Riparian forest composition affects stream litter decomposition despite similar microbial and invertebrate communities

Cross-boundary flows of energy and nutrients link biodiversity and functioning in adjacent ecosystems. The composition of forest tree species can affect the structure and functioning of stream ecosystems due to physical and chemical attributes, as well as changes in terrestrial resource subsidies. We examined how variation in riparian canopy composition (coniferous, deciduous, mixed) affects adjacent trophic levels (invertebrate and microbial consumers) and decomposition of organic matter in small, coastal rainforest streams in southwestern British Columbia. Breakdown rates of higher-quality red alder (Alnus rubra) litter were faster in streams with a greater percentage of deciduous than coniferous riparian canopy, whereas breakdown rates of lower-quality western hemlock (Tsuga heterophylla) litter were independent of riparian forest composition. When invertebrates were excluded using fine mesh, breakdown rates of both litter species were an order of magnitude less and were not significantly affected by riparian forest composition. Stream invertebrate and microbial communities were similar among riparian forest composition, with most variation attributed to leaf litter species. Invertebrate taxa richness and shredder biomass were higher in A. rubra litter; however, taxa evenness was greatest for T. heterophylla litter and both litter species in coniferous streams. Microbial community diversity (determined from terminal restriction fragment length polymorphisms) was unaffected by riparian forest or litter species. Fungal allele richness was higher than bacterial allele richness, and microbial communities associated with lower-quality T. heterophylla litter had higher diversity (allele uniqueness and richness) than those associated with higher-quality A. rubra litter. Percent variation in breakdown rates was mostly attributed to riparian forest composition in the presence of invertebrates and microbes; however, stream consumer biodiversity at adjacent trophic levels did not explain these patterns. Riparian and stream ecosystems and their biotic communities are linked through exchange and decomposition of detrital resources, and we provide evidence that riparian forest composition affects stream ecosystem catabolism despite similarities in microbial and invertebrate communities.     

Reciprocal subsidies between freshwater and terrestrial ecosystems structure consumer resource dynamics

Cross-ecosystem movements of material and energy, particularly reciprocal resource fluxes across the freshwater-land interface, have received major attention. Freshwater ecosystems may receive higher amounts of subsidies (i.e., resources produced outside the focal ecosystem) than terrestrial ecosystems, potentially leading to increased secondary production in freshwaters. Here we used a meta-analytic approach to quantify the magnitude and direction of subsidy inputs across the freshwater-land interface and to determine subsequent responses in recipient animals. Terrestrial and freshwater ecosystems differed in the magnitude of subsidies they received, with aquatic ecosystems generally receiving higher subsidies than terrestrial ecosystems. Surprisingly, and despite the large discrepancy in magnitude, the contribution of these subsidies to animal carbon inferred from stable isotope composition did not differ between freshwater and terrestrial ecosystems, likely due to the differences in subsidy quality. The contribution of allochthonous subsidies was highest to primary consumers and predators, suggesting that bottom-up and top-down effects may be affected considerably by the input of allochthonous resources. Future work on subsidies will profit from a food web dynamic approach including indirect trophic interactions and propagating effects.     

Short-term patch dynamics of macroinvertebrate colonization on decaying reed detritus in a Mediterranean lagoon (Lake Alimini Grande,Apulia, SE Italy)

In aquatic ecosystems, the ecology of macrophytal detritus aggregations has been investigated over a range of temporal scales. Yet, no attempts have been made to minimize the temporal resolution of the analysis, thus neglecting the occurrence of short-term colonization and decay dynamics for these ephemeral patches of high biological activity. The present study was carried out at two sampling sites in Lake Alimini Grande, a brackish lagoon in South-East Italy. The abundance of macroinvertebrate taxa colonizing prepared packs of reed leaf detritus and the ash-free dry mass of the packs were monitored daily for 40 days. Spectral analysis was used to investigate the fine-scale temporal dynamics of invertebrates colonization and leaf pack mass variations. The macrobenthos found on leaf packs was dominated at both sites by few taxa (i.e., the vagile Lekanesphaera monodi, Microdeutopus gryllotalpa, Neanthes caudata and the sessile Balanus sp.), whose general colonization patterns were described by unimodal curves. At a daily temporal resolution Balanus sp. was characterized by random short-term density variations, while vagile taxa showed non-random, highly site- and species-specific fluctuations generally characterized by an oscillation period comprised between 2 and 3 days. In addition, leaf packs were characterized by non-random, short-term mass oscillations, coherent and in phase with L. monodi abundance fluctuations. Our results highlight the occurrence of short-term, non-random density variations of invertebrate colonizers on leaf detritus patches. In addition, they emphasize a previously unexplored, fine-scale linkage between abundance patterns of invertebrate consumers and short-term mass variations of decaying detritus.     

An unusual trophic subsidy and species dominance in a tropical stream

In classical theory, species are assumed to achieve dominance through competitive exclusion, but if food resources are limiting, cross-habitat trophic subsidies could also underpin dominance. The impact of dominant species on community dynamics may depend on the energy base of population size. We report on an unusual, spatially subsidized population of a tropical, stream-dwelling crab that dominates the benthic fauna of a Kenyan stream. Diet and stable isotope analyses indicated that this crab is a true omnivore, with terrestrial subsidies dominating both plant and animal resources. Unusually, the animal prey included almost no aquatic invertebrates. Instead, a single species of ant constituted approximately 35% of the annual diet (stomach contents analysis) and up to 90% of assimilated nitrogen (estimates from stable isotope analysis). Ants may be pivotal to enabling crab dominance, and this crab may be largely disconnected from the local trophic network for its dietary needs. The paucity of other invertebrates in the stream community suggests that this super-dominant crab is a strong interactor that suppresses aquatic invertebrate populations. Common stabilizing attributes of spatially subsidized food webs (e.g., asynchronous prey availability, wide feeding niche, consumer migration) were absent from this system, and although apparently stable, it may be vulnerable to disturbance in the donor habitat.     

Meta-analysis: trophic level, habitat, and productivity shape the food web effects of resource subsidies

Studies of the effects of cross-habitat resource subsidies have been a feature of food web ecology over the past decade. To date, most studies have focused on demonstrating the magnitude of a subsidy or documenting its effect in the recipient habitat. Ecologists have yet to develop a satisfactory framework for predicting the magnitude of these effects. We used 115 data sets from 32 studies to compare consumer responses to resource subsidies across recipient habitat type, trophic level, and functional group. Changes in consumer density or biomass in response to subsidies were inconsistent across habitats, trophic, and functional groups. Responses in stream cobble bar and coastline habitats were larger than in other habitats. Contrary to expectation, the magnitude of consumer response was not affected by recipient habitat productivity or the ratio of productivity between donor and recipient habitats. However, consumer response was significantly related to the ratio of subsidy resources to equivalent resources in the recipient habitat. Broad contrasts in productivity are modified by subsidy type, vector, and the physical and biotic characteristics of both donor and recipient habitats. For this reason, the ratio of subsidy to equivalent resources is a more useful tool for predicting the possible effect of a subsidy than coarser contrasts of in situ productivity. The commonness of subsidy effects suggests that many ecosystems need to be studied as open systems.     

Terrestrial organic matter as subsidies that aid in the recovery of macroinvertebrates in industrially damaged lakes

The importance of allochthonous carbon to the productivity of stream ecosystems in temperate ecozones is well understood, but this relationship is less established in oligotrophic lakes. The nearshore littoral zones, at the interface of terrestrial and aquatic systems, are areas where the influence of terrestrial subsidies is likely greatest. We investigated the response of nearshore communities to variation in the quantity and composition of allochthonous materials, determined the landscape characteristics that regulate the variation of this subsidy, and explored the potential for terrestrial restoration practices to influence the export of organic matter to lakes. Stepwise multiple regressions revealed that diversity of nearshore macroinvertebrate families increased with the amount of fine particulate organic matter (FPOM) captured in sediment traps. The quantity of FPOM (g) increased with forest cover, and the relative amount of FPOM (percentage of total particulate material) in the traps increased with surface area of wetland in the catchments. These models suggest that terrestrially derived subsidies are important in smelter-impacted watersheds, and that the restoration of forests and wetlands will speed the return of nearshore consumer community diversity in industrially damaged lakes.     

Unexplored Diversity and Conservation Potential of Neotropical Hot Caves

The term hot cave is used to describe some subterranean chambers in the Neotropics that are characterized by constantly high ambient temperatures generated by the body heat of high densities of certain bat species. Many of these species have limited geographic ranges, and some occur only in the hot‐cave environment. In addition to the bats, the stable microclimate and abundant bat guano provides refuge and food for a high diversity of invertebrates. Hot caves have so far been described in the Caribbean and in a few isolated locations from Mexico to Brazil, although there is some evidence that similar caves may be present throughout the tropics. The existing literature suggests these poorly known ecosystems, with their unique combination of geomorphology and bat‐generated microclimate, are particularly sensitive to disturbance and face multiple threats from urbanization, agricultural development, mining, and tourism. Diversidad No Explorada y Potencial de Conservación de Cuevas Neotropicales Calientes     

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.     

Consumer-resource body-size relationships in natural food webs

It has been suggested that differences in body size between consumer and resource species may have important implications for interaction strengths, population dynamics, and eventually food web structure, function, and evolution. Still, the general distribution of consumer-'resource body-size ratios in real ecosystems, and whether they vary systematically among habitats or broad taxonomic groups, is poorly understood. Using a unique global database on consumer and resource body sizes, we show that the mean body-size ratios of aquatic herbivorous and detritivorous consumers are several orders of magnitude larger than those of carnivorous predators. Carnivorous predator-prey body-size ratios vary across different habitats and predator and prey types (invertebrates, ectotherm, and endotherm vertebrates). Predator-prey body-size ratios are on average significantly higher (1) in freshwater habitats than in marine or terrestrial habitats, (2) for vertebrate than for invertebrate predators, and (3) for invertebrate than for ectotherm vertebrate prey. If recent studies that relate body-size ratios to interaction strengths are general, our results suggest that mean consumer-resource interaction strengths may vary systematically across different habitat categories and consumer types.     

Predators, prey, and transient states in the assembly of spatially structured communities

Ecological theory suggests that both dispersal limitation and resource limitation can exert strong effects on community assembly. However, empirical studies of community assembly have focused almost exclusively on communities with a single trophic level. Thus, little is known about the combined effects of dispersal and resource limitation on assembly of communities with multiple trophic levels. We performed a landscape-scale experiment using spatially arranged mesocosms to study effects of dispersal and resource limitation on the assembly dynamics of aquatic invertebrate communities with two trophic levels. We found that interplay between dispersal and resource limitation regulated the assembly of predator and prey trophic levels in these pond communities. Early in assembly, predators and prey were strongly dispersal limited, and resource (i.e., prey) availability did not influence predator colonization. Later in assembly, after predators colonized, resource limitation was the strongest driver of predator abundance, and dispersal limitation played a negligible role. Thus, habitat isolation affected predators directly by reducing predator colonization rate, and indirectly through the effect of distance on prey availability. Dispersal and resource limitation of predators resulted in a transient period in which predators were absent or rare in isolated habitats. This period may be important for understanding population dynamics of vulnerable prey species. Our findings demonstrate that dispersal and resource limitation can jointly regulate assembly dynamics in multi-trophic systems. They also highlight the need to develop a temporal picture of the assembly process in multi-trophic communities because the availability and spatial distribution of limiting resources (i.e., prey) and the distribution of predators can shift radically over time.     

Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.     

Resources from another place and time: responses to pulses in a spatially subsidized system

As the theoretical bases for the dynamics of spatially subsidized communities emerge, ecologists question whether spatially subsidized communities exhibit similar structure or dynamics to communities that receive strongly pulsed resources. In both cases, communities may be structured by responses to resources that are potentially absent at any given point in time (pulsed communities) or space (subsidized communities), even if pulsed resources are part of the in situ productivity of the system or the subsidies arrive as a relatively constant input from a nearby system. The potential for significant spatial or temporal resource limitation, therefore, may be a key factor influencing in similar ways the persistence of populations, the structure and dynamics of communities, and the evolution of specific life history traits. In most complex systems, however, multiple resources may arrive for various trophic entities at various points in time and from various points in space, and thus it may be difficult to separate or compare the dynamics of spatially subsidized and pulsed systems. In this paper, we explore the effects of interactions between pulses and subsidies in plant and animal populations and communities on highly pulsed and variably subsidized islands in the Gulf of California. While many of the plant and animal communities on the unsubsidized islands in this system respond to pulses of rain in classic ways, responses to these rain pulses on islands subsidized by seabird guano or other marine resources are quite different and variable, and depend on a combination of life history characteristics, physiology, competitive interactions, and trophic relationships. These variable responses to rain pulses then translate into large differences in dynamics and community structure of subsidized vs. unsubsidized islands. Indeed, most systems experience both temporal pulses and spatial subsidies. When considered in tandem, complementary or synergistic effects of the multiple, temporally and spatially variable resources may emerge that help explain complex food web structure and dynamics.     

Decomposers (Lumbricidae, Collembola) affect plant performance in model grasslands of different diversity

Decomposer invertebrates influence soil structure and nutrient mineralization as well as the activity and composition of the microbial community in soil and therefore likely affect plant performance and plant competition. We established model grassland communities in a greenhouse to study the interrelationship between two different functional groups of decomposer invertebrates, Lumbricidae and Collembola, and their effect on plant performance and plant nitrogen uptake in a plant diversity gradient. Common plant species of Central European Arrhenatherion grasslands were transplanted into microcosms with numbers of plant species varying from one to eight and plant functional groups varying from one to four. Separate and combined treatments with earthworms and collembolans were set up. Microcosms contained 15N labeled litter to track N fluxes into plant shoots. Presence of decomposers strongly increased total plant and plant shoot biomass. Root biomass decreased in the presence of collembolans and even more in the presence of earthworms. However, it increased when both animal groups were present. Also, presence of decomposers increased total N concentration and 15N enrichment of grasses, legumes, and small herbs. Small herbs were at a maximum in the combined treatment with earthworms and collembolans. The impact of earthworms and collembolans on plant performance strongly varied with plant functional group identity and plant species diversity and was modified when both decomposers were present. Both decomposer groups generally increased aboveground plant productivity through effects on litter decomposition and nutrient mineralization leading to an increased plant nutrient acquisition. The non-uniform effects of earthworms and collembolans suggest that functional diversity of soil decomposer animals matters and that the interactions between soil animal functional groups affect the structure of plant communities.     

Nutrient subsidies to belowground microbes impact aboveground food web interactions

Historically, terrestrial food web theory has been compartmentalized into interactions among aboveground or belowground communities. In this study we took a more synthetic approach to understanding food web interactions by simultaneously examining four trophic levels and investigating how nutrient (nitrogen and carbon) and detrital subsidies impact the ability of the belowground microbial community to alter the abundance of aboveground arthropods (herbivores and predators) associated with the intertidal cord grass Spartina alterniflora. We manipulated carbon, nitrogen, and detrital resources in a field experiment and measured decomposition rate, soil nitrogen pools, plant biomass and quality, herbivore density, and arthropod predator abundance. Because carbon subsidies impact plant growth only indirectly (microbial pathways), whereas nitrogen additions both directly (plant uptake) and indirectly (microbial pathways) impact plant primary productivity, we were able to assess the effect of both belowground soil microbes and nutrient availability on aboveground herbivores and their predators. Herbivore density in the field was suppressed by carbon supplements. Carbon addition altered soil microbial dynamics (net potential ammonification, litter decomposition rate, DON [dissolved organic N] concentration), which limited inorganic soil nitrogen availability and reduced plant size as well as predator abundance. Nitrogen addition enhanced herbivore density by increasing plant size and quality directly by increasing inorganic soil nitrogen pools, and indirectly by enhancing microbial nitrification. Detritus adversely affected aboveground herbivores mainly by promoting predator aggregation. To date, the effects of carbon and nitrogen subsidies on salt marshes have been examined as isolated effects on either the aboveground or the belowground community. Our results emphasize the importance of directly addressing the soil microbial community as a factor that influences aboveground food web structure by affecting plant size and aboveground plant nitrogen.     

Plant genes link forests and streams

Although it is understood that the composition of riparian trees can affect stream function through leaf litter fall, the potential effects of genetic variation within species are less understood. Using a naturally hybridizing cottonwood system, we examined the hypothesis that genetic differences among two parental species (Populus fremontii and P. angustifolia) and two groups of their hybrids (F1 and backcrosses to P. angustifolia) would affect litter decomposition rates and the composition of the aquatic invertebrate community that colonizes leaves. Three major findings emerged: (1) parental and hybrid types differ in litter quality, (2) decomposition differs between two groups, a fast group (P. fremontii and F1 hybrid), and a slow group (P. angustifolia and backcross hybrids), and (3) aquatic invertebrate communities colonizing P. fremontii litter differed significantly in composition from all other cross types, even though P. fremontii and the F1 hybrid decomposed at similar rates. These findings are in agreement with terrestrial arthropod studies in the same cottonwood system. However, the effects are less pronounced aquatically than those observed in the adjacent terrestrial community, which supports a genetic diffusion hypothesis. Importantly, these findings argue that genetic interactions link terrestrial and aquatic communities and may have significant evolutionary and conservation implications.     

Biogeochemical consequences of rapid microbial turnover and seasonal succession in soil

Soil microbial communities have the metabolic and genetic capability to adapt to changing environmental conditions on very short time scales. In this paper we combine biogeochemical and molecular approaches to reveal this potential, showing that microbial biomass can turn over on time scales of days to months in soil, resulting in a succession of microbial communities over the course of a year. This new understanding of the year-round turnover and succession of microbial communities allows us for the first time to propose a temporally explicit N cycle that provides mechanistic hypotheses to explain both the loss and retention of dissolved organic N (DON) and inorganic N (DIN) throughout the year in terrestrial ecosystems. In addition, our results strongly support the hypothesis that turnover of the microbial community is the largest source of DON and DIN for plant uptake during the plant growing season. While this model of microbial biogeochemistry is derived from observed dynamics in the alpine, we present several examples from other ecosystems to indicate that the general ideas of biogeochemical fluxes being linked to turnover and succession of microbial communities are applicable to a wide range of terrestrial ecosystems.     

Lake to land subsidies: experimental addition of aquatic insects increases terrestrial arthropod densities

Aquatic insects are a common and important subsidy to terrestrial systems, yet little is known about how these inputs affect terrestrial food webs, especially around lakes. Myvatn, a lake in northern Iceland, has extraordinary midge (Chironomidae) emergences that result in large inputs of biomass and nutrients to terrestrial arthropod communities. We simulated this lake-to-land resource pulse by collecting midges from Myvatn and spreading their dried carcasses on 1-m2 plots at a nearby site that receives very little midge deposition. We hypothesized a positive bottom-up response of detritivores that would be transmitted to their predators and would persist into the following year. We sampled the arthropod community once per month for two consecutive summers. Midge addition resulted in significantly different arthropod communities and increased densities of some taxa in both years. Detritivores, specifically Diptera larvae, Collembola, and Acari increased in midge-addition plots, and so did some predators and parasitoids. Arthropod densities were still elevated a year after midge addition, and two years of midge addition further increased the density of higher-order consumers (e.g., Coleoptera and Hymenoptera). Midge addition increased arthropod biomass by 68% after one year and 108% after two years. By manipulating the nutrient pulse delivered by midges we were able to elucidate food web consequences of midge deposition and spatial and temporal dynamics that are difficult to determine based on comparative approaches alone. Resources cross ecosystem boundaries and are assimilated over time because of life-history strategies that connect aquatic and terrestrial food webs and these systems cannot be fully understood in isolation from each other.     

Environmental impact propagated by cross-system subsidy: chronic stream pollution controls riparian spider populations

Resource subsidies between habitats are common and create the potential for the propagation of environmental impacts across system boundaries. However, recent understanding of the potential for subsidy-mediated cross-system impact propagations is limited and primarily based on passive flows of nutrients and detritus or short-term effects. Here, we assess the effects of sustained alterations in aquatic insect emergence (active subsidy pathway), due to chronic stream pollution, for riparian spiders. The sustained reduction in aquatic insect densities at the polluted reaches resulted in a marked decline in web spider population density and a shift in spider community composition. Our results provide the first evidence that stream pollution can control populations and community structure of terrestrial predators via sustained alterations in aquatic subsidies, emphasizing the role of subtle trophic linkages in the transmission of environmental impacts across ecosystem boundaries.     

Introduced trout sever trophic connections in watersheds: consequences for a declining amphibian

Trophic linkages between terrestrial and aquatic ecosystems are increasingly recognized as important yet poorly known features of food webs. Here we describe research to understand the dynamics of lake food webs in relation to a native riparian amphibian and its interaction with introduced trout. The mountain yellow-legged frog Rana muscosa is endemic to alpine watersheds of the Sierra Nevada Mountains and the Transverse Ranges of California, but it has declined to a small fraction of its historical distribution and abundance. Although remaining frogs and introduced trout feed in different habitats of alpine lakes, our stable-isotope analyses clearly show that the same resource base of benthic invertebrates sustains their growth. During one period, insect emergence from naturally fishless lakes was nearly 20-fold higher compared to adjacent lakes with trout, showing that fish reduce availability of aquatic prey to amphibious and terrestrial consumers. Although trout cannot prey on adult frogs due to gape limitation, foraging post-metamorphic frogs are 10 times more abundant in the absence of trout, suggesting an important role for competition for prey by trout in highly unproductive alpine watersheds. Most Sierran lakes contain fish, and those that do not are usually small isolated ponds; in our study, these two lake types supported the lowest densities of post-metamorphic frogs, and these frogs were less reliant on local, benthic sources of productivity. Since Rana muscosa was formerly the most abundant vertebrate in the Sierra Nevada, the reduction in energy flow from lake benthos to this consumer due to fish introductions may have had negative consequences for its numerous terrestrial predators, many of which have also declined. We suggest that disruptions of trophic connections between aquatic and terrestrial food webs are an important but poorly understood consequence of fish introduction to many thousands of montane lakes and streams worldwide and may contribute to declines of native consumers in riparian habitats.     

Headwater riparian forest-floor invertebrate communities associated with alternative forest management practices.

Headwater streams and their riparian zones are a common, yet poorly understood, component of Pacific Northwest, USA, landscapes. We describe the ecological importance of headwater stream riparian zones as habitat for forest-floor invertebrate communities and assess how alternative management strategies for riparian zones may impact these communities. We compared community composition of forest-floor invertebrates at increasing distances along trans-riparian (stream edge to upslope) transects in mature forests, clearcuts, and riparian buffers of approximately 30-m width with upslope clearcuts. Invertebrates were collected using pitfall traps in five replicate blocks of three treatments each in the Willamette National Forest, Oregon, USA. We measured microclimate and microhabitat variables at pitfall locations. Despite strong elevation and block effects on community composition, community analyses revealed a distinct "riparian" invertebrate community within 1 m of the stream edge in mature forest treatments, which was strongly related to cool, humid microclimate conditions. Invertebrate community composition in buffer treatments was far more similar to that of mature forests than to clearcuts; a pattern mirrored by microclimate. These results suggest that, within our study sites, forest-floor invertebrate distributions are strongly associated with microclimate and that riparian buffers of approximately 30-m width do provide habitat for many riparian and forest species. Riparian reserves may serve as effective forest refugia and/or dispersal corridors for invertebrates and other taxa, and their incorporation into watershed management plans likely will contribute to meeting persistence and connectivity objectives.