Christopher M. Swan

Loss of foundation species: consequences for the structure and dynamics of forested ecosystems

In many forested ecosystems, the architecture and functional ecology of certain tree species define forest structure and their species-specific traits control ecosystem dynamics. Such foundation tree species are declining throughout the world due to introductions and outbreaks of pests and pathogens, selective removal of individual taxa, and over-harvesting. Through a series of case studies, we show that the loss of foundation tree species changes the local environment on which a variety of other species depend; how this disrupts fundamental ecosystem processes, including rates of decomposition, nutrient fluxes, carbon sequestration, and energy flow; and dramatically alters the dynamics of associated aquatic ecosystems. Forests in which dynamics are controlled by one or a few foundation species appear to be dominated by a small number of strong interactions and may be highly susceptible to alternating between stable states following even small perturbations. The ongoing decline of many foundation species provides a set of important, albeit unfortunate, opportunities to develop the research tools, models, and metrics needed to identify foundation species, anticipate the cascade of immediate, short- and long-term changes in ecosystem structure and function that will follow from their loss, and provide options for remedial conservation and management.

Diversity meets decomposition

Over 100 gigatons of terrestrial plant biomass are produced globally each year. Ninety percent of this biomass escapes herbivory and enters the dead organic matter pool, thus supporting complex detritus-based food webs that determine the critical balance between carbon mineralization and sequestration. How will changes in biodiversity affect this vital component of ecosystem functioning? Based on our analysis of concepts and experiments of leaf decomposition in forest floors and streams, we suggest that changes in species diversity within and across trophic levels can significantly alter decomposition. This happens through various mechanisms that are broadly similar in forest floors and streams. Differences in diversity effects between these systems relate to divergent habitat conditions and evolutionary trajectories of aquatic and terrestrial decomposers.

Dendritic network structure constrains metacommunity properties in riverine ecosystems

1. Increasingly, ecologists conceptualize local communities as connected to a regional species pool rather than as isolated entities. By this paradigm, community structure is determined through the relative strengths of dispersal-driven regional effects and local environmental factors. However, despite explicit incorporation of dispersal, metacommunity models and frameworks often fail to capture the realities of natural systems by not accounting for the configuration of space within which organisms disperse. This shortcoming may be of particular consequence in riverine networks which consist of linearly -arranged, hierarchical, branching habitat elements. Our goal was to understand how constraints of network connectivity in riverine systems change the relative importance of local vs. regional factors in structuring communities. 2. We hypothesized that communities in more isolated headwaters of riverine networks would be structured by local forces, while mainstem sections would be structured by both local and regional processes. We examined these hypotheses using a spatially explicit regional analysis of riverine macroinvertebrate communities, focusing on change in community similarity with distance between local communities [i.e., distance-decay relationships; (DDRs)], and the change in environmental similarity with distance. Strong DDRs frequently indicate dispersal-driven dynamics. 3. There was no evidence of a DDR in headwater communities, supporting our hypothesis that dispersal is a weak structuring force. Furthermore, a positive relationship between community similarity and environmental similarity supported dynamics driven by local environmental factors (i.e., species sorting). In mainstem habitats, significant DDRs and community x environment similarity relationships suggested both dispersal-driven and environmental constraints on local community structure (i.e., mass effects). 4. We used species traits to compare communities characterized by low vs. high dispersal taxa. In headwaters, neither strength nor mode (in-network vs. out of network) of dispersal changed our results. However, outcomes in mainstems changed substantially with both dispersal mode and strength, further supporting the hypothesis that regional forces drive community dynamics in mainstems. 5. Our findings demonstrate that the balance of local and regional effects changes depending on location within riverine network with local (environmental) factors dictating community structure in headwaters, and regional (dispersal driven) forces dominating in mainstems.

The influence of substrate heterogeneity on biofilm metabolism in a stream ecosystem

Simplification of natural habitats is a growing global concern demanding that ecologists better understand how habitat heterogeneity influences the structure and functioning of ecosystems. While there is extensive evidence that physical habitat hetero- geneity affects the structure of biotic communities (i.e., organismal abundance, distribution, diversity, etc.), ecologists know little about how variability in physical conditions within habitats regulates ecological processes that are important for the functioning of an eco- system. We performed a field experiment to assess the effects of geomorphic heterogeneity (i.e., variation in substrate size) on rates of benthic productivity and respiration at the scale of whole riffle habitats in a stream ecosystem. While holding median sizes constant, we manipulated variation in the size of stream bed sediments in replicate riffles to create two treatments representing increased and decreased levels of physical habitat heterogeneity relative to natural conditions in the stream. Physical habitat heterogeneity had an immediate and significant impact on the primary productivity of stream algae and on the respiration of the benthic biofilm. The rates of both ecological processes were elevated in the high- heterogeneity riffles, probably as a result of quantified alterations to near-bed flow velocity and turbulence intensity. Results presented here provide support for the widely held, but largely untested, assumption that physical habitat heterogeneity exhibits control over eco- system-level processes, and it suggests that human-induced simplification of habitats may indeed be altering the functioning of ecosystems.

Leaf diversity alters litter breakdown in a Piedmont stream

Abstract Work in terrestrial ecosystems has suggested that the breakdown rate of leaf litter may not change predictably with increasing plant species richness. Indeed, it may be that certain combinations of leaf species demonstrate significant non-additive effects on breakdown rates, mediated by the presence of a single key species. Such effects have not been explored in running-water ecosystems despite the strong interest in the conservation and restoration of riparian systems. We documented the magnitude and species composition of leaf litter inputs, the species richness and composition of leaf litter on the streambed, and estimated the breakdown rate of mixed litter in both the summer and autumn in a warmwater stream. We found that leaf species richness of litter packs on the streambed varied from 1 to 11 species, and leaf species composition reflected the composition of litter inputs from May through November. We did not find a general relationship between breakdown rate and leaf litter species richness. However, we did find a strong effect of species composition of leaf packs on the breakdown rate during the summer. Overall, breakdown rates of mixed-species leaf packs were non-additive during the summer, but very predictable in the autumn. In particular, leaf mixtures containing American sycamore always exhibited slower breakdown rates than predicted in summer. One explanation for the discrepancy between summer and autumn results may be decreased temperature in autumn; reduced temperature may have slowed breakdown rates across treatments to the extent that any non-additive effects found in summer were masked by the effect of temperature. Given the importance of detritus to stream food webs, the simplification of plant assemblages along intact or restored streams may have important implications for stream ecosystems.

Metacommunity theory as a multispecies, multiscale framework for studying the influence of river network structure on riverine communities and ecosystems

Abstract Explaining the mechanisms underlying patterns of species diversity and composition in riverine networks is challenging. Historically, community ecologists have conceived of communities as largely isolated entities and have focused on local environmental factors and interspecific interactions as the major forces determining species composition. However, stream ecologists have long embraced a multiscale approach to studying riverine ecosystems and have studied both local factors and larger-scale regional factors, such as dispersal and disturbance. River networks exhibit a dendritic spatial structure that can constrain aquatic organisms when their dispersal is influenced by or confined to the river network. We contend that the principles of metacommunity theory would help stream ecologists to understand how the complex spatial structure of river networks mediates the relative influences of local and regional control on species composition. From a basic ecological perspective, the concept is attractive because new evidence suggests that the importance of regional processes (dispersal) depends on spatial structure of habitat and on connection to the regional species pool. The role of local factors relative to regional factors will vary with spatial position in a river network. From an applied perspective, the long-standing view in ecology that local community composition is an indicator of habitat quality may not be uniformly applicable across a river network, but the strength of such bioassessment approaches probably will depend on spatial position in the network. The principles of metacommunity theory are broadly applicable across taxa and systems but seem of particular consequence to stream ecology given the unique spatial structure of riverine systems. By explicitly embracing processes at multiple spatial scales, metacommunity theory provides a foundation on which to build a richer understanding of stream communities.

Composition of speciose leaf litter alters stream detritivore growth, feeding activity and leaf breakdown

Leaf litter derived from riparian trees can control secondary production of detritivores in forested streams. Species-rich assemblages of leaf litter reflect riparian plant species richness and represent a heterogeneous resource for stream consumers. Such variation in resource quality may alter consumer growth and thus the feedback on leaf breakdown rate via changes in feeding activity. To assess the consequences of this type of resource heterogeneity on both consumer growth and subsequent litter breakdown, we performed a laboratory experiment where we offered a leaf-shredding stream detritivore (the stonefly Tallaperla maria, Peltoperlidae) ten treatments of either single- or mixed-species leaf litter. We measured consumer growth rate, breakdown rate and feeding activity both with and without consumers for each treatment and showed that all three variables responded to speciose leaf litter. However, the number of leaf species was not responsible for these results, but leaf species composition explained the apparent non-additive effects. T. maria growth responded both positively and negatively to litter composition, and growth on mixed-litter could not always be predicted by averaging estimates of growth in single-species treatments. Furthermore, breakdown and feeding rates in mixed litter treatments could not always be predicted from estimates of single-species rates. Given that species richness and composition of senesced leaves in streams reflects riparian plant species richness, in-stream secondary production of detritivores and organic matter dynamics may be related to species loss of trees in the riparian zone. Loss of key species may be more critical to maintaining such processes than species richness per se.

Forecasting functional implications of global changes in riparian plant communities

Riparian ecosystems support mosaics of terrestrial and aquatic plant species that enhance regional biodiversity and provide important ecosystem services to humans. Species composition and the distribution of functional traits – traits that define species in terms of their ecological roles – within riparian plant communities are rapidly changing in response to various global change drivers. Here, we present a conceptual framework illustrating how changes in dependent wildlife communities and ecosystem processes can be predicted by examining shifts in riparian plant functional trait diversity and redundancy (overlap). Three widespread examples of altered riparian plant composition are: shifts in the dominance of deciduous and coniferous species; increases in drought-tolerant species; and the increasing global distribution of plantation and crop species. Changes in the diversity and distribution of critical plant functional traits influence terrestrial and aquatic food webs, organic matter production and pro...

Exposure and Nontarget Effects of Transgenic Bt Corn Debris in Streams

ABSTRACT Corn (Zea mays L.) transformed with a gene from the bacterium Bacillus thuringiensis (Bt) comprises 49% of all corn in the United States. The input of senesced corn tissue expressing the Bt gene may impact stream-inhabiting invertebrates that process plant debris, especially trichopteran species related to the target group of lepidopteran pests. Our goal was to assess risk associated with transgenic corn debris entering streams. First, we show the input of corn tissue after harvest was extended over months in a stream. Second, using laboratory bioassays based on European corn borer [Ostrinia nubilalis (Hübner)], we found no bioactivity of Cry1Ab protein in senesced corn tissue after 2 wk of exposure to terrestrial or aquatic environments. Third, we show that Bt near-isolines modify growth and survivorship of some species of invertebrates. Of the four nontarget invertebrate species fed Bt near-isolines, growth of two closely related trichopterans was not negatively affected, whereas a tipulid crane fly exhibited reduced growth rates, and an isopod exhibited reduced growth and survivorship on the Cry1Ab near-isoline but not on the stacked Cry1Ab + Cry3Bb1 near-isoline. Because of lack of evidence of bioactivity of Bt after 2 wk and because of lack of nontarget effects on the stacked near-isoline, we suggest that tissue-mediated differences, and not the presence of the Cry1Ab protein, caused the different responses among the species. Overall, our results provide evidence that adverse effects to aquatic nontarget shredders involve complex interactions arising from plant genetics and environment that cannot be ascribed to the presence of Cry1Ab proteins.

Preferential feeding by an aquatic consumer mediates non-additive decomposition of speciose leaf litter

Forest soils and streams receive substantial inputs of detritus from deciduous vegetation. Decay of this material is a critical ecosystem process, recycling nutrients and supporting detrital-based food webs, and has been attributed, in part, to leaf litter species composition. However, research on why speciose leaf litter should degrade differently has relied on a bottom-up approach, embracing interspecific variation in litter chemistry. We hypothesized that preferential feeding by an aquatic detritivore interacts with species-specific leaf palatability and slows decay of speciose leaf litter. We addressed this by offering four single- and mixed-species leaf resources to field densities of a leaf-shredding consumer. Mixing leaf species resulted in slower total leaf decomposition. Decreases in mixed-species decomposition was partly explained by preferential feeding by the consumers in one case, but the lack of preferential feeding in other mixtures suggested an interactive effect of feeding and microbial degradation. Loss of riparian tree biodiversity may have implications for in-stream consumer-resource interactions.