CPOM stocks are determined by rates of supply and breakdown of detritus and by channel retentiveness (i.e. the capacity for the channel to trap and retain CPOM). We focussed on factors affecting the retentiveness of channels, which theoretically differs among streams with different sediment sizes and concomitant channel morphology.
2. We developed a new, rapid method to measure retentiveness using line-intercept surveys along transects. With this rapid approach, we surveyed 32 sites from three types of streams (smooth sandy channels, n = 10; gravel channels of intermediate roughness, n = 12; rough cobble channels, n = 10) in Victoria, south-eastern Australia, and tested the simple hypotheses that: (1) retentiveness increases in channels with increasing channel roughness (i.e. sandy versus gravel versus cobble-bed streams); (2) different types of channel features (e.g. log jams, cobbles, depositional areas) differ in the efficiency with which they retain CPOM. The line-intercept survey method was readily adapted to measure retentiveness as m of retentive structure per m of transect (i.e. the Linear Coverage Index) and trapping efficiency as m of CPOM per m of retentive element, for 10 different types of retentive elements.
3. Unexpectedly, the retentiveness of channels did not increase with channel roughness. This occurred because channels with different roughness were dominated by different types of retentive structure. Retentive structure in cobble sites was dominated by cobbles themselves, which were highly retentive in other studies but poorly retentive in our system. Gravel and sand sites had more log jams and depositional areas, such as pools and backwaters, and these features were more effective at trapping CPOM. Thus, retention of CPOM was highest in gravel and sand sites.
4. Our method provides a new tool for investigators testing hypotheses about CPOM retention in streams. The method is rapid, requires a minimum of equipment and personnel, and may be applied in any wadeable stream. Retentiveness is calculated in intuitive units that are directly comparable among sites and may have utility as variables in models of CPOM dynamics. We hope this method will open up new avenues for research that may shed light on how CPOM stocks vary among streams, with implications for diversity of aquatic fauna and ecosystem functions such as decomposition.

     

Effect of a leaf-shredding invertebrate on organic matter dynamics and phosphorus spiralling in heterotrophic laboratory streams

Summary The effect of invertebrate shredders on organic matter dynamics and phosphorus spiralling was studied over a 30-week period in laboratory streams. The streams were fed by groundwater, layered with cobble and gravel from a natural stream, covered with opaque material to eliminate algal growth, and initially contained 195 g/m² of autumn-shed leaves. Four weeks after leaf addition, leaf-shredding snails (Goniobasis clavaeformis) were added to each of three streams in densities of 75, 220, and 800/m². A fourth stream contained no snails and served as a control.Presence of snails increased the loss rates of coarse particulate organic matter (CPOM) and total organic matter (TOM), primarily by increasing leaf fragmentation and seston export. Although snail feeding increased specific metabolism of microbes associated with CPOM and cobble surfaces, it was not enough to compensate for reduction in bacterial cell numbers per unit surface area and in stream TOM. Consequently mineralization of detritus and whole stream phosphorus utilization rate were maximum in the stream with no snails and decreased with increasing snail density. From previous simulations of a stream model based on the nutrient spiralling concept, we predicted that there should be an intermediate shredder density which would minimize phosphorus spiralling length (maximize phosphorus utilization) in a natural stream nearby. Our current results conflict with the model-based predictions primarily because the increase in microbial metabolism was less important than reduction in bacterial cell numbers and total benthic organic matter resulting from snail feeding. Although our results indicate macroinvertebrate shredders reduce phosphorus utilization in headwater streams, shreders may increase nutrient utilization downstream where riparian inputs are lower, thus linking low- and high-order streams.Research supported by the National Science Foundation's Ecosystem Studies Program under Interagency Agreement No. BSR-8103181, A02 with the U.S. Department of Energy, under Contract No. DE-AC05-840R21400 with Martin Marietta Energy Systems, Inc.Publication No. 2394, Environmental Sciences Division, ORNL