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A test of geographic assignment using isotope tracers in feathers of known origin

We used feathers of known origin collected from across the breeding range of a migratory shorebird to test the use of isotope tracers for assigning breeding origins. We analyzed δD, δ 13 C, and δ 15 N in feathers from 75 mountain plover ( Charadrius montanus ) chicks sampled in 2001 and from 119 chicks sampled in 2002. We estimated parameters for continuous-response inverse regression models and for discrete-response Bayesian probability models from data for each year independently. We evaluated model predictions with both the training data and by using the alternate year as an independent test dataset. Our results provide weak support for modeling latitude and isotope values as monotonic functions of one another, especially when data are pooled over known sources of variation such as sample year or location. We were unable to make even qualitative statements, such as north versus south, about the likely origin of birds using both δD and δ 13 C in inverse regression models; results were no better than random assignment. Probability models provided better results and a more natural framework for the problem. Correct assignment rates were highest when considering all three isotopes in the probability framework, but the use of even a single isotope was better than random assignment. The method appears relatively robust to temporal effects and is most sensitive to the isotope discrimination gradients over which samples are taken. We offer that the problem of using isotope tracers to infer geographic origin is best framed as one of assignment, rather than prediction.

Oecologia

Controlling for anthropogenically induced atmospheric variation in stable carbon isotope studies

Increased use of stable isotope analysis to examine food-web dynamics, migration, transfer of nutrients, and behavior will likely result in expansion of stable isotope studies investigating human-induced global changes. Recent elevation of atmospheric CO2 concentration, related primarily to fossil fuel combustion, has reduced atmospheric CO2 ??13C (13C/12C), and this change in isotopic baseline has, in turn, reduced plant and animal tissue ??13C of terrestrial and aquatic organisms. Such depletion in CO2 ??13C and its effects on tissue ??13C may introduce bias into ??13C investigations, and if this variation is not controlled, may confound interpretation of results obtained from tissue samples collected over a temporal span. To control for this source of variation, we used a high-precision record of atmospheric CO2 ??13C from ice cores and direct atmospheric measurements to model modern change in CO2 ??13C. From this model, we estimated a correction factor that controls for atmospheric change; this correction reduces bias associated with changes in atmospheric isotopic baseline and facilitates comparison of tissue ??13C collected over multiple years. To exemplify the importance of accounting for atmospheric CO2 ??13C depletion, we applied the correction to a dataset of collagen ??13C obtained from mountain lion (Puma concolor) bone samples collected in California between 1893 and 1995. Before correction, in three of four ecoregions collagen ??13C decreased significantly concurrent with depletion of atmospheric CO2 ??13C (n ??? 32, P ??? 0.01). Application of the correction to collagen ??13C data removed trends from regions demonstrating significant declines, and measurement error associated with the correction did not add substantial variation to adjusted estimates. Controlling for long-term atmospheric variation and correcting tissue samples for changes in isotopic baseline facilitate analysis of samples that span a large temporal range. ?? Springer-Verlag 2005.

Oecologia

Incubation period and immune function: A comparative field study among coexisting birds

Developmental periods are integral components of life history strategies that can have important fitness consequences and vary enormously among organisms. However, the selection pressures and mechanisms causing variation in length of developmental periods are poorly understood. Particularly puzzling are prolonged developmental periods, because their selective advantage is unclear. Here we tested the hypotheses that immune function is stronger in species that are attacked at a higher rate by parasites and that prolonged embryonic development allows the development of this stronger immune system. Through a comparative field study among 12 coexisting passerine bird species, we show that species with higher blood parasite prevalence mounted stronger cellular immune responses than species with lower prevalence. These results provide support for the hypothesis that species facing greater selection pressure from parasites invest more in immune function. However, species with longer incubation periods mounted weaker cellular immune responses than species with shorter periods. Therefore, cellular immune responses do not support the hypothesis that longer development time enhances immunocompentence. Future studies should assess other components of the immune system and test alternative causes of variation in incubation periods among bird species. ?? Springer-Verlag 2005.

Oecologia

Spatiotemporal variability of stream habitat and movement of three species of fish

Relationships between environmental variability and movement are poorly understood, due to both their complexity and the limited ecological scope of most movement studies. We studied movements of fantail (Etheostoma flabellare), riverweed (E. podostemone), and Roanoke darters (Percina roanoka) through two stream systems during two summers. We then related movement to variability in measured habitat attributes using logistic regression and exploratory data plots. We indexed habitat conditions at both microhabitat (i.e., patches of uniform depth, velocity, and substrate) and mesohabitat (i.e., riffle and pool channel units) spatial scales, and determined how local habitat conditions were affected by landscape spatial (i.e., longitudinal position, land use) and temporal contexts. Most spatial variability in habitat conditions and fish movement was unexplained by a site's location on the landscape. Exceptions were microhabitat diversity, which was greater in the less-disturbed watershed, and riffle isolation and predator density in pools, which were greater at more-downstream sites. Habitat conditions and movement also exhibited only minor temporal variability, but the relative influences of habitat attributes on movement were quite variable over time. During the first year, movements of fantail and riverweed darters were triggered predominantly by loss of shallow microhabitats; whereas, during the second year, microhabitat diversity was more strongly related (though in opposite directions) to movement of these two species. Roanoke darters did not move in response to microhabitat-scale variables, presumably because of the species' preference for deeper microhabitats that changed little over time. Conversely, movement of all species appeared to be constrained by riffle isolation and predator density in pools, two mesohabitat-scale attributes. Relationships between environmental variability and movement depended on both the spatiotemporal scale of consideration and the ecology of the species. Future studies that integrate across scales, taxa, and life-histories are likely to provide greater insight into movement ecology than will traditional, single-season, single-species approaches. ?? 2006 Springer-Verlag.

Oecologia

Natal location influences movement and survival of a spatially structured population of snail kites

Despite the accepted importance of the need to better understand how natal location affects movement decisions and survival of animals, robust estimates of movement and survival in relation to the natal location are lacking. Our study focuses on movement and survival related to the natal location of snail kites in Florida and shows that kites, in addition to exhibiting a high level of site tenacity to breeding regions, also exhibit particular attraction to their natal region. More specifically, we found that estimates of movement from post-dispersal regions were greater toward natal regions than toward non-natal regions (differences were significant for three of four regions). We also found that estimates of natal philopatry were greater than estimates of philopatry to non-natal regions (differences were statistically significant for two of four regions). A previous study indicated an effect of natal region on juvenile survival; in this study, we show an effect of natal region on adult survival. Estimates of adult survival varied among kites that were hatched in different regions. Adults experienced mortality rates characteristic of the region occupied at the time when survival was measured, but because there is a greater probability that kites will return to their natal region than to any other regions, their survival was ultimately influenced by their natal region. In most years, kites hatched in southern regions had greater survival probabilities than did kites hatched in northern regions. However, during a multiregional drought, one of the northern regions served as a refuge from drought, and during this perturbation, survival was greater for birds hatched in the north. Our study shows that natal location may be important in influencing the ecological dynamics of kites but also highlights the importance of considering temporal variation in habitat conditions of spatially structured systems when attempting to evaluate the conservation value of habitats.

Oecologia

Comparative performance of invasive and native Celastrus species across environmental gradients

The ability to understand and predict the success of invasive plant species in their new ranges is increased when there is a sympatric native congener available for comparison. Celastrus orbiculatus (oriental bittersweet) is a liana introduced into the United States in the mid-1800s from East Asia as an ornamental plant. Its native congener, Celastrus scandens (American bittersweet), ranges from the east coast of the United States as far west as Wyoming. In the Northeastern United States, C. orbiculatus is continuing to expand its range while C. scandens appears to be in serious decline. One hypothesis for this decline is that C. scandens does not have such a wide range of ecological tolerances in the current landscape as C. orbiculatus, which seems to tolerate a greater range of resource conditions. To investigate this hypothesis, we transplanted these two species into ten sites that spanned a full range of light and soil moisture conditions to compare their establishment and performance in terms of aboveground growth (biomass and height) and mortality. After two years, C. orbiculatus showed significantly lower mortality and greater biomass across all resource conditions compared to C. scandens. In addition, C. orbiculatus preferred more mesic soil moisture conditions, while C. scandens performed better in drier soil moisture conditions. Since much of the Northeastern United States is now forested, this preference for mesic soil conditions could make it more successful than C. scandens in the region. This study shows the utility of manipulative experiments, particularly those using congeneric native species as benchmarks, for assessing the causes and predicting the course of invasions. ?? 2007 Springer-Verlag.

Oecologia

King eider use an income strategy for egg production: a case study for incorporating individual dietary variation into nutrient allocation research

The use of stored nutrients for reproduction represents an important component of life-history variation. Recent studies from several species have used stable isotopes to estimate the reliance on stored body reserves in reproduction. Such approaches rely on population-level dietary endpoints to characterize stored reserves (“capital”) and current diet (“income”). Individual variation in diet choice has so far not been incorporated in such approaches, but is crucial for assessing variation in nutrient allocation strategies. We investigated nutrient allocation to egg production in a large-bodied sea duck in northern Alaska, the king eider ( Somateria spectabilis ). We first used Bayesian isotopic mixing models to quantify at the population level the amount of endogenous carbon and nitrogen invested into egg proteins based on carbon and nitrogen isotope ratios. We then defined the isotopic signature of the current diet of every nesting female based on isotope ratios of eggshell membranes, because diets varied isotopically among individual king eiders on breeding grounds. We used these individual-based dietary isotope signals to characterize nutrient allocation for each female in the study population. At the population level, the Bayesian and the individual-based approaches yielded identical results, and showed that king eiders used an income strategy for the synthesis of egg proteins. The majority of the carbon and nitrogen in albumen (C: 86 ± 18%, N: 99 ± 1%) and the nitrogen in lipid-free yolk (90 ± 15%) were derived from food consumed on breeding grounds. Carbon in lipid-free yolk derived evenly from endogenous sources and current diet (exogenous C: 54 ± 24%), but source contribution was highly variable among individual females. These results suggest that even large-bodied birds traditionally viewed as capital breeders use exogenous nutrients for reproduction. We recommend that investigations of nutrient allocation should incorporate individual variation into mixing models to reveal intraspecific variation in reproductive strategies.

Alaska

Distribution patterns of wintering sea ducks in relation to the North Atlantic Oscillation and local environmental characteristics

Twelve species of North American sea ducks (Tribe Mergini) winter off the eastern coast of the United States and Canada. Yet, despite their seasonal proximity to urbanized areas in this region, there is limited information on patterns of wintering sea duck habitat use. It is difficult to gather information on sea ducks because of the relative inaccessibility of their offshore locations, their high degree of mobility, and their aggregated distributions. To characterize environmental conditions that affect wintering distributions, as well as their geographic ranges, we analyzed count data on five species of sea ducks (black scoters Melanitta nigra americana, surf scoters M. perspicillata, white-winged scoters M. fusca, common eiders Somateria mollissima, and long-tailed ducks Clangula hyemalis) that were collected during the Atlantic Flyway Sea Duck Survey for ten years starting in the early 1990s. We modeled count data for each species within ten-nautical-mile linear survey segments using a zero-inflated negative binomial model that included four local-scale habitat covariates (sea surface temperature, mean bottom depth, maximum bottom slope, and a variable to indicate if the segment was in a bay or not), one broad-scale covariate (the North Atlantic Oscillation), and a temporal correlation component. Our results indicate that species distributions have strong latitudinal gradients and consistency in local habitat use. The North Atlantic Oscillation was the only environmental covariate that had a significant (but variable) effect on the expected count for all five species, suggesting that broad-scale climatic conditions may be directly or indirectly important to the distributions of wintering sea ducks. Our results provide critical information on species-habitat associations, elucidate the complicated relationship between the North Atlantic Oscillation, sea surface temperature, and local sea duck abundances, and should be useful in assessing the impacts of climate change on seabirds.

Oecologia

Microbial community shifts influence patterns in tropical forest nitrogen fixation

The role of biodiversity in ecosystem function receives substantial attention, yet despite the diversity and functional relevance of microorganisms, relationships between microbial community structure and ecosystem processes remain largely unknown. We used tropical rain forest fertilization plots to directly compare the relative abundance, composition and diversity of free-living nitrogen (N)-fixer communities to in situ leaf litter N fixation rates. N fixation rates varied greatly within the landscape, and 'hotspots' of high N fixation activity were observed in both control and phosphorus (P)-fertilized plots. Compared with zones of average activity, the N fixation 'hotspots' in unfertilized plots were characterized by marked differences in N-fixer community composition and had substantially higher overall diversity. P additions increased the efficiency of N-fixer communities, resulting in elevated rates of fixation per nifH gene. Furthermore, P fertilization increased N fixation rates and N-fixer abundance, eliminated a highly novel group of N-fixers, and increased N-fixer diversity. Yet the relationships between diversity and function were not simple, and coupling rate measurements to indicators of community structure revealed a biological dynamism not apparent from process measurements alone. Taken together, these data suggest that the rain forest litter layer maintains high N fixation rates and unique N-fixing organisms and that, as observed in plant community ecology, structural shifts in N-fixing communities may partially explain significant differences in system-scale N fixation rates.

Oecologia

High mangrove density enhances surface accretion, surface elevation change, and tree survival in coastal areas susceptible to sea-level rise

Survival, growth, aboveground biomass accumulation, sediment surface elevation dynamics and nitrogen accumulation in sediments were studied in experimental treatments planted with four different densities (6.96, 3.26, 1.93 and 0.95 seedlings m −2 ) of the mangrove Rhizophora mucronata in Puttalam Lagoon, Sri Lanka. Measurements were taken over a period of 1,171 days and were compared with those from unplanted controls. Trees at the lowest density showed significantly reduced survival, whilst measures of individual tree growth did not differ among treatments. Rates of surface sediment accretion (means ± SE) were 13.0 (±1.3), 10.5 (±0.9), 8.4 (±0.3), 6.9 (±0.5) and 5.7 (±0.3) mm year −1 at planting densities of 6.96, 3.26, 1.93, 0.95, and 0 (unplanted control) seedlings m −2 , respectively, showing highly significant differences among treatments. Mean (±SE) rates of surface elevation change were much lower than rates of accretion at 2.8 (±0.2), 1.6 (±0.1), 1.1 (±0.2), 0.6 (±0.2) and −0.3 (±0.1) mm year −1 for 6.96, 3.26, 1.93, 0.95, and 0 seedlings m −2 , respectively. All planted treatments accumulated greater nitrogen concentrations in the sediment compared to the unplanted control. Sediment %N was significantly different among densities which suggests one potential causal mechanism for the facilitatory effects observed: high densities of plants potentially contribute to the accretion of greater amounts of nutrient rich sediment. While this potential process needs further research, this study demonstrated how higher densities of mangroves enhance rates of sediment accretion and surface elevation processes that may be crucial in mangrove ecosystem adaptation to sea-level rise. There was no evidence that increasing plant density evoked a trade-off with growth and survival of the planted trees. Rather, facilitatory effects enhanced survival at high densities, suggesting that managers may be able to take advantage of high plantation densities to help mitigate sea-level rise effects by encouraging positive sediment surface elevation.

Puttalam Lagoon

Population cycles are highly correlated over long time series and large spatial scales in two unrelated species: Greater sage-grouse and cottontail rabbits

Animal species across multiple taxa demonstrate multi-annual population cycles, which have long been of interest to ecologists. Correlated population cycles between species that do not share a predator–prey relationship are particularly intriguing and challenging to explain. We investigated annual population trends of greater sage-grouse ( Centrocercus urophasianus ) and cottontail rabbits ( Sylvilagus sp.) across Wyoming to explore the possibility of correlations between unrelated species, over multiple cycles, very large spatial areas, and relatively southern latitudes in terms of cycling species. We analyzed sage-grouse lek counts and annual hunter harvest indices from 1982 to 2007. We show that greater sage-grouse, currently listed as warranted but precluded under the US Endangered Species Act, and cottontails have highly correlated cycles ( r = 0.77). We explore possible mechanistic hypotheses to explain the synchronous population cycles. Our research highlights the importance of control populations in both adaptive management and impact studies. Furthermore, we demonstrate the functional value of these indices (lek counts and hunter harvest) for tracking broad-scale fluctuations in the species. This level of highly correlated long-term cycling has not previously been documented between two non-related species, over a long time-series, very large spatial scale, and within more southern latitudes.

Wyoming

Prey-mediated avoidance of an intraguild predator by its intraguild prey

Intraguild (IG) predation is an important factor influencing community structure, yet factors allowing coexistence of IG predator and IG prey are not well understood. The existence of spatial refuges for IG prey has recently been noted for their importance in allowing coexistence. However, reduction in basal prey availability might lead IG prey to leave spatial refuges for greater access to prey, leading to increased IG predation and fewer opportunities for coexistence. We determined how the availability of prey affected space-use patterns of bobcats (Lynx rufus, IG prey) in relation to coyote space-use patterns (Canis latrans, IG predators). We located animals from fall 2007 to spring 2009 and estimated bobcat home ranges and core areas seasonally. For each bobcat relocation, we determined intensity of coyote use, distance to water, small mammal biomass, and mean small mammal biomass of the home range during the season the location was collected. We built generalized linear mixed models and used Akaike Information Criteria to determine which factors best predicted bobcat space use. Coyote intensity was a primary determinant of bobcat core area location. In bobcat home ranges with abundant prey, core areas occurred where coyote use was low, but shifted to areas intensively used by coyotes when prey declined. High spatial variability in basal prey abundance allowed some bobcats to avoid coyotes while at the same time others were forced into more risky areas. Our results suggest that multiple behavioral strategies associated with spatial variation in basal prey abundance likely allow IG prey and IG predators to coexist. ?? 2010 Springer-Verlag.

Oecologia

Coexistence in streams: Do source-sink dynamics allow salamanders to persist with fish predators?

Theory suggests that source–sink dynamics can allow coexistence of intraguild predators and prey, but empirical evidence for this coexistence mechanism is limited. We used capture–mark–recapture, genetic methods, and stable isotopes to test whether source–sink dynamics promote coexistence between stream fishes, the intraguild predator, and stream salamanders ( Dicamptodon aterrimus ), the intraguild prey. Salamander populations from upstream reaches without fish were predicted to maintain or supplement sink populations in downstream reaches with fish. We found instead that downstream reaches with fish were not sinks even though fish consumed salamander larvae—apparent survival, recruitment, and population growth rate did not differ between upstream and downstream reaches. There was also no difference between upstream and downstream reaches in net emigration. We did find that D. aterrimus moved frequently along streams, but believe that this is a response to seasonal habitat changes rather than intraguild predation. Our study provides empirical evidence that local-scale mechanisms are more important than dispersal dynamics to coexistence of streams salamanders and fish. More broadly, it shows the value of empirical data on dispersal and gene flow for distinguishing between local and spatial mechanisms of coexistence.

Idaho

Species replacement by a nonnative salmonid alters ecosystem function by reducing prey subsidies that support riparian spiders

Replacement of a native species by a nonnative can have strong effects on ecosystem function, such as altering nutrient cycling or disturbance frequency. Replacements may cause shifts in ecosystem function because nonnatives establish at different biomass, or because they differ from native species in traits like foraging behavior. However, no studies have compared effects of wholesale replacement of a native by a nonnative species on subsidies that support consumers in adjacent habitats, nor quantified the magnitude of these effects. We examined whether streams invaded by nonnative brook trout ( Salvelinus fontinalis ) in two regions of the Rocky Mountains, USA, produced fewer emerging adult aquatic insects compared to paired streams with native cutthroat trout ( Oncorhynchus clarkii ), and whether riparian spiders that depend on these prey were less abundant along streams with lower total insect emergence. As predicted, emergence density was 36% lower from streams with the nonnative fish. Biomass of brook trout was higher than the cutthroat trout they replaced, but even after accounting for this difference, emergence was 24% lower from brook trout streams. More riparian spiders were counted along streams with greater total emergence across the water surface. Based on these results, we predicted that brook trout replacement would result in 6–20% fewer spiders in the two regions. When brook trout replace cutthroat trout, they reduce cross-habitat resource subsidies and alter ecosystem function in stream-riparian food webs, not only owing to increased biomass but also because traits apparently differ from native cutthroat trout.

Oecologia

δ 15 N constraints on long-term nitrogen balances in temperate forests

Biogeochemical theory emphasizes nitrogen (N) limitation and the many factors that can restrict N accumulation in temperate forests, yet lacks a working model of conditions that can promote naturally high N accumulation. We used a dynamic simulation model of ecosystem N and δ 15 N to evaluate which combination of N input and loss pathways could produce a range of high ecosystem N contents characteristic of forests in the Oregon Coast Range. Total ecosystem N at nine study sites ranged from 8,788 to 22,667 kg ha −1 and carbon (C) ranged from 188 to 460 Mg ha −1 , with highest values near the coast. Ecosystem δ 15 N displayed a curvilinear relationship with ecosystem N content, and largely reflected mineral soil, which accounted for 96–98% of total ecosystem N. Model simulations of ecosystem N balances parameterized with field rates of N leaching required long-term average N inputs that exceed atmospheric deposition and asymbiotic and epiphytic N 2 -fixation, and that were consistent with cycles of post-fire N 2 -fixation by early-successional red alder. Soil water δ 15 NO 3 − patterns suggested a shift in relative N losses from denitrification to nitrate leaching as N accumulated, and simulations identified nitrate leaching as the primary N loss pathway that constrains maximum N accumulation. Whereas current theory emphasizes constraints on biological N 2 -fixation and disturbance-mediated N losses as factors that limit N accumulation in temperate forests, our results suggest that wildfire can foster substantial long-term N accumulation in ecosystems that are colonized by symbiotic N 2 -fixing vegetation.

Oecologia

Genetic diversity and species diversity of stream fishes covary across a land-use gradient

Genetic diversity and species diversity are expected to covary according to area and isolation, but may not always covary with environmental heterogeneity. In this study, we examined how patterns of genetic and species diversity in stream fishes correspond to local and regional environmental conditions. To do so, we compared population size, genetic diversity and divergence in central stonerollers ( Campostoma anomalum ) to measures of species diversity and turnover in stream fish assemblages among similarly sized watersheds across an agriculture–forest land-use gradient in the Little Miami River basin (Ohio, USA). Significant correlations were found in many, but not all, pair-wise comparisons. Allelic richness and species richness were strongly correlated, for example, but diversity measures based on allele frequencies and assemblage structure were not. In-stream conditions related to agricultural land use were identified as significant predictors of genetic diversity and species diversity. Comparisons to population size indicate, however, that genetic diversity and species diversity are not necessarily independent and that variation also corresponds to watershed location and glaciation history in the drainage basin. Our findings demonstrate that genetic diversity and species diversity can covary in stream fish assemblages, and illustrate the potential importance of scaling observations to capture responses to hierarchical environmental variation. More comparisons according to life history variation could further improve understanding of conditions that give rise to parallel variation in genetic diversity and species diversity, which in turn could improve diagnosis of anthropogenic influences on aquatic ecosystems.

Ohio

Foraging segregation and genetic divergence between geographically proximate colonies of a highly mobile seabird

Foraging segregation may play an important role in the maintenance of animal diversity, and is a proposed mechanism for promoting genetic divergence within seabird species. However, little information exists regarding its presence among seabird populations. We investigated genetic and foraging divergence between two colonies of endangered Hawaiian petrels ( Pterodroma sandwichensis ) nesting on the islands of Hawaii and Kauai using the mitochondrial Cytochrome b gene and carbon, nitrogen and hydrogen isotope values (δ 13 C, δ 15 N and δD, respectively) of feathers. Genetic analyses revealed strong differentiation between colonies on Hawaii and Kauai, with Φ ST = 0.50 ( p < 0.0001). Coalescent-based analyses gave estimates of <1 migration event per 1,000 generations. Hatch-year birds from Kauai had significantly lower δ 13 C and δ 15 N values than those from Hawaii. This is consistent with Kauai birds provisioning chicks with prey derived from near or north of the Hawaiian Islands, and Hawaii birds provisioning young with prey from regions of the equatorial Pacific characterized by elevated δ 15 N values at the food web base. δ 15 N values of Kauai and Hawaii adults differed significantly, indicating additional foraging segregation during molt. Feather δD varied from −69 to 53‰. This variation cannot be related solely to an isotopically homogeneous ocean water source or evaporative water loss. Instead, we propose the involvement of salt gland excretion. Our data demonstrate the presence of foraging segregation between proximately nesting seabird populations, despite high species mobility. This ecological diversity may facilitate population coexistence, and its preservation should be a focus of conservation strategies.

Hawaii

Species traits and environmental conditions govern the relationship between biodiversity effects across trophic levels

Changing environments can have divergent effects on biodiversity–ecosystem function relationships at alternating trophic levels. Freshwater mussels fertilize stream foodwebs through nutrient excretion, and mussel species-specific excretion rates depend on environmental conditions. We asked how differences in mussel diversity in varying environments influence the dynamics between primary producers and consumers. We conducted field experiments manipulating mussel richness under summer (low flow, high temperature) and fall (moderate flow and temperature) conditions, measured nutrient limitation, algal biomass and grazing chironomid abundance, and analyzed the data with non-transgressive overyielding and tripartite biodiversity partitioning analyses. Algal biomass and chironomid abundance were best explained by trait-independent complementarity among mussel species, but the relationship between biodiversity effects across trophic levels (algae and grazers) depended on seasonal differences in mussel species’ trait expression (nutrient excretion and activity level). Both species identity and overall diversity effects were related to the magnitude of nutrient limitation. Our results demonstrate that biodiversity of a resource-provisioning (nutrients and habitat) group of species influences foodweb dynamics and that understanding species traits and environmental context are important for interpreting biodiversity experiments.

Oecologia