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.     

Riparian plant species loss alters trophic dynamics in detritus-based stream ecosystems

Riparian vegetation is closely connected to stream food webs through input of leaf detritus as a primary energy supply, and therefore, any alteration of plant diversity may influence aquatic ecosystem functioning. We measured leaf litter breakdown rate and associated biological parameters in mesh bags in eight headwater streams bordered either with mixed deciduous forest or with beech forest. The variety of leaf litter types in mixed forest results in higher food quality for large-particle invertebrate detritivores (‘shredders’) than in beech forest, which is dominated by a single leaf species of low quality. Breakdown rate of low quality (oak) leaf litter in coarse mesh bags was lower in beech forest streams than in mixed forest streams, a consequence of lower shredder biomass. In contrast, high quality (alder) leaf litter broke down at similar rates in both stream categories as a result of similar shredder biomass in coarse mesh bags. Microbial breakdown rate of oak and alder leaves, determined in fine mesh bags, did not differ between the stream categories. We found however aquatic hyphomycete species richness on leaf litter to positively co-vary with riparian plant species richness. Fungal species richness may enhance leaf litter breakdown rate through positive effects on resource quality for shredders. A feeding experiment established a positive relationship between fungal species richness per se and leaf litter consumption rate by an amphipod shredder (Gammarus fossarum). Our results show therefore that plant species richness may indirectly govern ecosystem functioning through complex trophic interactions. Integrating microbial diversity and trophic dynamics would considerably improve the prediction of the consequences of species loss.     

Riparian land use and the relationship between the benthos and litter decomposition in tropical montane streams

1. Although stream–catchment interactions have been analysed in some detail in temperate environments, little is known about the effects of land‐use changes in the tropics. Here, we analyse differences in benthic communities (macroinvertebrates and fungi) under two contrasting land uses (mature secondary forest and pasture) in montane streams in north‐western Ecuador and their influence on the rates of litter processing. 2. Between 2005 and 2006, we used a combination of coarse and fine mesh bags to study the relative contribution of macroinvertebrates and fungi to processing of two types of litter, Alnus acuminata and Inga spectabilis, in three‐first‐order streams running through mature secondary montane forests and adjacent downstream reaches running through pastures. At the same time, we characterised the assemblages of shreddering macroinvertebrates and fungi communities and the litter processing rates in stream reaches under both vegetation types. 3. Litter processing rates attributable to invertebrate feeding (coarse mesh bags) were significantly slower in streams running through pastures. Nevertheless, shredder diversity and richness were similar between pasture and forest sections, while shredder abundance was significantly higher in forest streams (mainly Phylloicus sp. :Trichoptera). Fungal reproductive activity and litter processing rates were low (fine mesh bags) and did not differ significantly between pasture and forest stream reaches. 4. Phylloicus sp. abundance was the best predictor of the percentage of litter remaining in coarse mesh bags across pasture and forest sites. Neither shredder diversity nor their species richness was a significant predictor of mass loss, as most of the decomposition was performed by a single keystone species. Although litter decomposition by microbial decomposers was low, fungal biomass (but not diversity) was the best variable explaining the percentage of litter remaining in fine mesh bags. 5. Our data suggest that, in these Neotropical montane streams, land use can have a significant impact on the rates of critical ecosystem processes, such as litter decomposition. In this study, this effect was not mediated by a major shift in the structure of the benthos, but by a decrease in the abundance and relative representation of a single species whose life history makes it critical to litter processing. 6. This study highlights the significant role that macroinvertebrate fauna can have in the processing of litter in Neotropical streams and the predominant role that single species can have in terms of controlling stream ecosystem‐level processes. Understanding the extent to which these patterns affect the long‐term and large‐scale functioning of stream ecosystems still needs further research and will become increasingly important in terms of managing lotic ecosystems in the context of rapid land‐use change.     

Beyond cool: adapting upland streams for climate change using riparian woodlands

Managed adaptation could reduce the risks of climate change to the world's ecosystems, but there have been surprisingly few practical evaluations of the options available. For example, riparian woodland is advocated widely as shade to reduce warming in temperate streams, but few studies have considered collateral effects on species composition or ecosystem functions. Here, we use cross‐sectional analyses at two scales (region and within streams) to investigate whether four types of riparian management, including those proposed to reduce potential climate change impacts, might also affect the composition, functional character, dynamics and energetic resourcing of macroinvertebrates in upland Welsh streams (UK). Riparian land use across the region had only small effects on invertebrate taxonomic composition, while stable isotope data showed how energetic resources assimilated by macroinvertebrates in all functional guilds were split roughly 50:50 between terrestrial and aquatic origins irrespective of riparian management. Nevertheless, streams draining the most extensive deciduous woodland had the greatest stocks of coarse particulate matter (CPOM) and greater numbers of ‘shredding’ detritivores. Stream‐scale investigations showed that macroinvertebrate biomass in deciduous woodland streams was around twice that in moorland streams, and lowest of all in streams draining non‐native conifers. The unexpected absence of contrasting terrestrial signals in the isotopic data implies that factors other than local land use affect the relative incorporation of allochthonous subsidies into riverine food webs. Nevertheless, our results reveal how planting deciduous riparian trees along temperate headwaters as an adaptation to climate change can modify macroinvertebrate function, increase biomass and potentially enhance resilience by increasing basal resources where cover is extensive (>60 m riparian width). We advocate greater urgency in efforts to understand the ecosystem consequences of climate change adaptation to guide future actions.     

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.     

Scale and the detection of land-use effects on morphology, vegetation and macroinvertebrate communities of grassland streams

1. Land‐use studies are challenging because of the difficulty of finding catchments that can be used as replicates and because land‐use effects may be obscured by sources of variance acting over spatial scales smaller than the catchment. To determine the extent to which land‐use effects on stream ecosystems are scale dependent, we designed a whole‐catchment study of six matched pairs (pasture versus native tussock) of second‐order stream catchments, taking replicate samples from replicate bedforms (pools and riffles) in each stream. 2. Pasture streams had a smaller representation of endemic riparian plant species, particularly tussock grasses, higher bank erosion, a somewhat deeper layer of fine sediment, lower water velocities in riffles, less moss cover and higher macroinvertebrate biodiversity. At the bedform scale, suspendable inorganic sediment (SIS) was higher in pools than riffles and in pasture streams there was a negative relationship between SIS and the percentage of the bed free of overhanging vegetation. Differences between stream reaches (including any interactions between land use and stream pair) were significant for SIS, substrate depth and characteristics of riparian vegetation. There were also significant differences between replicate bedforms in the same stream reaches in percentage exotic species in overhanging vegetation, percentage moss cover, QMCI (Quantitative Macroinvertebrate Community Index – a macroinvertebrate‐based stream health index) and macroinvertebrate density. 3. Significant differences among stream reaches and among replicate bedform units within the same reach, as well as interactions between these spatial units and land‐use effects, are neither trivial nor ‘noise’ but represent real differences among spatial units that typically are unaccounted for in stream studies. Our multi‐scale study design, accompanied by an investigation of the explanatory power of different factors operating at different scales, provides an improved understanding of variability in nature.     

Composition of riparian litter input regulates organic matter decomposition: Implications for headwater stream functioning in a managed forest landscape

Although the importance of stream condition for leaf litter decomposition has been extensively studied, little is known about how processing rates change in response to altered riparian vegetation community composition. We investigated patterns of plant litter input and decomposition across 20 boreal headwater streams that varied in proportions of riparian deciduous and coniferous trees. We measured a suite of in‐stream physical and chemical characteristics, as well as the amount and type of litter inputs from riparian vegetation, and related these to decomposition rates of native (alder, birch, and spruce) and introduced (lodgepole pine) litter species incubated in coarse‐ and fine‐mesh bags. Total litter inputs ranged more than fivefold among sites and increased with the proportion of deciduous vegetation in the riparian zone. In line with differences in initial litter quality, mean decomposition rate was highest for alder, followed by birch, spruce, and lodgepole pine (12, 55, and 68% lower rates, respectively). Further, these rates were greater in coarse‐mesh bags that allow colonization by macroinvertebrates. Variance in decomposition rate among sites for different species was best explained by different sets of environmental conditions, but litter‐input composition (i.e., quality) was overall highly important. On average, native litter decomposed faster in sites with higher‐quality litter input and (with the exception of spruce) higher concentrations of dissolved nutrients and open canopies. By contrast, lodgepole pine decomposed more rapidly in sites receiving lower‐quality litter inputs. Birch litter decomposition rate in coarse‐mesh bags was best predicted by the same environmental variables as in fine‐mesh bags, with additional positive influences of macroinvertebrate species richness. Hence, to facilitate energy turnover in boreal headwaters, forest management with focus on conifer production should aim at increasing the presence of native deciduous trees along streams, as they promote conditions that favor higher decomposition rates of terrestrial plant litter.     

Land-use effects on structural and functional composition of benthic and leaf-associated macroinvertebrates in four Andean streams

The replacement of native forests by pastures takes place widely in the Andes. The effects of such land-use change on aquatic assemblages are poorly understood. We conducted a comparative analysis of the effects of forest conversion to pastures on the taxonomic, structural, and functional composition of macroinvertebrates (benthic and leaf-associated) in montane and upper montane streams (ecosystem type) of the south Ecuadorian Andes. Taxonomic composition of benthic and leaf-associated macroinvertebrates was different between ecosystem type and land use. Also, major differences in the structural and functional composition of benthic and leaf-associated macroinvertebrates were mainly promoted by land use in both ecosystem types. Forested streams showed higher diversity than pasture streams, sustaining more shredder, scraper, and predatory invertebrates. We also observed differences in the macroinvertebrate communities between benthic and leaf-bag samples. Leaf bags had lower diversity and more collector invertebrates than benthic samples. This study highlights the large effect of riparian forest conversion to pasture land on macroinvertebrate communities, and the importance of using appropriate sampling techniques to characterize aquatic assemblages. We also recommend the maintenance and restoration of riparian vegetation to mitigate the effects of deforestation on stream communities and ecosystem processes.     

Ecosystem Functioning and Community Structure as Indicators for Assessing Environmental Impacts: Leaf Processing and Macroinvertebrates in Atlantic Forest Streams

Programs for evaluating environmental impacts are often carried out with chemical analyses that are inadequate for evaluating ecological aspects of rivers. In recent times there has been a strong movement towards biomonitoring using indices based on the structure of communities of organisms. Less attention has been given to functional parameters (ecosystem level processes such as decomposition, primary production, nutrient cycling, etc.), despite recent work that demonstrates their applicability. The breakdown of leaf material in streams provides a system in which an aspect of ecosystem functioning (decomposition) can be measured along with the community structure of the fauna associated with the leaves. We measured rate of leaf processing and the associated macroinvertebrates in 9 streams in 3 categories of environmental impact: Reference (pristine forest), Intermediate (pasture land‐use with intact riparian vegetation) and Impacted (pasture land‐use with degraded banks and stream‐bed). Leaf processing was fastest in the least impacted, “Reference”, streams. The abundance and taxon diversity of the associated fauna were not different among categories of impact. Thus this aspect of ecosystem functioning was more sensitive to perturbation than was the structure of the community in this case. We suggest that leaf processing can be a cheap and indicative parameter for biological assessment and monitoring. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Recent land‐use changes affect stream ecosystem processes in a subtropical island in Brazil

Land‐use changes such as conversion of natural forest to rural and urban areas have been considered as main drivers of ecosystem functions decline, and a large variety of indicators has been used to investigate these effects. Here, we used a replicated litter‐bag experiment to investigate the effects of land‐use changes on the leaf‐litter breakdown process and leaf‐associated invertebrates along the forest–pasture–urban gradient located in a subtropical island (Florianópolis, SC, Brazil). We identified the invertebrates and measured the litter breakdown rates using the litter bags approach. Litter bags containing 3 g of dry leaf of Alchornea triplinervia were deployed on forest rural and urban streams. Principal component analysis, based on physico‐chemical variables which, confirmed a gradient of degradation from forest to urban streams with intermediate values in rural areas. In accordance, shredder richness and abundance were lower in rural and urban than in forest streams. The land‐use changes led also to the dominance of tolerant generalist taxa (Chironomidae and Oligochaeta) reducing the taxonomic and functional diversity in these sites. Leaf‐litter breakdown rates decreased from forest to rural and finally to urban areas and were associated with changes in pH, water velocity, dissolved oxygen and abundance of leaf‐shredding invertebrates, although global decomposition rates did not differ between rural and urban streams. Overall, this study showed that land‐use changes, namely to rural and urban areas, have a strong impact on tropical streams ecosystems, in both processes and communities composition and structure. Despite of being apparently a smaller transformation of landscape, rural land use is comparable to urbanisation in terms of impact in stream functioning. It is thus critical to carefully plan urban development and maintain forest areas in the island of Florianópolis in order to preserve its natural biodiversity and aquatic ecosystems functioning.     

Effects of eucalyptus afforestation on leaf litter dynamics and macroinvertebrate community structure of streams in Central Portugal

To test the hypothesis whether afforestation with Eucalyptus globulus affects litter dynamics in streams and the structure of macroinvertebrate aquatic communities, we compared streams flowing through eucalyptus and deciduous forests, paying attention to: (i) litterfall dynamics, (ii) accumulation of organic matter, (iii) processing rates of two dominant leaf species: eucalyptus and chestnut, and (iv) macroinvertebrate community structure. The amount of allochthonous inputs was similar in both vegetation types, but the seasonality of litter inputs differed between eucalyptus and natural deciduous forests. Eucalyptus forest streams accumulated more organic matter than deciduous forest streams. Decomposition of both eucalyptus and chestnut leaf litter was higher in streams flowing through deciduous forests. The eucalyptus forest soils were highly hydrophobic resulting in strong seasonal fluctuations in discharge. In autumn the communities of benthic macroinvertebrates of the two stream types were significantly different. Deciduous forest streams contained higher numbers of invertebrates and more taxa than eucalyptus forest streams. Mixed forest streams (streams flowing through eucalyptus forests but bordered by deciduous vegetation) were intermediate between the two other vegetation types in all studied characteristics (accumulation of benthic organic matter, density and diversity of aquatic invertebrates). These results suggest that monocultures of eucalyptus affect low order stream communities. However, the impact may be attenuated if riparian corridors of original vegetation are kept in plantation forestry.     

Leaf breakdown in streams differing in catchment land use

1. The impact of changes in land use on stream ecosystem function is poorly understood. We studied leaf breakdown, a fundamental process of stream ecosystems, in streams that represent a range of catchment land use in the Piedmont physiographic province of the south‐eastern United States. 2. We placed bags of chalk maple (Acer barbatum) leaves in similar‐sized streams in 12 catchments of differing dominant land use: four forested, three agricultural, two suburban and three urban catchments. We measured leaf mass, invertebrate abundance and fungal biomass in leaf bags over time. 3. Leaves decayed significantly faster in agricultural (0.0465 day^?1) and urban (0.0474 day^?1) streams than in suburban (0.0173 day^?1) and forested (0.0100 day^?1) streams. Additionally, breakdown rates in the agricultural and urban streams were among the fastest reported for deciduous leaves in any stream. Nutrient concentrations in agricultural streams were significantly higher than in any other land‐use type. Fungal biomass associated with leaves was significantly lower in urban streams; while shredder abundance in leaf bags was significantly higher in forested and agricultural streams than in suburban and urban streams. Storm runoff was significantly higher in urban and suburban catchments that had higher impervious surface cover than forested or agricultural catchments. 4. We propose that processes accelerating leaf breakdown in agricultural and urban streams were not the same: faster breakdown in agricultural streams was due to increased biological activity as a result of nutrient enrichment, whereas faster breakdown in urban streams was a result of physical fragmentation resulting from higher storm runoff.     

The influences of climatic variation and vegetation on stream biota: lessons from the Big Dry in southeastern Australia

Aquatic biodiversity faces increasing threats from climate change, escalating exploitation of water and land use intensification. Loss of vegetation in catchments (= watersheds) has been identified as a substantial problem for many river basins, and there is an urgent need to better understand how climate change may interact with changes in catchment vegetation to influence the ecological condition of freshwater ecosystems. We used 20 years of biological monitoring data from Victoria, southeastern Australia, to explore the influences of catchment vegetation and climate on stream macroinvertebrate assemblages. Southeastern Australia experienced a severe drought from 1997 to 2009, with reductions of stream flows >50% in some areas. The prolonged drying substantially altered macroinvertebrate assemblages, with reduced prevalence of many flow‐dependent taxa and increased prevalence of taxa that are tolerant of low‐flow conditions and poor water quality. Stream condition, as assessed by several commonly used macroinvertebrate indices, was consistently better in reaches with extensive native tree cover in upstream catchments. Prolonged drought apparently caused similar absolute declines in macroinvertebrate condition indices regardless of vegetation cover, but streams with intact catchment and riparian vegetation started in better condition and remained so throughout the drought. The largest positive effects of catchment tree cover on both water quality and macroinvertebrate assemblages occurred above a threshold of ca. 60% areal tree cover in upstream catchments and in higher rainfall areas. Riparian tree cover also had positive effects on macroinvertebrate assemblages, especially in warmer catchments. Our results suggest that the benefits of extensive tree cover via improved water quality and in‐channel habitat persist during drought and show the potential for vegetation management to reduce negative impacts of climatic extremes for aquatic ecosystems.     

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

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Effects of agriculture on wood breakdown and microbial biofilm respiration in southern Appalachian streams

1. Agriculture causes high sediment, nutrient and light input to streams, which may affect rates of ecosystem processes, such as organic matter decay. In the southern Appalachians, socioeconomic trends over the past 50 years have caused widespread abandonment of farmland with subsequent reforestation. Physical and chemical properties of streams in these reforested areas may be returning to pre‐agriculture levels thereby creating the potential for recovery of ecosystem processes. 2. We examined wood breakdown and microbial activity on wood substrata in streams with different historical and current agricultural activity in their catchments. We analysed historical (1950) and recent (1998) forested land cover from large areas of the southern Appalachians and categorized streams based on percent forested land cover in these two time periods. Categories included a gradient of current agriculture from forested to heavily agricultural and reforestation from agriculture due to land abandonment. We compared microbial respiration on wood veneer substrata and breakdown of wood veneers among these land‐use categories. We also compared temperature, sediment accumulation and nitrogen and phosphorus concentrations. 3. Streams with current agriculture had higher concentrations of dissolved inorganic nitrogen than forested streams. Despite reforestation from agriculture, nitrogen concentrations were also elevated in streams with agricultural histories relative to forested streams. Temperature was also higher in agricultural streams but appeared to recover from historical agriculture through reforestation and stream shading. 4. Wood breakdown rates ranged from 0.0015 to 0.0076 day^?1 and were similar to other studies using wood veneers to determine breakdown rate. Microbial respiration increased with incubation time in streams up to approximately 150 days, after which it remained constant. Neither wood breakdown nor microbial respiration was significantly different among land‐use categories, despite the observed physical and chemical differences in streams based on land‐use. Wood breakdown rates could be predicted by microbial respiration indicating microbial control of wood breakdown in these streams. Both breakdown and microbial respiration were negatively correlated with the amount of inorganic sediment accumulated on wood veneers. 5. Higher nutrients and temperature led us to expect faster breakdown and higher microbial respiration in agricultural streams, but sediment in these streams may be limiting microbial activity and breakdown of organic material resulting in little net effect of agriculture on wood breakdown. Wood may not be desirable as a tool for functional assessment of stream integrity due to its unpredictable response to agriculture.     

Relationships between land use, spatial scale and stream macroinvertebrate communities

1. The structure of lotic macroinvertebrate communities may be strongly influenced by land‐use practices within catchments. However, the relative magnitude of influence on the benthos may depend upon the spatial arrangement of different land uses in the catchment. 2. We examined the influence of land‐cover patterns on in‐stream physico‐chemical features and macroinvertebrate assemblages in nine southern Appalachian headwater basins characterized by a mixture of land‐use practices. Using a geographical information system (GIS)/remote sensing approach, we quantified land‐cover at five spatial scales; the entire catchment, the riparian corridor, and three riparian ‘sub‐corridors’ extending 200, 1000 and 2000 m upstream of sampling reaches. 3. Stream water chemistry was generally related to features at the catchment scale. Conversely, stream temperature and substratum characteristics were strongly influenced by land‐cover patterns at the riparian corridor and sub‐corridor scales. 4. Macroinvertebrate assemblage structure was quantified using the slope of rank‐abundance plots, and further described using diversity and evenness indices. Taxon richness ranged from 24 to 54 among sites, and the analysis of rank‐abundance curves defined three distinct groups with high, medium and low diversity. In general, other macroinvertebrate indices were in accord with rank‐abundance groups, with richness and evenness decreasing among sites with maximum stream temperature. 5. Macroinvertebrate indices were most closely related to land‐cover patterns evaluated at the 200 m sub‐corridor scale, suggesting that local, streamside development effectively alters assemblage structure. 6. Results suggest that differences in macroinvertebrate assemblage structure can be explained by land‐cover patterns when appropriate spatial scales are employed. In addition, the influence of riparian forest patches on in‐stream habitat features (e.g. the thermal regime) may be critical to the distribution of many taxa in headwater streams draining catchments with mixed land‐use practices.     

Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.     

Deforestation and Benthic Indicators: How Much Vegetation Cover Is Needed to Sustain Healthy Andean Streams?

Deforestation in the tropical Andes is affecting ecological conditions of streams, and determination of how much forest should be retained is a pressing task for conservation, restoration and management strategies. We calculated and analyzed eight benthic metrics (structural, compositional and water quality indices) and a physical-chemical composite index with gradients of vegetation cover to assess the effects of deforestation on macroinvertebrate communities and water quality of 23 streams in southern Ecuadorian Andes. Using a geographical information system (GIS), we quantified vegetation cover at three spatial scales: the entire catchment, the riparian buffer of 30 m width extending the entire stream length, and the local scale defined for a stream reach of 100 m in length and similar buffer width. Macroinvertebrate and water quality metrics had the strongest relationships with vegetation cover at catchment and riparian scales, while vegetation cover did not show any association with the macroinvertebrate metrics at local scale. At catchment scale, the water quality metrics indicate that ecological condition of Andean streams is good when vegetation cover is over 70%. Further, macroinvertebrate community assemblages were more diverse and related in catchments largely covered by native vegetation (>70%). Our results suggest that retaining an important quantity of native vegetation cover within the catchments and a linkage between headwater and riparian forests help to maintain and improve stream biodiversity and water quality in Andean streams affected by deforestation. This research proposes that a strong regulation focused to the management of riparian buffers can be successful when decision making is addressed to conservation/restoration of Andean catchments.     

Resource quality and stoichiometric constraints on stream ecosystem functioning

1. Resource quality and stoichiometric imbalances in carbon : nutrient ratios between consumers and resources can influence key ecosystem processes. In many streams, this has important implications for food webs that are based largely upon the utilization of terrestrial leaf‐litter, which varies widely among litter types in its value as a food source for detritivores and as a substrate for microbial decomposers. 2. We measured breakdown rates and macroinvertebrate colonization of leaf‐litter from a range of native and exotic plants of differing resource quality and palatability to consumers [e.g. carbon : nitrogen : phosphorus (C : N : P) ratios, lignin and cellulose content], in a field experiment. We also measured C : N : P ratios of the principal leaf‐shredding invertebrates, which revealed strong stoichiometric imbalances across trophic levels: C : N and C : P ratios typically differed by at least one order of magnitude between consumers and resources, whereas N : P imbalances were less marked. Application of the threshold elemental ratio approach, which integrates animal bioenergetics and body elemental composition in examining nutrient deficiency between consumers and resources, revealed less marked C : P imbalances than those based on the simpler arithmetic differences described above. 3. Litter breakdown rates declined as nutrient imbalances widened and resource quality fell, but they were independent of whether resources were exotic or native. The principal drivers of total, microbial and invertebrate‐mediated breakdown rates were lignin : N, lignin : P and fungal biomass, respectively. However, multiple regression using orthogonal predictors yielded even more efficient models of litter breakdown, as consumers responded to more than one aspect of resource quality. For example, fungal biomass and litter C : N both influenced invertebrate‐mediated breakdown. 4. Large stoichiometric imbalances and changes in resource quality are likely to have serious consequences for stream ecosystem functioning, especially when riparian zones have been invaded by exotic plant species whose chemical composition differs markedly from that of the native flora. Consequently, the magnitude and direction of change in breakdown rates and, thus, resource depletion, will be driven to a large extent by the biochemical traits (rather than taxonomic identity per se) of the resident and invading flora.     

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.