Shifts in leaf litter breakdown along a forest–pasture–urban gradient in Andean streams

Tropical montane ecosystems of the Andes are critically threatened by a rapid land‐use change which can potentially affect stream variables, aquatic communities, and ecosystem processes such as leaf litter breakdown. However, these effects have not been sufficiently investigated in the Andean region and at high altitude locations in general. Here, we studied the influence of land use (forest–pasture–urban) on stream physico‐chemical variables (e.g., water temperature, nutrient concentration, and pH), aquatic communities (macroinvertebrates and aquatic fungi) and leaf litter breakdown rates in Andean streams (southern Ecuador), and how variation in those stream physico‐chemical variables affect macroinvertebrates and fungi related to leaf litter breakdown. We found that pH, water temperature, and nutrient concentration increased along the land‐use gradient. Macroinvertebrate communities were significantly different between land uses. Shredder richness and abundance were lower in pasture than forest sites and totally absent in urban sites, and fungal richness and biomass were higher in forest sites than in pasture and urban sites. Leaf litter breakdown rates became slower as riparian land use changed from natural to anthropogenically disturbed conditions and were largely determined by pH, water temperature, phosphate concentration, fungal activity, and single species of leaf‐shredding invertebrates. Our findings provide evidence that leaf litter breakdown in Andean streams is sensitive to riparian land‐use change, with urban streams being the most affected. In addition, this study highlights the role of fungal biomass and shredder species (Phylloicus; Trichoptera and Anchytarsus; Coleoptera) on leaf litter breakdown in Andean streams and the contribution of aquatic fungi in supporting this ecosystem process when shredders are absent or present low abundance in streams affected by urbanization. Finally, we summarize important implications in terms of managing of native vegetation and riparian buffers to promote ecological integrity and functioning of tropical Andean stream ecosystems.     

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

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

     

The legacy of forest logging on organic matter inputs and storage in tropical streams

Riparian forests play an important role in stream ecosystems, as they support biodiversity, reduce water erosion, and provide litter that fuels aquatic biota. However, they are affected by great array of anthropogenic threats (e.g., fire, logging, and organic pollution), which alter species composition and their physical structure. Although forest recovery after disturbance such as logging can take decades, the legacy of forest clear-cut logging on key processes in tropical riparian ecosystems is mostly unknown. Here, we investigated how litter inputs (leaves, twigs, and reproductive parts) and storage, key processes for carbon and nutrient recycling and for forest and stream biota, are influenced by riparian vegetation undergoing succession (after 28 years from logging) through the comparison of reference and logged forest sites in the Cerrado biome. Litterfall was overall similar between forest types, but litterfall of twigs was twofold higher at logged than reference sites. Similarly, litter inputs from the bank to the stream (i.e., lateral inputs) and streambed storage were 50–60% higher at logged than reference sites. The higher litterfall observed in logged forests could be related to higher proportion of tree species that are characteristic of primary and secondary successional stages, including fast-growing and liana species, which often are more productive and common in anthropogenic areas. Our results showed that the legacy impact of clear-cut logging, even if residual woody vegetation is maintained in riparian buffers, can shift the type, quantity, and seasonality of litter subsidies to tropical streams. This knowledge should be considered within the context of management and conservation of communities and ecosystem processes in the forest-stream interfaces.     

Influences of logging history and riparian forest characteristics on macroinvertebrates and brook trout (Salvelinus fontinalis) in headwater streams (New Hampshire, U.S.A.)

1. Logging can strongly affect stream macroinvertebrate communities, but the direction and magnitude of these effects and their implications for trout abundance are frequently region‐specific and difficult to predict. 2. In first‐order streams in northern New England (U.S.A.) representing a chronosequence of logging history (<2 to >80 years since logging), we measured riparian forest conditions, stream macroinvertebrate community characteristics and brook trout (Salvelinus fontinalis) abundance. Principal component analysis was used to collapse forest data into two independent variables representing variation in logging history, riparian forest structure and canopy cover. We used these data to test whether logging history and associated forest conditions were significant predictors of macroinvertebrate abundance and functional feeding group composition, and whether brook trout abundance was related to logging‐associated variation in invertebrate communities. 3. Catchments with high PC1 scores (recently logged, high‐density stands with low mean tree diameter) and low PC2 scores (low canopy cover) had significantly higher total macroinvertebrate abundance, particularly with respect to chironomid larvae (low PC2 scores) and invertebrates in the grazer functional feeding group (high PC1 scores). In contrast, proportional representation of macroinvertebrates in the shredder functional feeding group increased with time since logging and canopy cover (high PC2 scores). Brook trout density and biomass was significantly greater in young, recently logged stands (high PC1 scores) and was positively related to overall macroinvertebrate abundance. In addition, three variables – trout density, invertebrate abundance and shredder abundance – successfully discriminated between streams that were less‐impacted versus more‐impacted by forestry. 4. These results indicate that timber harvest in northern New England headwater streams may shift shredder‐dominated macroinvertebrate communities supporting low trout abundance to a grazer/chironomid‐dominated macroinvertebrate community supporting higher trout abundance. However, while local effects on brook trout abundance may be positive, these benefits may be outweighed by negative effects of brook trout on co‐occurring species, as well as impairment of habitat quality downstream. Research testing the generality of these patterns will improve understanding of how aquatic ecosystems respond to anthropogenic and natural trajectories of forest change.     

The impacts of habitat disturbance on adult and larval dragonflies (Odonata) in rainforest streams in Sabah,Malaysian Borneo

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Ecosystem Linkages between Southern Appalachian Headwater Streams and Their Banks: Leaf Litter Breakdown and Invertebrate Assemblages

We examined red maple (Acer rubrum L.) leaf litter breakdown in streams and riparian zones at two sites in the southern Appalachian Mountains to understand how differences in abiotic and biotic factors influence leaf breakdown rates. Litterbags were placed in three riparian habitats differing in litter layer moisture: stream > bank > upland. Invertebrates colonizing litterbags at one site were also examined to determine how variations in community and functional structure affect breakdown rates. Leaves broke down fastest in streams and slowest in upland habitats, whereas bank habitats were intermediate and characterized by high variability. Faster leaf breakdown rates in streams appeared to be a function of greater moisture availability, a more stable thermal regime, and a higher biomass of leaf-shredding invertebrates, especially the stonefly Tallaperla. In addition, patterns of leaf breakdown and invertebrate community structure provided evidence for a stronger than expected ecological connection between the stream and the bank. Overall, detritus processing within this narrow riparian ecosystem varied considerably depending on the availability of moisture. Results from this study show that stream channel–floodplain interactions in riparian ecosystems of steep forested mountains are analogous to ones in larger downstream or low-gradient systems. Riparian zones throughout a river network display a remarkable heterogeneity in their ability to process organic matter, which is ultimately driven by changes in hydrological conditions. Received 6 March 2001; accepted 3 July 2001.     

Leaf‐litter breakdown in pasture and deciduous woodland streams: a comparison among three European regions

1. Human land‐use has altered catchments on a large scale in most parts of the world, with one of the most profound changes relevant for streams and rivers being the widespread clearance of woody riparian vegetation to make way for livestock grazing pasture. Increasingly, environmental legislation, such as the EU Water Framework Directive (EU WFD), calls for bioassessment tools that can detect such anthropogenic impacts on ecosystem functioning. 2. We conducted a large‐scale field experiment in 30 European streams to quantify leaf‐litter breakdown, a key ecosystem process, in streams whose riparian zones and catchments had been cleared for pasture compared with those in native deciduous woodland. The study encompassed a west–east gradient, from Ireland to Switzerland to Romania, with each of the three countries representing a distinct region. We used coarse‐mesh and fine‐mesh litter bags (10 and 0.5 mm, respectively) to assess total, microbial and, by difference, macroinvertebrate‐mediated breakdown. 3. Overall, total breakdown rates did not differ between land‐use categories, but in some regions macroinvertebrate‐mediated breakdown was higher in deciduous woodland streams, whereas microbial breakdown was higher in pasture streams. This result suggests that overall ecosystem functioning is maintained by compensatory increases in microbial activity in pasture streams. 4. We suggest that simple coefficients of breakdown rates on their own often might not be powerful enough as a bioassessment tool for detecting differences related to land‐use such as riparian vegetation removal. However, shifts in the relative contributions to breakdown by microbial decomposers versus invertebrate detritivores, as revealed by the ratios of their associated breakdown rate coefficients, showed clear responses to land‐use.     

Invertebrate community structure and ecosystem functioning in European conifer plantation streams

1. Terrestrial leaf‐litter is the dominant energy input to many headwater streams and consequently the nature of the riparian vegetation can have profound effects on in‐stream processes. The impact of conifer plantations on community structure and ecosystem functioning (litter breakdown) was investigated in field experiments in three countries (Britain, Ireland, Poland), each representing a distinct European ecoregion. Twenty‐six streams were used in the trial: half were bordered with broadleaved and the other half with conifer riparian vegetation. 2. In a leaf breakdown study using litter bags, two leaf types (oak and alder) were used to assess the impact of resource quality and two mesh sizes (10 and 0.5 mm aperture) were used to gauge the relative importance of invertebrate detritivores and microbial decomposers respectively. Comparisons were made between vegetation types and among regions; pH varied among individual streams but, unlike many previous studies, it was not confounded with vegetation type, enabling us to isolate the effect of vegetation more effectively. 3. Overall, riparian vegetation type did not affect breakdown rates but strong regional differences were observed. There was also a significant interaction between these two variables, but this disappeared after fitting pH as a covariable, demonstrating its importance in determining breakdown rates and raising the possibility that in previous studies the impacts of conifer plantations might have been confounded with pH. 4. Shredder species composition differed between vegetation types. Small stoneflies were most strongly associated with conifer streams; broadleaved streams generally had a higher proportion of larger taxa, such as limnephilid caddisflies and gammarid shrimps, although the latter were excluded from sites with low pH. However, breakdown rates were maintained irrespective of shredder community composition, suggesting a high degree of functional redundancy in these communities. Similar processing rates were observed between streams with high numbers of nemourids and those with only a few limnephilids or gammarids, suggesting that density compensation among consumers might stabilise process rates. 5. Our results suggest that leaf‐litter breakdown can be an effective proxy for assessing stream ecosystem functioning, as rates differed significantly across spatial scales, from between streams to across regions and responded to an environmental gradient (pH). The litter bag technique can also complement traditional assessment methods by providing valuable information on the composition of consumer guilds, thereby providing an important link between structure and function that is needed to help inform management practices.     

Effects of Historical Selective Logging on Anuran Communities in a Wet Evergreen Forest,South India

Vast areas of tropical evergreen forests have been selectively logged in the past, and many areas continue to be logged. The impacts of such logging on amphibians are poorly understood. I examined the response of anuran communities to historical selective logging in a wet evergreen forest in south India. Anuran assemblages in unlogged forest were compared with assemblages in selectively logged forest. Forty 10 m × 10 m quadrats in forest, riparian zones, and streams of unlogged and selectively logged forests were searched at night for anurans. Species richness did not appear to be affected by logging. However, anuran density varied significantly and was 42 percent lower in selectively logged forests compared to unlogged forests. Anuran densities also varied significantly across microhabitats, with highest densities in streams of both selectively logged and unlogged forests. Patterns of niche overlap varied with selective logging as niche breadth either expanded, contracted, or remained neutral for different species. Ordination analysis explained 95 percent of the variation in species assemblage across selectively logged and unlogged forests. The assemblage in selectively logged forest was nested within unlogged forest. Among the habitat characteristics, litter thickness and water depth had the highest influence on the assemblage. This was followed by litter/water temperature, air temperature, and lastly relative humidity. It appears that species richness and composition of anurans in selectively logged forests is converging with unlogged forests, but the effects of historical logging seem to persist on anuran densities and their niche characteristics even ca 40 yr since logging ceased.     

Ecoregion and land‐use influence invertebrate and detritus transport from headwater streams

Summary 1. Habitats are often connected by fluxes of energy and nutrients across their boundaries. For example, headwater streams are linked to surrounding riparian vegetation through invertebrate and leaf litter inputs, and there is evidence that consumers in downstream habitats are subsidised by resources flowing from headwater systems. However, the strength of these linkages and the manner in which potential headwater subsidies vary along climatic and disturbance gradients are unknown. 2. We quantified the downstream transport of invertebrates, organic matter and inorganic sediment from 60 fishless headwater streams in the Wenatchee River Basin located on the eastern slope of the Cascade Range in Washington, U.S.A. Streams were classified into four groups (each n = 15) based on their position within two ecological subregions (wet and dry) and the extent of past timber harvest and road development (logged and unlogged). 3. Time and ecoregion were significant for all response variables as transport varied across sampling periods, and dry ecoregion streams displayed significantly higher mean values. Logged sites also generally showed higher mean transport, but only inorganic sediment transport was significantly higher in logged sites. Both ecoregion and land‐use interacted significantly with time depending on the response variable. Differences among stream categories were driven by relatively low levels of transport in unlogged drainages of the wet ecoregion. Interestingly, unlogged dry ecoregion streams showed comparable transport rates to logged sites in the wet ecoregion. Dominance by deciduous riparian vegetation in all but unlogged streams in the wet ecoregion is a primary hypothesised mechanism determining transport dynamics in our study streams. 4. Understanding the quantity and variation of headwater subsidies across climate and disturbance gradients is needed to appreciate the significance of ecological linkages between headwaters and associated downstream habitats. This will enable the accurate assessment of resource management impacts on stream ecosystems. Predicting the consequences of natural and anthropogenic disturbances on headwater stream transport rates will require knowledge of how both local and regional factors influence these potential subsidies. Our results suggest that resources transported from headwater streams reflect both the meso‐scale land‐use surrounding these areas and the constraints imposed by the ecoregion in which they are embedded.     

Influences of upland timber harvest on aquatic invertebrate communities in seasonal ponds: efficacy of forested buffers

We assessed community responses of aquatic invertebrates in 16 small, seasonal ponds in a forested region of north central Minnesota, USA, to evaluate potential influences of timber harvest and efficacy of uncut forested buffers in adjacent uplands. Invertebrate data gathered before (2000) and during the first 4 years following clearcut timber harvest (2001–2004) indicated that tree removal was followed by shifts in aquatic invertebrate communities in adjacent seasonal ponds. Retention of forested buffers appeared to partially mitigate influences of tree removal, but benefits of buffers may be limited by wind throw or other factors. Additional research is needed to clarify relationships between ecological characteristics of seasonal ponds and upland silviculture activities, and to better document efficacy and longevity of forested buffers.     

Differences in the Effects of Selective Logging on Amphibian Assemblages in Three West African Forest Types

Making generalizations about the impact of commercial selective logging on biodiversity has so far remained elusive. Species responses to logging depend on a number of factors, many of which have not been studied in detail. These factors may include the natural forest conditions (forest types) under which logging impacts are investigated; but this question has so far remained unexamined. In a large‐scale replicate study we aimed at clarifying the relationship between logging and forest types on leaf litter frogs. We contrast three distinct and naturally occurring forest types, including wet evergreen, moist evergreen and semi‐deciduous forests. Selectively logged sites were compared with primary forest sites for each forest type. We found that the response of frog communities to logging varies in different forest types. In the wet evergreen forest, richness was higher in logged forest than primary forest, while diversity measures were not different between logged and primary forest habitats. In the moist evergreen, richness and diversity were higher in selectively logged areas compared with primary forest habitats. In the semi‐deciduous, logged forests were characterized by drastic loss of forest specialists, reduced richness, and diversity. These results indicate that the net effect of logging varies with respect to forest type. Forest types that are characterized by adverse climatic conditions (i.e., low rainfall and protracted dry seasons) are more likely to produce negative effects on leaf litter anuran communities. For comparisons of the impact of logging on species to be effective, future research must endeavor to include details of forest type.     

Effects of past logging and grazing on understory plant communities in a montane Colorado forest

Throughout Pinus ponderosa–Pseudotsuga menziesii forests of the southern Colorado Front Range, USA, intense logging and domestic grazing began at the time of Euro-American settlement in the late 1800s and continued until the early 1900s. We investigated the long-term impacts of these settlement-era activities on understory plant communities by comparing understory composition at a historically logged and grazed site to that of an environmentally similar site which was protected from past use. We found that species richness and cover within functional groups rarely differed between sites in either upland or riparian areas. Multivariate analyses revealed little difference in species composition between sites on uplands, though compositional differences were apparent in riparian zones. Our findings suggest that settlement-era logging and grazing have had only minor long-term impacts on understories of upland Front Range P. ponderosa–P. menziesii forests, though they have had a greater long-term influence on riparian understories, where these activities were likely the most intense. This article was written and prepared by US Government employees on official time, and therefore it is in the public domain and not subject to copyright.     

Ecology of leaf pack macroinvertebrate communities in streams of the Fraser River Basin, British Columbia

1. To characterise geographic and small scale variation in the structure of macroinvertebrate communities in stream leaf packs, we collected one to three natural leaf pack communities from 119 reference streams in the Fraser River Basin and quantified their variability and correlation with aspects of the stream environment at several scales. We also sampled leaf packs in 19 test streams in the same geographic area exposed to stressors (nine logged, seven farmed, three mined catchments) to evaluate the leaf pack community as a tool for bioassessment. 2. There was substantial variation in the composition of invertebrate communities in leaf packs among reference streams of the Fraser River Basin. Capnia and Zapada (stoneflies), Baetis and Ephemerella (mayflies) and Tvetnia (midge) were the most common taxa found in the leaf packs. There were three types of assemblages identified by non‐metric multidimensional scaling; Capnia, Baetis and Ephemerella communities. 3. Leaf pack communities from the 19 test streams were plotted on a non‐metric multidimensional scaling ordination of the reference communities, and 14 of 19 sites fell outside the 80% confidence ellipse of the reference sites, including eight of nine logged, four of seven farmed and one of three mined catchments. Most of these streams plotted on the ordination near the Ephemerella reference communities. Reference stream communities had a similar number of genera per leaf pack (12.0) and genera per site (18.7) as the test streams (12.6 genera per leaf pack and 18.7 genera per site). Among the test sites, the farmed catchments had higher genera per leaf pack (17.8) and genera per site (21.9) than either the logged (11.5 genera per leaf pack; 19.9 genera per site) or mined (3.4 genera per leaf pack; 7.7 genera per site) catchments. 4. Heterogeneity of leaf pack communities within a site decreased as the number of genera found at the site increased. This was determined by allometric regression of the number of genera found at a site on the maximum number of genera possible, given the average number found per leaf pack. 5. There was a significant relationship between the composition of the leaf pack invertebrate community and stream geography (latitude, longitude, altitude, stream order). Canonical correspondence analysis showed differences among ‘big river’, ‘mountain stream’ and ‘southern’ communities. 6. There was no relationship between the composition of the leaf pack invertebrate community and stream channel and flow characteristics (bank dimensions, flow, slope). There was a significant relationship between the composition of the leaf pack invertebrate community and water quality of the stream (oxygen, nitrogen, phosphorus, conductivity, pH, temperature). ‘Cold, oxygen rich water’ communities were distinguishable from communities in streams with warmer, lower oxygen concentration. ‘High nutrient water’ communities were also distinct from communities in low nutrient streams. There was no relationship between the composition of the leaf pack invertebrate community and the nature of the leaf pack itself (i.e. morphology, decomposition, coniferous needle content). 7. Invertebrate communities in leaf packs show substantial, interpretable variation among reference streams. They are sensitive to human stressors at a landscape scale such as forestry and agriculture. Their diversity and composition varies at different spatial scales in a way that is at least partially explained by the environment of the stream and its catchment area.     

Leaf litter breakdown, microbial respiration and shredder production in metal-polluted streams

1. If species disproportionately influence ecosystem functioning and also differ in their sensitivities to environmental conditions, the selective removal of species by anthropogenic stressors may lead to strong effects on ecosystem processes. We evaluated whether these circumstances held for several Colorado, U.S.A. streams stressed by Zn. 2. Benthic invertebrates and chemistry were sampled in five second–third order streams for 1 year. Study streams differed in dissolved metal concentrations, but were otherwise similar in chemical and physical characteristics. Secondary production of leaf‐shredding insects was estimated using the increment summation and size‐frequency methods. Leaf litter breakdown rates were estimated by retrieving litter‐bags over a 171 day period. Microbial activity on leaf litter was measured in the laboratory using changes in oxygen concentration over a 48 h incubation period. 3. Dissolved Zn concentrations varied eightfold among two reference and three polluted streams. Total secondary production of shredders was negatively associated with metal contamination. Secondary production in reference streams was dominated by Taenionema pallidum. Results of previous studies and the current investigation demonstrate that this shredder is highly sensitive to metals in Colorado headwater streams. Leaf litter breakdown rates were similar between reference streams and declined significantly in the polluted streams. Microbial respiration at the most contaminated site was significantly lower than at reference sites. 4. Our results supported the hypothesis that some shredder species contribute disproportionately to leaf litter breakdown. Furthermore, the functionally dominant taxon was also the most sensitive to metal contamination. We conclude that leaf litter breakdown in our study streams lacked functional redundancy and was therefore highly sensitive to contaminant‐induced alterations in community structure. We argue for the necessity of simultaneously measuring community structure and ecosystem function in anthropogenically stressed ecosystems.     

Importance of scale,land‐use,and stream network properties for riparian plant communities along an urban gradient

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Effects of forest disturbances on aquatic insect assemblages

Natural and human‐made disasters such as floods and logging occur in and around rivers. Stream‐dwelling aquatic insects respond to these disturbances in various ways. Primary consumers among them rely greatly on algae and leaf litter from riparian vegetation as food. Therefore, once a disturbance such as a flood has occurred, insects may find it difficult to find food in a stream, and the aquatic insect assemblage can be impacted greatly as a result. Disturbances in riparian areas also increase fine sediment loads into streams, damaging habitat and altering the aquatic insect assemblage. Deforestation impacts not only terrestrial but also aquatic animals. In this review paper, aquatic insect assemblages are assessed according to alterations in land use in and around streams. Following this paper, it is expected that clarification of aquatic insect fauna and their life cycles will progress and that the distribution and habitat use of aquatic insects will be afforded greater attention in forest management.     

Planted Riparian Buffer Zones in New Zealand: Do They Live Up to Expectations?

Abstract River and stream rehabilitation projects are increasing in number, but the success or failure of these projects has rarely been evaluated, and the extent to which buffers can restore riparian and stream function and species composition is not well understood. In New Zealand the widespread conversion of forest to agricultural land has caused degradation of streams and riparian ecosystems. We assessed nine riparian buffer zone schemes in North Island, New Zealand that had been fenced and planted (age range from 2 to 24 years) and compared them with unbuffered control reaches upstream or nearby. Macroinvertebrate community composition was our prime indicator of water and habitat quality and ecological functioning, but we also assessed a range of physical and water quality variables within the stream and in the riparian zone. Generally, streams within buffer zones showed rapid improvements in visual water clarity and channel stability, but nutrient and fecal contamination responses were variable. Significant changes in macroinvertebrate communities toward “clean water” or native forest communities did not occur at most of the study sites. Improvement in invertebrate communities appeared to be most strongly linked to decreases in water temperature, suggesting that restoration of in‐stream communities would only be achieved after canopy closure, with long buffer lengths, and protection of headwater tributaries. Expectations of riparian restoration efforts should be tempered by (1) time scales and (2) spatial arrangement of planted reaches, either within a catchment or with consideration of their proximity to source areas of recolonists.