Kyle F. Edwards

A meta‐analysis of resource pulse–consumer interactions

Resource pulses are infrequent, large-magnitude, and short-duration events of increased resource availability. They include a diverse set of extreme events in a wide range of ecosystems, but identifying general patterns among the diversity of pulsed resource phenomena in nature remains an important challenge. Here we present a meta-analysis of resource pulse-consumer interactions that addresses four key questions: (1) Which characteristics of pulsed resources best predict their effects on consumers? (2) Which characteristics of consumers best predict their responses to resource pulses? (3) How do the effects of resource pulses differ in different ecosystems? (4) What are the indirect effects of resource pulses in communities? To investigate these questions, we built a data set of diverse pulsed resource-consumer interactions from around the world, developed metrics to compare the effects of resource pulses across disparate systems, and conducted multilevel regression analyses to examine the manner in which variation in the characteristics of resource pulse- consumer interactions affects important aspects of consumer responses. Resource pulse magnitude, resource trophic level, resource pulse duration, ecosystem type and subtype, consumer response mechanisms, and consumer body mass were found to be key explanatory factors predicting the magnitude, duration, and timing of consumer responses. Larger consumers showed more persistent responses to resource pulses, and reproductive responses were more persistent than aggregative responses. Aquatic systems showed shorter temporal lags between peaks of resource availability and consumer response compared to terrestrial systems, and temporal lags were also shorter for smaller consumers compared to larger consumers. The magnitude of consumer responses relative to their resource pulses was generally smaller for the direct consumers of primary resource pulses, compared to consumers at greater trophic distances from the initial resource pulse. In specific systems, this data set showed both attenuating and amplifying indirect effects. We consider the mechanistic processes behind these patterns and their implications for the ecology of resource pulses.

The biogeography of marine plankton traits

Changes in marine plankton communities driven by environmental variability impact the marine food web and global biogeochemical cycles of carbon and other elements. To predict and assess these community shifts and their consequences, ecologists are increasingly investigating how the functional traits of plankton determine their relative fitness along environmental and biological gradients. Laboratory, field and modelling studies are adopting this trait-based approach to map the biogeography of plankton traits that underlies variations in plankton communities. Here, we review progress towards understanding the regulatory roles of several key plankton functional traits, including cell size, N2 -fixation and mixotrophy among phytoplankton, and body size, ontogeny and feeding behaviour for zooplankton. The trait biogeographical approach sheds light on what structures plankton communities in the current ocean, as well as under climate change scenarios, and also allows for finer resolution of community function because community trait composition determines the rates of significant processes, including carbon export. Although understanding of trait biogeography is growing, uncertainties remain that stem, in part, from the paucity of observations describing plankton functional traits. Thus, in addition to recommending widespread adoption of the trait-based approach, we advocate for enhanced collection, standardisation and dissemination of plankton functional trait data.

Volatile communication between plants that affects herbivory: a meta‐analysis

Volatile communication between plants causing enhanced defence has been controversial. Early studies were not replicated, and influential reviews questioned the validity of the phenomenon. We collected 48 well-replicated studies and found overall support for the hypothesis that resistance increased for individuals with damaged neighbours. Laboratory or greenhouse studies and those conducted on agricultural crops showed stronger induced resistance than field studies on undomesticated species, presumably because other variation had been reduced. A cumulative analysis revealed that early, non-replicated studies were more variable and showed less evidence for communication. Effects of habitat and plant growth form were undetectable. In most cases, the mechanisms of resistance and alternative hypotheses were not considered. There was no indication that some response variables were more likely to produce large effects. These results indicate that plants of diverse taxonomic affinities and ecological conditions become more resistant to herbivores when exposed to volatiles from damaged neighbours.

Functional traits explain phytoplankton community structure and seasonal dynamics in a marine ecosystem

A fundamental yet elusive goal of ecology is to predict the structure of communities from the environmental conditions they experience. Trait-based approaches to terrestrial plant communities have shown that functional traits can help reveal the mechanisms underlying community assembly, but such approaches have not been tested on the microbes that dominate ecosystem processes in the ocean. Here, we test whether functional traits can explain community responses to seasonal environmental fluctuation, using a time series of the phytoplankton of the English Channel. We show that interspecific variation in response to major limiting resources, light and nitrate, can be well-predicted by lab-measured traits characterising light utilisation, nitrate utilisation and maximum growth rate. As these relationships were predicted a priori, using independently measured traits, our results show that functional traits provide a strong mechanistic foundation for understanding the structure and dynamics of ecological communities.

Evidence for a three‐way trade‐off between nitrogen and phosphorus competitive abilities and cell size in phytoplankton

Trade-offs among functional traits are essential for explaining community structure and species coexistence. While two-way trade-offs have been investigated in many systems, higher-dimensional trade-offs remain largely hypothetical. Here we demonstrate a three-way trade-off between cell size and competitive abilities for nitrogen and phosphorus in marine and freshwater phytoplankton. At a given cell size, competitive abilities for N and P are negatively correlated, but as cell size increases, competitive ability decreases for both nutrients. The relative importance of the two trade-off axes appears to be environment dependent, suggesting different selective pressures: freshwater phytoplankton separate more along the N vs. P competition axis, and marine phytoplankton separate more along the nutrient competition vs. cell size axis. Our results demonstrate the multidimensional nature of key trade-offs among traits and suggest that such trade-offs may drive species interactions and structure ecological communities.

Prey diversity is associated with weaker consumer effects in a meta-analysis of benthic marine experiments

A rapidly accumulating body of research has shown that species diversity consistently affects the functioning of ecosystems. The incorporation of trophic complexity and the extension of this research to larger scales and natural ranges in species diversity remain as important challenges for understanding the true magnitude of these effects in natural systems. Here, we test whether the diversity of prey communities affects the magnitude of aggregate consumer effects. We conducted a meta-analysis of 57 consumer removal field experiments from a range of intertidal and subtidal hard substrate marine communities. We found that the richness of the prey community was the strongest predictor of the magnitude of consumer effects while controlling for habitat type, taxonomic composition, and other variables. Consumer removal increased aggregate prey abundance on average by 1200% at the lower limit of prey diversity (two species), but only 200% at the upper limit of 37 species. Importantly, compositional change was substantial at both high and low prey diversity, suggesting predation intensity did not vary with prey richness. Rather diverse prey communities appear to be more capable of maintaining abundance via compensatory responses, by containing prey species that are resistant to (or tolerant of) predators. These results suggest that the effects of species diversity on trophic interactions may scale consistently from small-scale manipulations to cross-community comparisons.

Microbial resource utilization traits and trade-offs: implications for community structure, functioning, and biogeochemical impacts at present and in the future

Trait-based approaches provide a mechanistic framework to understand and predict the structure and functioning of microbial communities. Resource utilization traits and trade-offs are among key microbial traits that describe population dynamics and competition among microbes. Several important trade-offs have been identified for prokaryotic and eukaryotic microbial taxa that define contrasting ecological strategies and contribute to species coexistence and diversity. The shape, dimensionality, and hierarchy of trade-offs may determine coexistence patterns and need to be better characterized. Laboratory measured resource utilization traits can be used to explain temporal and spatial structure and dynamics of natural microbial communities and predict biogeochemical impacts. Global environmental change can alter microbial community composition through altering resource utilization by different microbes and, consequently, may modify biogeochemical impacts of microbes.

Spatially stochastic settlement and the coexistence of benthic marine animals

For sessile organisms, dispersal and recruitment are typically spatially stochastic, but there is little understanding of how this variability scales up to influence processes such as competitive coexistence. Here we argue that coexistence of benthic marine animals is enhanced by stochastic differences between species in the spatial distribution of larval settlement. Differentiation of settlement distributions among competitors results in intraspecifically aggregated settlement, which can reduce overall interspecific competition and increase overall intraspecific competition. We test for the components of this mechanism using a pair of subtidal invertebrates, and we find that the mean interspecific effect of the dominant competitor is substantially reduced by natural settlement variability. Using a simulation parameterized with experimental data, we find that variable settlement could play an important role in long-term coexistence between these species. This mechanism may apply broadly to benthic marine communities, which can be highly diverse and typically exhibit large settlement fluctuation over a range of scales.

A three-way trade-off maintains functional diversity under variable resource supply.

The resources that organisms depend on often fluctuate over time, and a variety of common traits are thought to be adaptations to variable resource supply. To understand the trait structure of communities, it is necessary to understand the functional trade-offs that determine what trait combinations are possible and which species can persist and coexist in a given environment. We compare traits across phytoplankton species in order to test for proposed trade-offs between maximum growth rate, equilibrium competitive ability for phosphorus (P), and ability to store P. We find evidence for a three-way trade-off between these traits, and we use empirical trait covariation to parameterize a mechanistic model of competition under pulsed P supply. The model shows that different strategies are favored under different conditions of nutrient supply regime, productivity, and mortality. Furthermore, multiple strategies typically coexist, and the range of traits that persist in the model is similar to the range of traits found in real species. These results suggest that mechanistic models informed by empirical trait variation, in combination with data on the trait structure of natural communities, will play an important role in uncovering the mechanisms that underlie the diversity and structure of ecological communities.

Community trait structure in phytoplankton: seasonal dynamics from a method for sparse trait data

The distribution of functional traits in communities, and how trait distributions shift over time and space, is critical information for understanding community structure, the maintenance of diversity, and community effects on ecosystem function. It is often the case that traits tightly linked to ecological performance, such as physiological capacities, are laborious to measure and largely unknown for speciose communities; however, these traits are particularly important for unraveling the mechanistic basis of community structure. Here I develop a method combining sparse trait data with a statistical niche model to infer trait distributions for phytoplankton communities and how they vary over 10 yr in the western English Channel. I find that community-average nitrate affinity, light-limited growth rate, and maximum growth rate all show major seasonal patterns, reflecting alternate limitation by light vs. nitrogen. Trait diversity exhibits a variety of patterns distinct from community trait means, which suggests complex regulation of functional diversity. Patterns such as these are important for predicting how ocean ecosystems will respond to global change, and for developing trait-based models of emergent community structure. The statistical approach used here could be applied to any kind of organism, if it exhibits strong relationships between traits and statistical niche estimates.