Laurie B. Marczak

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.

THE MUTUALISM–PARASITISM CONTINUUM IN ECTOMYCORRHIZAS: A QUANTITATIVE ASSESSMENT USING META‐ANALYSIS

Context dependency is deemed to position the outcomes of species interactions along a continuum of mutualism to parasitism. Thus, it is imperative to understand which factors determine where a particular interspecific interaction falls along the continuum. Over the past 20 years research on the ectomycorrhizal symbiosis has resulted in sufficient independent studies to now generalize about the factors and mechanisms that affect host response to ectomycorrhizas. Using meta-analysis we quantitatively evaluated the role of biotic (partner identity and colonization levels of ectomycorrhizal fungi) and abiotic (phosphorus levels) factors in determining host biomass, height, and shoot:root responses to ectomycorrhizal associations. On average, seedlings across multiple host genera increased in total biomass when inoculated with ectomycorrhizal fungi regardless of the identity of the fungal associate; host genera differed in the magnitude of response for both total biomass and shoot:root ratio. Association with different fungal genera modified only host allocation of biomass to shoots and roots. Neither level of colonization on inoculated seedlings nor the level of contamination on control seedlings relative to colonization levels by target fungi on inoculated seedlings was important in explaining variation in effect sizes for any growth response. None of our proposed factors (identity of partners, colonization level, magnitude of contamination, or duration of association) explained variation in effect sizes for shoot height, although in general seedlings were taller when inoculated with ectomycorrhizal fungi. Phosphorus additions did not influence effect sizes. Although the general trend across studies was for a positive response of hosts to ectomycorrhizal inoculation, publication bias and methodological issues effectively reduce and distort the spectrum on which we evaluate host responses to ectomycorrhizal inoculation. Our results indicate that the variation in ectomycorrhizal fungi perceived by the host may be of a discrete (presence/absence of ectomycorrhizal fungi) rather than continuous nature (variation in identity or abundance of ectomycorrhizal fungi).

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.

Are forested buffers an effective conservation strategy for riparian fauna? An assessment using meta-analysis

Historically, forested riparian buffers have been created to provide protection for aquatic organisms and aquatic ecosystem functions. Increasingly, new and existing riparian buffers are being used also to meet terrestrial conservation requirements. To test the effectiveness of riparian buffers for conserving terrestrial fauna, we conducted a meta-analysis using published data from 397 comparisons of species abundance in riparian buffers and unharvested (reference) riparian sites. The response of terrestrial species to riparian buffers was not consistent between taxonomic groups; bird and arthropod abundances were significantly greater in buffers relative to unharvested areas, whereas amphibian abundance decreased. Edge-preferring species were more abundant in buffer sites than reference sites, whereas species associated with interior habitat were not significantly different in abundance. The degree of buffer effect on animal abundance was unrelated to buffer width; wider buffers did not result in greater similarity between reference and buffer sites. However, responses to buffer treatment were more variable in buffers <50 m wide, a commonly prescribed width in many management plans. Our results indicate that current buffer prescriptions do not maintain most terrestrial organisms in buffer strips at levels comparable to undisturbed sites.

Growth and development rates in a riparian spider are altered by asynchrony between the timing and amount of a resource subsidy

Rapid growth in response to increased prey abundance may be induced by environmental variability associated with resource subsidies. Spiders living in riparian areas are subject to frequent, episodic bursts of aquatic prey (subsidies). These periods of high resource abundance may occur at different points in recipient consumers’ development through variation in emergence patterns of prey between years or across a landscape. We examine how variable timing of subsidy abundance intersects with life history scheduling to produce different growth and development outcomes for individuals within a population. Through a series of controlled feeding experiments, we tested the hypotheses that the spider Tetragnatha versicolor: (1) exhibits compensatory growth in response to subsidy variability, (2) that rapid increases in mass may result in a greater risk of mortality, and (3) that the timing of subsidy resources relative to the development schedule of this spider may produce different outcomes for individual growth patterns and adult condition. Spiders fed at very high rates grew fastest but also showed evidence of increased mortality risk during moulting. T. versicolor is capable of exhibiting strong growth compensation—individuals suffering initial growth restriction were able to catch up completely with animals on a constant diet utilising the same amount of food. Spiders that received an early pulse of resources (simulating an early arrival of an aquatic insect subsidy to riparian forests) did worse on all measures of development and fitness than spiders that received either a constant supply of food or a late pulse of resources. Importantly, receiving large amounts of food early in life appears to actually confer relative disadvantages in terms of later performance compared with receiving subsidies later in development. Subsidies may provide greater benefits to individuals or age cohorts encountering this resource abundance closer to the onset of reproductive efforts than subsidies arriving early in development.

Effects of an Omnivorous Katydid, Salinity, and Nutrients on a Planthopper-Spartina Food Web

Top–down and bottom–up effects interact to structure communities, especially in salt marshes, which contain strong gradients in bottom–up drivers such as salinity and nutrients. How omnivorous consumers respond to variation in prey availability and plant quality is poorly understood. We used a mesocosm experiment to examine how salinity, nutrients, an omnivore (the katydid Orchelimum fidicinium) and an herbivore (the planthopper Prokelisia spp.) interacted to structure a simplified salt marsh food web based on the marsh grass Spartina alterniflora. Bottom–up effects were strong, with both salinity and nutrients decreasing leaf C/N and increasing Prokelisia abundance. Top–down effects on plants were also strong, with both the herbivore and the omnivore affecting S. alterniflora traits and growth, especially when nutrients or salt were added. In contrast, top–down control by Orchelimum of Prokelisia was independent of bottom–up conditions. Orchelimum grew best on a diet containing both Spartina and Prokelisia, and in contrast to a sympatric omnivorous crab, did not shift to an animal-based diet when prey were present, suggesting that it is constrained to consume a mixed diet. These results suggest that the trophic effects of omnivores depend on omnivore behavior, dietary constraints, and ability to suppress lower trophic levels, and that omnivorous katydids may play a previously unrecognized role in salt marsh food webs.

The role of stranding and inundation on leaf litterdecomposition in headwater streams

Abstract Discharge-driven shifts in the wetted area of streams can modify the amount of leaf litter resources available to stream consumers as well as the physical conditions to which leaf litter is exposed. The consequences of this continual movement from wet to dry storage for rates of organic matter processing and resource availability to benthic communities are poorly understood. We used a 30-day field experiment during the period of maximum stream contraction to examine the effects of stranding on black cottonwood (Populus trichocarpa) leaf litter decomposition rates and associated changes in microbial respiration in a forested stream in western Montana. Leaf litter was enclosed in both coarse and fine mesh bags and moved from the wetted area of the stream to the stream bank in six treatments designed to mimic a gradient of dry exposure due to stranding. We also measured existing accumulations of organic material in quadrats placed in both wet and dry areas of the stream. The total storage of litter resources (ash-free dry mass, g m2) retained on dry stream banks increased steadily as stream flow decreased, resulting from reductions in wetted width and continuous inputs from terrestrial zones. In contrast, total mass of stored litter submerged in the stream channel remained relatively constant. Leaf decomposition rates increased as a function of time inundated and were fastest in the presence of macroinvertebrates. Our results suggest that prolonged stranding can alter fundamental processes and energy pathways in stream food webs by shifting pools of resources from the active channel to dry storage on riverbanks where decomposition is driven primarily by microbial processes. Since the length of time that leaf litter is inundated prior to stranding alters decomposition rates, changes in stream hydrograph variability (as a consequence of land management practices or incipient climate change) has the potential to alter energy flow through stream systems. In particular, dry storage may function as a type of ‘temporal subsidy’ for stream organisms particularly if slowly decomposing stranded leaf litter is re-entrained during periods when in-stream detrital resources are otherwise scarce.

Importance of local vs. geographic variation in salt marsh plant quality for arthropod herbivore communities

Summary 1. An important recent advance in food web ecology has been the application of theory regarding spatial gradients to studies of the factors that affect animal population dynamics. Building on extensive studies of the Spartina alterniflora food web at the local scale, we hypothesized that geographic variation in S. alterniflora quality is an important bottom-up control on food web structure and that geographic variation in S. alterniflora quality would interact with the presence of predators and top omnivores to mediate herbivore densities. 2. We employed a four-factor fully crossed experiment in which we (i) collected plants from highand low-latitude locations and grew them in a common garden and varied (ii) plant fertilization status (mimicking the plant quality differences due to marsh elevation), (iii) mesopredator density and (iv) omnivore density. 3. Our results suggest that the single most important factor mediating insect herbivore densities is local variation in plant quality – induced in our experiment by fertilization and demonstrated repeatedly as a consequence of marsh elevation. 4. Top-down effects were generally weak and in those cases where predators did exert a significant suppressing effect on herbivores, that impact was itself mediated by host-plant characteristics. 5. Finally, despite observed variation in plant quality with latitude, and the separately measurable effects of this variation on herbivores, geographic-scale variation in plant quality was overwhelmed by local conditions in our experiments. 6. Synthesis. We suggest that a first-order understanding of variation across large latitudinal ranges in the Spartina alterniflora arthropod food web must begin with local variation in plant quality, which provides strong bottom-up forcing to herbivore populations. A second-order understanding of the arthropod food web should consider the role of predation in controlling herbivores feeding on lowquality plants. Finally, while latitudinal variation in plant quality probably explains some variation in herbivore densities, it is probably more of a response to herbivore pressure than a driver of the herbivore dynamics. Although extrapolating from local to geographic scales presents multiple challenges, it is an essential task in order for us to develop an understanding that is general rather than site-specific.

Geographic variation in plant community structure of salt marshes: species, functional and phylogenetic perspectives.

In general, community similarity is thought to decay with distance; however, this view may be complicated by the relative roles of different ecological processes at different geographical scales, and by the compositional perspective (e.g. species, functional group and phylogenetic lineage) used. Coastal salt marshes are widely distributed worldwide, but no studies have explicitly examined variation in salt marsh plant community composition across geographical scales, and from species, functional and phylogenetic perspectives. Based on studies in other ecosystems, we hypothesized that, in coastal salt marshes, community turnover would be more rapid at local versus larger geographical scales; and that community turnover patterns would diverge among compositional perspectives, with a greater distance decay at the species level than at the functional or phylogenetic levels. We tested these hypotheses in salt marshes of two regions: The southern Atlantic and Gulf Coasts of the United States. We examined the characteristics of plant community composition at each salt marsh site, how community similarity decayed with distance within individual salt marshes versus among sites in each region, and how community similarity differed among regions, using species, functional and phylogenetic perspectives. We found that results from the three compositional perspectives generally showed similar patterns: there was strong variation in community composition within individual salt marsh sites across elevation; in contrast, community similarity decayed with distance four to five orders of magnitude more slowly across sites within each region. Overall, community dissimilarity of salt marshes was lowest on the southern Atlantic Coast, intermediate on the Gulf Coast, and highest between the two regions. Our results indicated that local gradients are relatively more important than regional processes in structuring coastal salt marsh communities. Our results also suggested that in ecosystems with low species diversity, functional and phylogenetic approaches may not provide additional insight over a species-based approach.