Evidence of climate-driven selection on tree traits and trait plasticity across the climatic range of a riparian foundation species.

Selection on quantitative traits by heterogeneous climatic conditions can lead to substantial trait variation across a species range. In the context of rapidly changing environments, however, it is equally important to understand selection on trait plasticity. To evaluate the role of selection in driving divergences in traits and their associated plasticities within a widespread species, we compared molecular and quantitative trait variation in Populus fremontii (Fremont cottonwood), a foundation riparian distributed throughout Arizona. Using SNP data and genotypes from 16 populations reciprocally planted in three common gardens, we first performed QST -FST analyses to detect selection on traits and trait plasticity. We then explored the environmental drivers of selection using trait-climate and plasticity-climate regressions. Three major findings emerged: (1) There was significant genetic variation in traits expressed in each of the common gardens and in the phenotypic plasticity of traits across gardens, both of which were heritable. (2) Based on QST -FST comparisons, there was evidence of selection in all traits measured; however, this result varied from no effect in one garden to highly significant in another, indicating that detection of past selection is environmentally dependent. We also found strong evidence of divergent selection on plasticity across environments for two traits. (3) Traits and/or their plasticity were often correlated with population source climate (R2 up to .77 and .66, respectively). These results suggest that steep climate gradients across the Southwest have played a major role in shaping the evolution of divergent phenotypic responses in populations and genotypes now experiencing climate change.

Genetic divergence along a climate gradient shapes chemical plasticity of a foundation tree species to both changing climate and herbivore damage.

Climate change is threatening the persistence of many tree species via independent and interactive effects on abiotic and biotic conditions. In addition, changes in temperature, precipitation, and insect attacks can alter the traits of these trees, disrupting communities and ecosystems. For foundation species such as Populus, phytochemical traits are key mechanisms linking trees with their environment and are likely jointly determined by interactive effects of genetic divergence and variable environments throughout their geographic range. Using reciprocal Fremont cottonwood (Populus fremontii) common gardens along a steep climatic gradient, we explored how environment (garden climate and simulated herbivore damage) and genetics (tree provenance and genotype) affect both foliar chemical traits and the plasticity of these traits. We found that (1) Constitutive and plastic chemical responses to changes in garden climate and damage varied among defense compounds, structural compounds, and leaf nitrogen. (2) For both defense and structural compounds, plastic responses to different garden climates depended on the climate in which a population or genotype originated. Specifically, trees originating from cool provenances showed higher defense plasticity in response to climate changes than trees from warmer provenances. (3) Trees from cool provenances growing in cool garden conditions expressed the lowest constitutive defense levels but the strongest induced (plastic) defenses in response to damage. (4) The combination of hot garden conditions and simulated herbivory switched the strategy used by these genotypes, increasing constitutive defenses but erasing the capacity for induction after damage. Because Fremont cottonwood chemistry plays a major role in shaping riparian communities and ecosystems, the effects of changes in phytochemical traits can be wide reaching. As the southwestern US is confronted with warming temperatures and insect outbreaks, these results improve our capacity to predict ecosystem consequences of climate change and inform selection of tree genotypes for conservation and restoration purposes.

Dominance determines fish community biomass in a temperate seagrass ecosystem.

Biodiversity and ecosystem function are often correlated, but there are multiple hypotheses about the mechanisms underlying this relationship. Ecosystem functions such as primary or secondary production may be maximized by species richness, evenness in species abundances, or the presence or dominance of species with certain traits. Here, we combine surveys of natural fish communities (conducted in July and August 2016) with morphological trait data to examine relationships between biodiversity and ecosystem function (quantified as fish community biomass) across 14 subtidal eelgrass meadows in the Northeast Pacific (54°N, 130°W). We employ both taxonomic and functional trait measures of diversity to investigate whether ecosystem function is best predicted by species diversity (complementarity hypothesis) or by the presence or dominance of species with particular trait values (selection or dominance hypotheses). After controlling for environmental variation, we find that fish community biomass is maximized when taxonomic richness and functional evenness are low, and in communities dominated by species with particular trait values, specifically those associated with benthic habitats and prey capture. While previous work on fish communities has found that species richness is often positively correlated with ecosystem function, our results instead highlight the capacity for regionally prevalent and locally dominant species to drive ecosystem function in moderately diverse communities. We discuss these alternate links between community composition and ecosystem function and consider their divergent implications for ecosystem valuation and conservation prioritization.

Plastic responses to hot temperatures homogenize riparian leaf litter, speed decomposition, and reduce detritivores.

Efforts to maintain the function of critical ecosystems under climate change often begin with foundation species. In the southwestern United States, cottonwood trees support diverse communities in riparian ecosystems that are threatened by rising temperatures. Genetic variation within cottonwoods shapes communities and ecosystems, but these effects may be modified by phenotypic plasticity, where genotype traits change in response to environmental conditions. Here, we investigated plasticity in Fremont cottonwood (Populus fremontii) leaf litter traits as well as the consequences of plasticity for riparian ecosystems. We used three common gardens each planted with genotypes from six genetically divergent populations spanning a 12°C temperature gradient, and a decomposition experiment in a common stream environment. We found that leaf litter area, specific leaf area, and carbon to nitrogen ratio (C:N) were determined by interactions between genetics and growing environment, as was the subsequent rate of litter decomposition. Most of the genetic variation in leaf litter traits appeared among rather than within source populations with distinct climate histories. Source populations from hotter climates generally produced litter that decomposed more quickly, but plasticity varied the magnitude of this effect. We also found that hotter growing conditions reduced the variation in litter traits produced across genotypes, homogenizing the litter inputs to riparian ecosystems. All genotypes in the hottest garden produced comparatively small leaves that decomposed quickly and supported lower abundances of aquatic invertebrates, whereas the same genotypes in the coldest garden produced litter with distinct morphologies and decomposition rates. Our results suggest that plastic responses to climate stress may constrict the expression of genetic variation in predictable ways that impact communities and ecosystems. Understanding these interactions between genetic and environmental variation is critical to our ability to plan for the role of foundation species when managing and restoring riparian ecosystems in a warming world.

An experimental test of how parasites of predators can influence trophic cascades and ecosystem functioning.

Parasites can shape the structure and function of ecosystems by influencing both the density and traits of their hosts. Such changes in ecosystems are particularly likely when the host is a predator that mediates the dynamics of trophic cascades. Here, we experimentally tested how parasite load of a small predatory fish, the threespine stickleback, can affect the occurrence and strength of trophic cascades and ecosystem functioning. In a factorial mesocosm experiment, we manipulated the density of stickleback (low vs. high), and the level of parasite load (natural vs. reduced). In addition, we used two stickleback populations from different lineages: an eastern European lineage with a more pelagic phenotype (Lake Constance) and a western European lineage with a more benthic phenotype (Lake Geneva). We found that stickleback caused trophic cascades in the pelagic but not the benthic food chain. Evidence for pelagic trophic cascades was stronger in treatments where parasite load of stickleback was reduced with an antihelmintic medication, and where fish originated from Lake Constance (i.e., the more pelagic lineage). A structural equation model revealed that differences in stickleback lineage and parasite load were most likely to impact trophic cascades via changes in the composition, rather than overall biomass, of zooplankton communities. Overall, our results provide experimental evidence that parasites of predators can influence the cascading effects of fish on lower trophic levels with consequences on ecosystem functioning.

Rapid Divergence of Predator Functional Traits Affects Prey Composition in Aquatic Communities.

Identifying traits that underlie variation in individual performance of consumers (i.e., trait utility) can help reveal the ecological causes of population divergence and the subsequent consequences for species interactions and community structure. Here, we document a case of rapid divergence (over the past 100 generations, or ∼150 years) in foraging traits and feeding efficiency between a lake and stream population pair of threespine stickleback. Building on predictions from functional trait models of fish feeding, we analyzed foraging experiments with a Bayesian path analysis and elucidated the traits explaining variation in foraging performance and the species composition of ingested prey. Despite extensive previous research on the divergence of foraging traits among populations and ecotypes of stickleback, our results provide novel experimental evidence of trait utility for jaw protrusion, gill raker length, and gill raker spacing when foraging on a natural zooplankton assemblage. Furthermore, we discuss how these traits might contribute to the differential effects of lake and stream stickleback on their prey communities, observed in both laboratory and mesocosm conditions. More generally, our results illustrate how the rapid divergence of functional foraging traits of consumers can impact the biomass, species composition, and trophic structure of prey communities.

The role of plasticity in the evolution of cryptic pigmentation in a freshwater isopod.

Cryptic pigmentation of prey is often thought to evolve in response to predator-mediated selection, but pigmentation traits can also be plastic, and change with respect to both abiotic and biotic environmental conditions. In such cases, identifying the presence of, and drivers of trait plasticity is useful for understanding the evolution of crypsis. Previous work suggests that cryptic pigmentation of freshwater isopods (Asellus aquaticus) has evolved in response to predation pressure by fish in habitats with varying macrophyte cover and coloration. However, macrophytes can potentially influence the distribution of pigmentation by altering not only habitat-specific predation susceptibility, but also dietary resources and abiotic conditions. The goals of this study were to experimentally test how two putative agents of selection, namely macrophytes and fish, affect the pigmentation of A. aquaticus, and to assess whether pigmentation is plastic, using a diet manipulation in a common garden. We performed two experiments: (a) in an outdoor mesocosm experiment, we investigated how different densities of predatory fish (0/30/60 three-spined stickleback [Gasterosteus aculeatus] per mesocosm) and macrophytes (presence/absence) affected the abundance, pigmentation and body size structure of isopod populations. (b) In a subsequent laboratory experiment, we reared isopods in a common garden experiment on two different food sources (high/low protein content) to test whether variation in pigmentation of isopods can be explained by diet-based developmental plasticity. We found that fish presence strongly reduced isopod densities, particularly in the absence of macrophytes, but had no effect on pigmentation or size structure of the populations. However, we found that isopods showed consistently higher pigmentation in the presence of macrophytes, regardless of fish presence or absence. Our laboratory experiment, in which we manipulated the protein content of the isopods' diet, revealed strong plasticity of pigmentation and weak plasticity of growth rate. The combined results of both experiments suggest that pigmentation of A. aquaticus is a developmentally plastic trait and that multiple environmental factors (e.g. macrophytes, diet and predation) might jointly influence the evolution of cryptic pigmentation of A. aquaticus in nature on relatively short time-scales.

Genotypic variation in phenological plasticity: Reciprocal common gardens reveal adaptive responses to warmer springs but not to fall frost.

Species faced with rapidly shifting environments must be able to move, adapt, or acclimate in order to survive. One mechanism to meet this challenge is phenotypic plasticity: altering phenotype in response to environmental change. Here, we investigated the magnitude, direction, and consequences of changes in two key phenology traits (fall bud set and spring bud flush) in a widespread riparian tree species, Populus fremontii. Using replicated genotypes from 16 populations from throughout the species' thermal range, and reciprocal common gardens at hot, warm, and cool sites, we identified four major findings: (a) There are significant genetic (G), environmental (E), and GxE components of variation for both traits across three common gardens; (b) The magnitude of phenotypic plasticity is correlated with provenance climate, where trees from hotter, southern populations exhibited up to four times greater plasticity compared to the northern, frost-adapted populations; (c) Phenological mismatches are correlated with higher mortality as the transfer distances between provenance and garden increase; and (d) The relationship between plasticity and survival depends not only on the magnitude and direction of environmental transfer, but also on the type of environmental stress (i.e., heat or freezing), and how particular traits have evolved in response to that stress. Trees transferred to warmer climates generally showed small to moderate shifts in an adaptive direction, a hopeful result for climate change. Trees experiencing cooler climates exhibited large, non-adaptive changes, suggesting smaller transfer distances for assisted migration. This study is especially important as it deconstructs trait responses to environmental cues that are rapidly changing (e.g., temperature and spring onset) and those that are fixed (photoperiod), and that vary across the species' range. Understanding the magnitude and adaptive nature of phenotypic plasticity of multiple traits responding to multiple environmental cues is key to guiding restoration management decisions as climate continues to change.

Assessing Feeding Preferences of a Consumer Guild: Partitioning Variation Among versus Within Species.

Interspecific variation in resource use is critical to understanding species diversity, coexistence, and ecosystem functioning. A growing body of research describes analogous intraspecific variation and its potential importance for population dynamics and community outcomes. However, the magnitude of intraspecific variation relative to interspecific variation in key dimensions of consumer-resource interactions remains unknown, hampering our understanding of the importance of this variation for population and community processes. In this study, we examine feeding preference through repeated laboratory choice feeding assays of 444 wild-caught individuals of eight invertebrate grazer species on rocky reefs in northern California. Between-species variation accounted for 25%-33% of the total variation in preference for the preferred resource, while between-individual variation accounted for 4%-5% of total variation. For two of the eight species, between-individual variation was significantly different from zero and on average contributed 14% and 17% of the total diet variation, even after accounting for differences due to size and sex. Therefore, even with clearly distinguishable between-species differences in mean preference, diet variation between and within individuals can contribute to the dietary niche width of species and guilds, which may be overlooked by focusing solely on species' mean resource use patterns.

Evolution as an ecosystem process: insights from genomics.

Evolution is a fundamental ecosystem process. The study of genomic variation of organisms can not only improve our understanding of evolutionary processes, but also of contemporary and future ecosystem dynamics. We argue that integrative research between the fields of genomics and ecosystem ecology could generate new insights. Specifically, studies of biodiversity and ecosystem functioning, evolutionary rescue, and eco-evolutionary dynamics could all benefit from information about variation in genome structure and the genetic architecture of traits, whereas genomic studies could benefit from information about the ecological context of evolutionary dynamics. We propose new ways to help link research on functional genomic diversity with (reciprocal) interactions between phenotypic evolution and ecosystem change. Despite numerous challenges, we anticipate that the wealth of genomic data being collected on natural populations will improve our understanding of ecosystems.

Transgenerational selection driven by divergent ecological impacts of hybridizing lineages.

Dynamic interactions between ecological conditions and the phenotypic composition of populations likely play an important role in evolution, but the direction and strength of these feedbacks remain difficult to characterize. We investigated these dynamics across two generations of threespine sticklebacks from two evolutionary lineages undergoing secondary contact and hybridization. Independently manipulating the density and lineage of adults in experimental mesocosms led to contrasting ecosystem conditions with strong effects on total survival in a subsequent generation of juveniles. Ecosystem modifications by adults also varied the strength of selection on competing hybrid and non-hybrid juveniles. This variation in selection indicated (1) a negative eco-evolutionary feedback driven by lineage-specific resource depletion and dependence and (2) a large performance advantage of hybrid juveniles in depleted environments. This work illustrates the importance of interactions between phenotype, population density and the environment in shaping selection and evolutionary trajectories, especially in the context of range expansion with secondary contact and hybridization.

Stoichiometric traits of stickleback: Effects of genetic background, rearing environment, and ontogeny.

Phenotypes can both evolve in response to, and affect, ecosystem change, but few examples of diverging ecosystem-effect traits have been investigated. Bony armor traits of fish are good candidates for this because they evolve rapidly in some freshwater fish populations, and bone is phosphorus rich and likely to affect nutrient recycling in aquatic ecosystems. Here, we explore how ontogeny, rearing environment, and bone allocation among body parts affect the stoichiometric phenotype (i.e., stoichiometric composition of bodies and excretion) of threespine stickleback. We use two populations from distinct freshwater lineages with contrasting lateral plating phenotypes (full vs. low plating) and their hybrids, which are mostly fully plated. We found that ontogeny, rearing environment, and body condition were the most important predictors of organismal stoichiometry. Although elemental composition was similar between both populations and their hybrids, we found significant divergence in phosphorus allocation among body parts and in phosphorus excretion rates. Overall, body armor differences did not explain variation in whole body phosphorus, phosphorus allocation, or phosphorus excretion. Evolutionary divergence between these lineages in both allocation and excretion is likely to have important direct consequences for ecosystems, but may be mediated by evolution of multiple morphological or physiological traits beyond plating phenotype.

Evolution in a Community Context: On Integrating Ecological Interactions and Macroevolution.

Despite a conceptual understanding that evolution and species interactions are inextricably linked, it remains challenging to study ecological and evolutionary dynamics together over long temporal scales. In this review, we argue that, despite inherent challenges associated with reconstructing historical processes, the interplay of ecology and evolution is central to our understanding of macroevolution and community coexistence, and cannot be safely ignored in community and comparative phylogenetic studies. We highlight new research avenues that foster greater consideration of both ecological and evolutionary dynamics as processes that occur along branches of phylogenetic trees. By promoting new ways forward using this perspective, we hope to inspire further integration that creatively co-utilizes phylogenies and ecological data to study eco-evolutionary dynamics over time and space.

Plant-animal diversity relationships in a rocky intertidal system depend on invertebrate body size and algal cover.

Considerable research has examined the influence of herbivores on the maintenance of plant diversity, but fewer studies have examined the reciprocal effect of plant diversity on the animals that use the plant community for food and shelter, particularly in marine systems. Several mechanisms could underlie such effects. Animal diversity and abundance could be increased by complementary use of different plants by different animals, or by an indirect effect of plant diversity on plant production that results in more total plant biomass in high plant-diversity communities. Alternatively, plant species identity could play a dominant role leading to sampling effects or no effect of diversity at all. We conducted a six-year field manipulation of the richness of rocky shore seaweeds in northern California and measured the effects of algal richness and identity on the invertebrate community, from meiofauna to macrofauna. We found that diverse algal communities hosted more species of both large and small invertebrates than the average algal monoculture but that the mechanisms underlying this pattern differed substantially for organisms of different size. More species of macrofauna occurred in the polycultures than in any of the monocultures, likely due to the greater total cover of algae produced in polycultures. Rare and common macrofaunal taxa responded to host plant species richness in opposite ways, with more occurrences of rare taxa and lower abundance of very common taxa in the polycultures. In contrast, meiofaunal richness in polycultures was no different than that of monocultures of finely branched species, leading to strong effects of algal identity. Our findings are similar to those from terrestrial systems in that the effects of plant diversity we observed were most related to the greater amount of habitat in polycultures as a result of overyielding in algal biomass. However, our findings differ from those in terrestrial systems in that the primary mechanisms for both richness and identity effects appear related to the value of plants as shelter from harsh abiotic conditions or predation rather than food, and in that animal body size altered the mechanisms underlying diversity effects.

Phenotypic and phylogenetic evidence for the role of food and habitat in the assembly of communities of marine amphipods.

The study of community assembly processes currently involves (a) longstanding questions about the relative importance of environmental filtering vs. niche partitioning in a wide range of ecosystems, and (b) more recent questions about methodology. The rapidly growing field of community phylogenetics has generated debate about the choice between functional traits and phylogenetic relationships for understanding species similarities, and has raised additional questions about the contribution of experimental vs. observational approaches to understanding evolutionary constraints on community assembly. In this study, we use traits, a phylogeny, and field surveys to identify the forces structuring communities of herbivorous marine amphipods and isopods living in adjacent seagrass and macroalgae. In addition, we compare our field results to a recently published mesocosm experiment that tested the effects of both trait and phylogenetic diversity on coexistence using the same species and system. With respect to community assembly processes, we found that environmental filtering was the dominant process in macroalgae habitats, that niche partitioning was the dominant process in seagrass habitats, and that the strength of these assembly mechanisms varied with seasonal fluctuations in environmental conditions and resource availability. These patterns are indicated by both phylogenetic relationships and trait distances, but the type of resources being partitioned in seagrass habitats can only be deciphered using trait data. Species coexisting in seagrass in the field differed not in their feeding niche but in traits related to microhabitat use, providing novel evidence of the relative importance of competition for food vs. habitat in structuring communities of phytophagous invertebrates. With respect to methodology, the results for seagrass habitats conflict with those obtained in mesocosms, where feeding trait diversity did promote coexistence and phylogenetic diversity had no effect. This contrast arises because a greater range of traits (some of which have much stronger phylogenetic signal than feeding traits) contribute to community assembly in the field. This highlights a mismatch between the processes that drive community assembly in the field and the processes we isolated in experimental tests, and illustrates that using phylogeny as a single proxy in both contexts may impede the synthesis of observational and experimental results.

Phylogeny as a proxy for ecology in seagrass amphipods: which traits are most conserved?

Increasingly, studies of community assembly and ecosystem function combine trait data and phylogenetic relationships to gain novel insight into the ecological and evolutionary constraints on community dynamics. However, the key to interpreting these two types of information is an understanding of the extent to which traits are phylogenetically conserved. In this study, we develop the necessary framework for community phylogenetics approaches in a system of marine crustacean herbivores that play an important role in the ecosystem functioning of seagrass systems worldwide. For 16 species of amphipods and isopods, we (1) reconstructed phylogenetic relationships using COI, 16S, and 18S sequences and Bayesian analyses, (2) measured traits that are potentially important for assembling species between and within habitats, and (3) compared the degree to which each of these traits are evolutionarily conserved. Despite poor phylogenetic resolution for the order Amphipoda as a whole, we resolved almost all of the topology for the species in our system, and used a sampling of ultrametric trees from the posterior distribution to account for remaining uncertainty in topology and branch lengths. We found that traits varied widely in their degree of phylogenetic signal. Body mass, fecundity, and tube building showed very strong phylogenetic signal, and temperature tolerance and feeding traits showed much less. As such, the degree of signal was not predictable based on whether the trait is related to environmental filtering or to resource partitioning. Further, we found that even with strong phylogenetic signal in body size, (which may have large impacts on ecosystem function), the predictive relationship between phylogenetic diversity and ecosystem function is not straightforward. We show that patterns of phylogenetic diversity in communities of seagrass mesograzers could lead to a variety of interpretations and predictions, and that detailed study of trait similarities and differences will be necessary to interpret these patterns.

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.

Exotic herbivores directly facilitate the exotic grasses they graze: mechanisms for an unexpected positive feedback between invaders.

The ability of an exotic species to establish in a system may depend not only on the invasibility of the native community, but also on its interactions with other exotic species. Though examples of mutually beneficial interactions between exotic species are known, few studies have quantified these effects or identified specific mechanisms. We used the co-invasion of an endangered island ecosystem by exotic Canada geese (Branta canadensis) and nine exotic annual grasses to study the effects of an invading herbivore on the success of invading grasses. On our study islands in southwestern Canada, we found that geese fed selectively on the exotic grasses and avoided native forbs. Counter to current theory suggesting that the grasses should be limited by a selective enemy, however, the grasses increased in proportional abundance under grazing whereas forbs showed declining abundance. Testing potential mechanisms for the effects of grazing on grasses, we found that the grasses produced more stems per area when grazing reduced vegetation height and prevented litter accumulation. Forming dense mats of short stems appeared to be an efficient reproductive and competitive strategy that the Eurasian grasses have evolved in the presence of grazers, conferring a competitive advantage in a system where the native species pool has very few annual grasses and no grazers. Germination trials further demonstrated that selective herbivory by geese enables their dispersal of exotic grass seed between heavily invaded feeding areas and the small islands used for nesting. In summary, the exotic geese facilitated both the local increase and the spatial spread of exotic grasses, which in turn provided the majority of their diet. This unexpected case of positive feedback between exotic species suggests that invasion success may depend on the overall differences between the evolutionary histories of the invaders and the evolutionary history of the native community they enter.

Complementarity in marine biodiversity manipulations: reconciling divergent evidence from field and mesocosm experiments.

Mounting concern over the loss of marine biodiversity has increased the urgency of understanding its consequences. This urgency spurred the publication of many short-term studies, which often report weak effects of diversity (species richness) driven by the presence of key species (the sampling effect). Longer-term field experiments are slowly accumulating, and they more often report strong diversity effects driven by species complementarity, calling into question the generality of earlier findings. However, differences among study systems in which short- and long-term studies are conducted currently limit our ability to assess whether these differences are simply due to biological or environmental differences among systems. In this paper, we compared the effect of intertidal seaweed species richness on biomass accumulation in mesocosms and field experiments using the same pool of species. We found that seaweed species richness increased biomass accumulation in field experiments in both short (2-month) and long (3-year) experiments, although effects were stronger in the long-term experiment. In contrast, richness had no effect in mesocosm experiments, where biomass accumulation was completely a function of species identity. We argue that the short-term experiments, like many published experiments on the topic, detect only a subset of possible mechanisms that operate in the field over the longer term because they lack sufficient environmental heterogeneity to allow expression of niche differences, and they are of insufficient length to capture population-level responses, such as recruitment. Many published experiments, therefore, likely underestimate the strength of diversity on ecosystem processes in natural ecosystems.

Exotic grasses and feces deposition by an exotic herbivore combine to reduce the relative abundance of native forbs.

Increased resource availability can facilitate establishment of exotic plant species, especially when coincident with propagule supply. Following establishment, increased resource availability may also facilitate the spread of exotic plant species if it enhances their competitive abilities relative to native species. Exotic Canada geese (Branta canadensis) introduce both exotic grass seed and nutrients to an endangered plant community on the Gulf Islands of southwestern British Columbia, Canada. I used greenhouse experiments to assess the competitive advantage of the exotic grasses relative to native and exotic forbs in this community and to test the impacts of nutrient addition from goose feces on competitive outcomes. I grew experimental communities varying in their proportion of forbs versus exotic grasses, and added goose feces as a nutrient source. I found that both native and exotic forbs produced significantly more biomass in competition with conspecifics than in competition with the grasses, and that the proportional abundance of two out of three native forbs was lowest in the combined presence of exotic grasses and nutrient addition. In a second experiment, I found that in monoculture all species of forbs and grasses showed equal growth responses to nutrients. The exotic species did not convert additional nutrients into additional biomass at a higher rate, but did germinate earlier and grow larger than the native species regardless of nutrient availability. This suggests that the exotic species may have achieved their competitive advantage partly by pre-empting resources in community mixtures. Small and late-germinating native forbs may be particularly vulnerable to competitive suppression from exotic grasses and forbs and may be at an even greater disadvantage if their competitors are benefiting from early access to additional nutrients. In combination, the input of exotic propagules and additional nutrients by nesting geese may compromise efforts to maintain native community composition in this system.