From bottom-up to top-down control of invertebrate herbivores in a retrogressive chronosequence

In the long-term absence of disturbance, ecosystems often enter a decline or retrogressive phase which leads to reductions in primary productivity, plant biomass, nutrient cycling and foliar quality. However, the consequences of ecosystem retrogression for higher trophic levels such as herbivores and predators, are less clear. Using a post-fire forested island-chronosequence across which retrogression occurs, we provide evidence that nutrient availability strongly controls invertebrate herbivore biomass when predators are few, but that there is a switch from bottom-up to top-down control when predators are common. This trophic flip in herbivore control probably arises because invertebrate predators respond to alternative energy channels from the adjacent aquatic matrix, which were independent of terrestrial plant biomass. Our results suggest that effects of nutrient limitation resulting from ecosystem retrogression on trophic cascades are modified by nutrient-independent variation in predator abundance, and this calls for a more holistic approach to trophic ecology to better understand herbivore effects on plant communities.     

Trophic‐level dependent effects on CO2 emissions from experimental stream ecosystems

Concern over accelerating rates of species invasions and losses have initiated investigations into how local and global changes to predator abundance mediate trophic cascades that influence CO₂ fluxes of aquatic ecosystems. However, to date, no studies have investigated how species additions or losses at other consumer trophic levels influence the CO₂ flux of aquatic ecosystems. In this study, we added a large predatory stonefly, detritivorous stonefly, or grazer tadpole to experimental stream food webs and over a 70‐day period quantified their effects on community composition, leaf litter decomposition, chlorophyll‐a concentrations, and stream CO₂ emissions. In general, streams where the large grazer or large detritivore were added showed no change in total invertebrate biomass, leaf litter loss, chlorophyll‐a concentrations, or stream CO₂ emissions compared with controls; although we did observe a spike in CO₂ emissions in the large grazer treatment following a substantial reduction in chlorophyll‐a concentrations on day 28. However, the large grazer and large detritivore altered the community composition of streams by reducing the densities of other grazer and detritivore taxa, respectively, compared with controls. Conversely, the addition of the large predator created trophic cascades that reduced total invertebrate biomass and increased primary producer biomass. The cascading effects of the predator additions on the food web ultimately led to decreased CO₂ emissions from stream channels by up to 95%. Our results suggest that stream ecosystem processes were more influenced by changes in large predator abundance than large grazer or detritivore abundance, because of a lack of functionally similar large predators. Our study demonstrates that the presence/absence of species with unique functional roles may have consequences for the exchange of CO₂ between the ecosystem and the atmosphere.     

Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.     

Increasing donor ecosystem productivity decreases terrestrial consumer reliance on a stream resource subsidy

Because nutrient enrichment can increase ecosystem productivity, it may enhance resource flows to adjacent ecosystems as organisms cross ecosystem boundaries and subsidize predators in recipient ecosystems. Here, we quantified the biomass and abundance of aquatic emergence and terrestrial spiders in a reference and treatment stream that had been continuously enriched with nitrogen and phosphorus for 5 years. Because we previously showed that enrichment increased secondary production of stream consumers, we predicted that aquatic emergence flux would be higher in the treatment stream, subsequently increasing the biomass and abundance of terrestrial spiders. Those increases were predicted to be greatest for spiders specializing on aquatic emergence subsidies (e.g., Tetragnathidae). By adding a ¹⁵N stable isotope tracer to both streams, we also quantified nitrogen flow from the stream into the riparian community. Emergence biomass, but not abundance, was higher in the treatment stream. The average body size of emerging adult insects and the relative dominance of Trichoptera adults were also greater in the treatment stream. However, spider biomass did not differ between streams. Spiders also exhibited substantially lower reliance on aquatic emergence nitrogen in the treatment stream. This reduced reliance likely resulted from shifts in the body size distributions and community composition of insect emergence that may have altered predator consumption efficiency in the treatment stream. Despite nutrient enrichment approximately doubling stream productivity and associated cross-ecosystem resource flows, the response of terrestrial predators depended more on the resource subsidy’s characteristics that affected the predator’s ability to capitalize on such increases.     

Nutrient Enrichment and Food Web Composition Affect Ecosystem Metabolism in an Experimental Seagrass Habitat

Background

Food web composition and resource levels can influence ecosystem properties such as productivity and elemental cycles. In particular, herbivores occupy a central place in food webs as the species richness and composition of this trophic level may simultaneously influence the transmission of resource and predator effects to higher and lower trophic levels, respectively. Yet, these interactions are poorly understood.

Methodology/Principal Findings

Using an experimental seagrass mesocosm system, we factorially manipulated water column nutrient concentrations, food chain length, and diversity of crustacean grazers to address two questions: (1) Does food web composition modulate the effects of nutrient enrichment on plant and grazer biomasses and stoichiometry? (2) Do ecosystem fluxes of dissolved oxygen and nutrients more closely reflect above-ground biomass and community structure or sediment processes? Nutrient enrichment and grazer presence generally had strong effects on biomass accumulation, stoichiometry, and ecosystem fluxes, whereas predator effects were weaker or absent. Nutrient enrichment had little effect on producer biomass or net ecosystem production but strongly increased seagrass nutrient content, ecosystem flux rates, and grazer secondary production, suggesting that enhanced production was efficiently transferred from producers to herbivores. Gross ecosystem production (oxygen evolution) correlated positively with above-ground plant biomass, whereas inorganic nutrient fluxes were unrelated to plant or grazer biomasses, suggesting dominance by sediment microbial processes. Finally, grazer richness significantly stabilized ecosystem processes, as predators decreased ecosystem production and respiration only in the zero- and one- species grazer treatments.

Conclusions/Significance

Overall, our results indicate that consumer presence and species composition strongly influence ecosystem responses to nutrient enrichment, and that increasing herbivore diversity can stabilize ecosystem flux rates in the face of perturbations.     

Cascading predator control interacts with productivity to determine the trophic level of biomass accumulation in a benthic food web

Large-scale exploitation of higher trophic levels by humans, together with global-scale nutrient enrichment, highlights the need to explore interactions between predator loss and resource availability. The hypothesis of exploitation ecosystems suggests that top–down and bottom–up control alternate between trophic levels, resulting in a positive relationship between primary production and the abundance of every second trophic level. Specifically, in food webs with three effective trophic levels, primary producers and predators should increase with primary production, while in food webs with two trophic levels, only herbivores should increase. We provided short-term experimental support for these model predictions in a natural benthic community with three effective trophic levels, where the number of algal recruits, but not the biomass of gastropod grazers, increased with algal production. In contrast, when the food web was reduced to two trophic levels by removing larger predators, the number of algal recruits was unchanged while gastropod grazer biomass increased with algal production. Predator removal only affected the consumer-controlled early life-stages of algae, indicating that both the number of trophic levels and the life-stage development of the producer trophic level determine the propagation of trophic cascades in benthic systems. Our results support the hypothesis that predators interact with resource availability to determine food-web structure.     

A cross-system synthesis of consumer and nutrient resource control on producer biomass

Nutrient availability and herbivory control the biomass of primary producer communities to varying degrees across ecosystems. Ecological theory, individual experiments in many different systems, and system-specific quantitative reviews have suggested that (i) bottom-up control is pervasive but top-down control is more influential in aquatic habitats relative to terrestrial systems and (ii) bottom-up and top-down forces are interdependent, with statistical interactions that synergize or dampen relative influences on producer biomass. We used simple dynamic models to review ecological mechanisms that generate independent vs. interactive responses of community-level biomass. We calibrated these mechanistic predictions with the metrics of factorial meta-analysis and tested their prevalence across freshwater, marine and terrestrial ecosystems with a comprehensive meta-analysis of 191 factorial manipulations of herbivores and nutrients. Our analysis showed that producer community biomass increased with fertilization across all systems, although increases were greatest in freshwater habitats. Herbivore removal generally increased producer biomass in both freshwater and marine systems, but effects were inconsistent on land. With the exception of marine temperate rocky reef systems that showed positive synergism of nutrient enrichment and herbivore removal, experimental studies showed limited support for statistical interactions between nutrient and herbivory treatments on producer biomass. Top-down control of herbivores, compensatory behaviour of multiple herbivore guilds, spatial and temporal heterogeneity of interactions, and herbivore-mediated nutrient recycling may lower the probability of consistent interactive effects on producer biomass. Continuing studies should expand the temporal and spatial scales of experiments, particularly in understudied terrestrial systems; broaden factorial designs to manipulate independently multiple producer resources (e.g. nitrogen, phosphorus, light), multiple herbivore taxa or guilds (e.g. vertebrates and invertebrates) and multiple trophic levels; and - in addition to measuring producer biomass - assess the responses of species diversity, community composition and nutrient status.     

Environmental context determines multi‐trophic effects of consumer species loss

Loss of biodiversity and nutrient enrichment are two of the main human impacts on ecosystems globally, yet we understand very little about the interactive effects of multiple stressors on natural communities and how this relates to biodiversity and ecosystem functioning. Advancing our understanding requires the following: (1) incorporation of processes occurring within and among trophic levels in natural ecosystems and (2) tests of context‐dependency of species loss effects. We examined the effects of loss of a key predator and two groups of its prey on algal assemblages at both ambient and enriched nutrient conditions in a marine benthic system and tested for interactions between the loss of functional diversity and nutrient enrichment on ecosystem functioning. We found that enrichment interacted with food web structure to alter the effects of species loss in natural communities. At ambient conditions, the loss of primary consumers led to an increase in biomass of algae, whereas predator loss caused a reduction in algal biomass (i.e. a trophic cascade). However, contrary to expectations, we found that nutrient enrichment negated the cascading effect of predators on algae. Moreover, algal assemblage structure varied in distinct ways in response to mussel loss, grazer loss, predator loss and with nutrient enrichment, with compensatory shifts in algal abundance driven by variation in responses of different algal species to different environmental conditions and the presence of different consumers. We identified and characterized several context‐dependent mechanisms driving direct and indirect effects of consumers. Our findings highlight the need to consider environmental context when examining potential species redundancies in particular with regard to changing environmental conditions. Furthermore, non‐trophic interactions based on empirical evidence must be incorporated into food web‐based ecological models to improve understanding of community responses to global change.     

Competitive displacement alters top-down effects on carbon dioxide concentrations in a freshwater ecosystem

Climate change and invasive species have the potential to alter species diversity, creating novel species interactions. Interspecific competition and facilitation between predators may either enhance or dampen trophic cascades, ultimately influencing total predator effects on communities and biogeochemical cycling of ecosystems. However, previous studies have only investigated the effects of a single predator species on CO₂ flux of aquatic ecosystems. In this study, we measured and compared the individual and joint effects of predatory damselfly larvae and diving beetles on total prey biomass, leaf litter processing, and dissolved CO₂ concentrations of experimental bromeliad ecosystems. Damselfly larvae created strong trophic cascades that reduced CO₂ concentrations by ~46 % relative to no-predator treatments. Conversely, the effects of diving beetles on prey biomass, leaf litter processing, and dissolved CO₂ were not statistically different to no-predator treatments. Relative to multiplicative null models, the presence of damselfly larvae and diving beetles together resulted in antagonistic relations that eliminated trophic cascades and top-down influences on CO₂ concentrations. Furthermore, we showed that the antagonistic interactions between predators occurred due to a tactile response that culminated in competitive displacement of damselfly larvae. Our results demonstrate that predator identity and predator–predator interactions can influence CO₂ concentrations of an aquatic ecosystem. We suggest that predator effects on CO₂ fluxes may depend on the particular predator species removed or added to the ecosystem and their interactions with other predators.     

Cascading effects of belowground predators on plant communities are density‐dependent

Soil food webs comprise a multitude of trophic interactions that can affect the composition and productivity of plant communities. Belowground predators feeding on microbial grazers like Collembola could decelerate nutrient mineralization by reducing microbial turnover in the soil, which in turn could negatively influence plant growth. However, empirical evidences for the ecological significance of belowground predators on nutrient cycling and plant communities are scarce. Here, we manipulated predator density (Hypoaspis aculeifer: predatory mite) with equal densities of three Collembola species as a prey in four functionally dissimilar plant communities in experimental microcosms: grass monoculture (Poa pratensis), herb monoculture (Rumex acetosa), legume monoculture (Trifolium pratense), and all three species as a mixed plant community. Density manipulation of predators allowed us to test for density‐mediated effects of belowground predators on Collembola and lower trophic groups. We hypothesized that predator density will reduce Collembola population causing a decrease in nutrient mineralization and hence detrimentally affect plant growth. First, we found a density‐dependent population change in predators, that is, an increase in low‐density treatments, but a decrease in high‐density treatments. Second, prey suppression was lower at high predator density, which caused a shift in the soil microbial community by increasing the fungal: bacterial biomass ratio, and an increase of nitrification rates, particularly in legume monocultures. Despite the increase in nutrient mineralization, legume monocultures performed worse at high predator density. Further, individual grass shoot biomass decreased in monocultures, while it increased in mixed plant communities with increasing predator density, which coincided with elevated soil N uptake by grasses. As a consequence, high predator density significantly increased plant complementarity effects indicating a decrease in interspecific plant competition. These results highlight that belowground predators can relax interspecific plant competition by increasing nutrient mineralization through their density‐dependent cascading effects on detritivore and soil microbial communities.     

Benthic community responses to nutrient enrichment and predator exclusion: Influence of background nutrient concentrations and interactive effects

Potential community effects of nutrient enhancement are a topic of theoretical interest and increasing management concern in coastal marine systems. While increased nutrient levels may lead to increased microalgal production and biomass, studies have provided variable evidence regarding the existence of upward cascade effects on macrofauna. In benthic marine communities, limitation by predation or factors preventing recruitment response may contribute to weak coupling between resource availability and macrobenthos abundances. We conducted blocked nutrient addition and predator exclusion experiments in the intertidal of two estuaries that varied in background nutrient concentrations (Cape Fear and White Oak, southeastern North Carolina). Benthic community comparisons were also made among these and two other North Carolina estuaries to examine correlations in distribution patterns. Cape Fear, which had the highest background nitrogen and phosphorus concentrations, also had highest ambient benthic microalgal biomass. There was no significant response of microalgal biomass to local nutrient additions in Cape Fear and only one macrofaunal taxon during one season exhibited abundance responses to nutrient additions. White Oak, with lower background nutrient levels, was characterized by significant microalgal responses to nutrient additions and significant macrofauna abundance responses for 50% of the species examined during summer experiments. However, all of these macrofauna declined in abundance with nutrient enhancement while biomass remained constant or significantly increased with nutrient additions. This suggests a complex response of macrofauna to nutrient additions in this estuary with greater biomass per individual but a corresponding decline in abundances. Top-down/bottom-up interactive effects were observed for haustoriid amphipods, which were uncommon or absent when predators had access, but exhibited strong biomass responses to nutrient enhancement when predators were excluded. These results support a growing body of literature that indicates the importance of background conditions in regulating benthic community responses to nutrient enhancement. However, responses may be complex with biomass per individual rather than densities being the primary response variable for some taxa and predator moderation of responses occurring for some taxa but not others.     

Seasonal influence of brook trout and mottled sculpin on lower trophic levels in an Appalachian stream

1. In some situations fish have strong top‐down effects in stream communities while in others they seem to be relatively unimportant. Differences in the impact of fish may depend on a variety of factors including the foraging mode of the fish, interactions among fish species and temporal variation in environmental conditions and species interactions. 2. We investigated the effect of brook trout (Salvelinus fontinalis) and mottled sculpin (Cottus bairdi) on lower trophic levels in Appalachian streams and whether or not interactions between these fish changed their influence. Mesocosms were placed in a headwater stream in a randomized complete block design. Within blocks, mesocosms were randomly assigned to one of the following treatments: (i) no fish; (ii) sculpin only; (iii) trout only and (iv) both sculpin and trout. Fish biomass was the same in all three fish treatments. Invertebrate density and algal biomass in mesocosms were determined after 3 weeks. We repeated the experiment in the autumn, spring and summer to test for seasonality of fish effects. 3. The effect of fish on invertebrate assemblages was seasonal and depended on prey identity. Sculpin strongly suppressed grazer abundance in spring while trout had little effect on grazers in any season. The influence of both fish on insect predators was similar and relatively constant across seasons. We found little evidence of an interaction between sculpin and trout that strongly influenced their effect on prey across seasons. 4. None of the fish treatments influenced algal biomass during any of the seasons. Algal growth was also seasonal, with a two‐ to four‐fold increase in algal biomass in spring compared to autumn and summer. 5. Our results indicate that benthic and drift feeding fish differ in their effects on some, but not all prey. Furthermore, fish effects on prey were strongly seasonal for some, but not all prey types. While the temporal context is not commonly considered, our results indicate seasonality can be an important component of predator–prey interactions in streams.     

Influence of herbaceous riparian buffers on physical habitat, water chemistry, and stream communities within channelized agricultural headwater streams

Herbaceous riparian buffers (CP 21 grass filter strips) are a widely used agricultural conservation practice in the United States for reducing nutrient, pesticide, and sediment loadings to agricultural streams. The ecological impacts of herbaceous riparian buffers on the channelized agricultural headwater streams that are common throughout the midwestern United States have not been evaluated. We sampled riparian habitat, geomorphology, instream habitat, water chemistry, fishes, and amphibians for 4 years from three channelized agricultural headwater streams without herbaceous riparian buffers and three channelized streams with herbaceous riparian buffers in central Ohio. Only seven of 55 response variables exhibited differences between buffer types. Riparian widths were greater in channelized headwater streams with herbaceous riparian buffers than streams without herbaceous riparian buffers. Percent insectivores and minnows were greater in channelized streams without herbaceous riparian buffers than streams with herbaceous riparian buffers. Percent clay, turbidity, specific conductance, and pH differed between buffer types only during one sampling period. No differences in geomorphology and amphibian communities occurred between buffer types. Our results suggest channelized agricultural headwater streams with and without herbaceous riparian buffers are similar physically, chemically, and biologically. Installation of herbaceous riparian buffers alone adjacent to channelized agricultural headwater streams in central Ohio and other parts of the midwestern United States may only provide limited environmental benefits for these stream ecosystems in the first 4-6 years after establishment. Alternative implementation designs combining the use of herbaceous riparian buffers with other practices capable of altering nutrient and pesticide loads, riparian hydrology, and instream habitat are needed.     

Benthic invertebrate communities in six Danish forest streams: Impact of forest type on structure and function

I investigated the impact of riparian vegetation type on stream invertebrate communities in SIX Danish forest streams during the period 1992–1993 Two of the streams ran through beech forest Fagus sylvatica , two through mixed, mainly deciduous forest, and two through conifer plantations Algal biomass and detritus standing stock differed significantly depending on forest type In mixed deciduous forest streams algal biomass was higher and detritus standing stock lower than in beech and conifer forest streams Benthic invertebrate community composition was functionally similar in all streams, with detritivores comprising >80% by numbers of the community Grazer abundance was low in all streams Shredder abundance in the four deciduous streams was significantly correlated to coarse detritus standing stock When taking into account both quantitative and qualitative POM parameters for all six streams combined, shredder abundance correlated significantly to the amount of CPOM There were significantly more invertebrates with a ≥ 2 yr life cycle in the conifer and beech forest streams than m the mixed forest streams, probably reflecting the larger and more stable food resource For all streams combined there was a significant correlation between predator abundance and the abundance of potential prey My findings suggest that Danish forest streams are regulated by "bottom-up control" at all trophic levels within the invertebrate community, and hence that forest type can structure benthic communities in forest streams     

Herbivore and predator diversity interactively affect ecosystem properties in an experimental marine community

Interacting changes in predator and prey diversity likely influence ecosystem properties but have rarely been experimentally tested. We manipulated the species richness of herbivores and predators in an experimental benthic marine community and measured their effects on predator, herbivore and primary producer performance. Predator composition and richness strongly affected several community and population responses, mostly via sampling effects. However, some predators survived better in polycultures than in monocultures, suggesting complementarity due to stronger intra- than interspecific interactions. Predator effects also differed between additive and substitutive designs, emphasizing that the relationship between diversity and abundance in an assemblage can strongly influence whether and how diversity effects are realized. Changing herbivore richness and predator richness interacted to influence both total herbivore abundance and predatory crab growth, but these interactive diversity effects were weak. Overall, the presence and richness of predators dominated biotic effects on community and ecosystem properties.     

Effect of emergent aquatic insects on bat foraging in a riparian forest

1. Riparian zones serve several ecological functions for bats. They provide a source of prey and likely provide favourable structural habitats and shelter from predators. Many studies have shown that bats use the space above streams, ponds or riparian vegetation as feeding habitat. These studies, however, have never distinguished between the effects of habitat structure and prey availability on the foraging activities of bats. Such effects can only be distinguished by an experimental approach. We predicted that bat activity along a stream is influenced by the number of emerged aquatic insects. 2. We evaluated the response of terrestrial consumers, insectivorous bats, to changes in the abundance of emergent aquatic insects by conducting a manipulative field experiment. In a deciduous riparian forest in Japan, aquatic insect flux from the stream to the riparian zone was controlled with an insect-proof cover over a 1.2 km stream reach. 3. We estimated the abundance of emergent aquatic and flying terrestrial arthropods near the treatment and control reaches using Malaise traps. The foraging activity of bats was evaluated in both treatment and control reaches using ultrasonic detectors. 4. The insect-proof cover effectively reduced the flux of emergent aquatic insects to the riparian zone adjacent to the treatment reach. Adjacent to the control reach, adult aquatic insect biomass was highest in spring, and then decreased gradually. Terrestrial insect biomass increased gradually during the summer at both treatment and control reaches. 5. Foraging activity of bats was correlated with insect abundance. In spring, foraging activity of bats at the control reach was significantly greater than at the treatment reach, and increased at both sites with increasing terrestrial insect abundance. 6. Our result suggests that the flux of aquatic insects emerging from streams is one of the most important factors affecting the distribution of riparian-foraging bats. As is the case with other riparian consumers, resource subsidies from streams can directly enhance the performance or population density of riparian-dependent bats. To conserve and manage bat populations, it is important to protect not only forest ecosystems, but also adjacent aquatic systems such as streams.     

Prey size diversity hinders biomass trophic transfer and predator size diversity promotes it in planktonic communities

Body size exerts multiple effects on plankton food-web interactions. However, the influence of size structure on trophic transfer remains poorly quantified in the field. Here, we examine how the size diversity of prey (nano-microplankton) and predators (mesozooplankton) influence trophic transfer efficiency (using biomass ratio as a proxy) in natural marine ecosystems. Our results support previous studies on single trophic levels: transfer efficiency decreases with increasing prey size diversity and is enhanced with greater predator size diversity. We further show that communities with low nano-microplankton size diversity and high mesozooplankton size diversity tend to occur in warmer environments with low nutrient concentrations, thus promoting trophic transfer to higher trophic levels in those conditions. Moreover, we reveal an interactive effect of predator and prey size diversities: the positive effect of predator size diversity becomes influential when prey size diversity is high. Mechanistically, the negative effect of prey size diversity on trophic transfer may be explained by unicellular size-based metabolic constraints as well as trade-offs between growth and predation avoidance with size, whereas increasing predator size diversity may enhance diet niche partitioning and thus promote trophic transfer. These findings provide insights into size-based theories of ecosystem functioning, with implications for ecosystem predictive models.     

The Effect of Forest Type on Benthic Macroinvertebrate Structure and Ecological Function in a Pine Plantation in the North Carolina Piedmont

We examined the impact of small-scale commercial forestry on the structure and function of 6 headwater streams in the North Carolina Piedmont. During 2001–2003 terrestrial organic matter inputs, temperature, macroinvertebrate community composition and tolerance, leaf breakdown rate, and food web structure were quantified for 2 streams draining mature stands of managed loblolly pine, 2 streams draining mature hardwood forests, and 2 streams draining 3-year-old clear cuts, which had been replanted with loblolly pine. Streams in the clear-cuts and pine plantations were bordered by a 15 m hardwood buffer. Despite differences in watershed land-use, there were no significant differences in the organic matter supply or temperature between streams draining different forest types. However, algal biomass was significantly higher in clear-cut sites than forested sites, and was also higher in hardwood sites than pine sites. Streams draining the clear-cut sites contained lower macroinvertebrate richness and diversity, and fewer intolerant species, than streams draining pine and hardwood stands. Despite the differences in macroinvertebrates community composition, there was no difference among forest types in leaf-pack breakdown rates. Analysis of δ¹⁵N and δ¹³C natural abundance of functional feeding group indicated that the shredders and predators collected from streams draining clear-cuts had a δ¹⁵N value that was enriched relative to the macroinvertebrates of streams draining pine and hardwood forests. This difference in δ¹⁵N signature appears to be the result of the incorporation of riparian grass species in the clear-cuts, which have a higher δ¹⁵N, into the diet of shredders. Pine sites had similar food webs to natural hardwood sites. Our results suggest that clear-cutting changes both the trophic dynamics and macroinvertebrate composition of low-order Piedmont streams in North Carolina despite the presence of hardwood buffers. However, large differences were not found between older pine and hardwood stands, indicating rapid recovery following re-growth of forest vegetation, when hardwood buffer strips were present.     

Non-native earthworms in riparian soils increase nitrogen flux into adjacent aquatic ecosystems

Riparian zones are an important transition between terrestrial and aquatic ecosystems, and they function in nutrient cycling and removal. Non-native earthworms invading earthworm-free areas of North America can affect nutrient cycling in upland soils and have the potential to affect it in riparian soils. We examined how the presence of earthworms can affect riparian nutrient cycling and nutrient delivery to streams. Two mesocosm experiments were conducted to determine how (1) the biomass of earthworms and (2) earthworm species can affect nutrient flux from riparian zones to nearby streams and how this flux can affect streamwater nutrients and periphyton growth. In separate experiments, riparian soil cores were amended with one of four mixed earthworm biomasses (0, 4, 10, or 23 g m(-2) ash-free dry mass) or with one of three earthworm species (Aporrectodea caliginosa, Lumbricus terrestris, L. rubellus) or no earthworm species. Riparian soil cores were coupled to artificial streams, and over a 36-day period, we measured nutrient leaching rates, in-stream nutrient concentrations, and periphyton growth. Ammonium leaching increased with increasing biomass and was greatest from the A. caliginosa treatments. Nitrate leaching increased through time and increased at a greater rate with higher biomass and from cores containing A. caliginosa. We suggest that the overall response of increased nitrate leaching [90% of total nitrogen (N)] was due to a combination of ammonium excretion and burrowing by earthworms, which increased nitrification rates. During both experiments, periphyton biomass increased through time but did not differ across treatments despite high in-stream inorganic N. Through time, in-stream phosphorus (P) concentration declined to <5 microg l(-1), and periphyton growth was likely P-limited. We conclude that activities of non-native earthworms (particularly A. caliginosa) can alter biogeochemical cycling in riparian zones, potentially reducing the N-buffering capacity of riparian zones and altering stoichiometric relationships in adjacent aquatic ecosystems.