A hybrid beachgrass (Ammophila arenaria × A. breviligulata) is more productive and outcompetes its non-native parent species.

The ability of non-native species to successfully invade new ecosystems sometimes involves evolutionary processes such as hybridization. Hybridization can produce individuals with superior traits that give them a competitive advantage over their parent species, allowing for rapid spread. Here we assess growth, functional morphology, and species interactions between two non-native beachgrass species (Ammophila arenaria and A. breviligulata) and their recently discovered hybrid (A. arenaria × A. breviligulata) on the U.S. Pacific Northwest coast. We asked whether the hybrid beachgrass differs from its parent species in morphology and growth, whether it competes with its parent species, and, if so, what are the potential mechanisms of competition. Plant taxa were grown in low- and high-density monocultures and in two-way interactions in a common garden environment. We show that the hybrid grew taller and more densely, with greater total biomass, than either parent species. The hybrid was also the better competitor, resulting in the model prediction of competitive exclusion against A. breviligulata and, depending on its relative abundance, A. arenaria. The hybrid displays a mixed 'guerilla-phalanx' growth form that allows it to spread laterally and achieve high shoot densities, giving it a competitive advantage. Given the current dominance of A. breviligulata compared to A. arenaria in most of the region where these taxa co-occur, we suggest that the hybrid will grow, compete, and spread quickly with potentially widespread consequences for the two non-native Ammophila congeners and the dunes they build.

Top ten priorities for global saltmarsh restoration, conservation and ecosystem service research.

Coastal saltmarshes provide globally important ecosystem services including 'blue carbon' sequestration, flood protection, pollutant remediation, habitat provision and cultural value. Large portions of marshes have been lost or fragmented as a result of land reclamation, embankment construction, and pollution. Sea level rise threatens marsh survival by blocking landward migration where coastlines have been developed. Research-informed saltmarsh conservation and restoration efforts are helping to prevent further loss, yet significant knowledge gaps remain. Using a mixed methods approach, this paper identifies ten research priorities through an online questionnaire and a residential workshop attended by an international, multi-disciplinary network of 35 saltmarsh experts spanning natural, physical and social sciences across research, policy, and practitioner sectors. Priorities have been grouped under four thematic areas of research: Saltmarsh Area Extent, Change and Restoration Potential (including past, present, global variation), Spatio-social contexts of Ecosystem Service delivery (e.g. influences of environmental context, climate change, and stakeholder groups on service provisioning), Patterns and Processes in saltmarsh functioning (global drivers of saltmarsh ecosystem structure/function) and Management and Policy Needs (how management varies contextually; challenges/opportunities for management). Although not intended to be exhaustive, the challenges, opportunities, and strategies for addressing each research priority examined here, providing a blueprint of the work that needs to be done to protect saltmarshes for future generations.

Warming alters the interaction of two invasive beachgrasses with implications for range shifts and coastal dune functions.

Forecasting the effects of climate change on the distribution of invasive species can be difficult, because invaders often thrive under novel physical conditions and biotic interactions that differ from those in their native range. In this study, we experimentally examined how rising temperatures and sand burial could alter the abundance and biotic interactions of two invasive beachgrasses, Ammophila arenaria and A. breviligulata, along the U.S. Pacific Northwest coast. We asked whether the current geographic ranges of the two congeners, and thus their effects on dune morphology and coastal ecosystem services, might shift as a consequence of climate driven changes in warming and sand supply. Our results show that A. breviligulata had lower biomass and tiller production when exposed to warming and high rates of sand burial, while A. arenaria showed neutral or positive responses to those treatments. Nevertheless, under all experimental combinations, A. breviligulata had strong negative effects on A. arenaria, while A. arenaria had weaker effects on A. breviligulata. Our models predict that although A. breviligulata mostly excludes A. arenaria, elevated temperatures and high rates of sand burial also increase the likelihood of species coexistence. We suggest that under climate change, the differences in physiological tolerance and the mediation of species interactions could expand the northern distributional limit of A. arenaria but restrict the southern limit of A. breviligulata. Moreover, because beachgrass abundance has direct effects on biophysical functions of dunes, reductions in vigor from warming could alter coastal protection, biodiversity, and carbon sequestration.

The non-market benefits of early and partial gains in managing threatened salmon.

Threatened species are increasingly dependent on conservation investments for persistence and recovery. Information that resource managers could use to evaluate investments-such as the public benefits arising from alternative conservation designs-is typically scarce because conservation benefits arise outside of conventional markets. Moreover, existing studies that measure the public benefits of conserving threatened species often do not measure the benefits from partial gains in species abundance that fall short of official recovery, or the benefits from achieving gains in species abundance that happen earlier in time. We report on a stated preference choice experiment designed to quantify the non-market benefits for conservation investments aimed at threatened Pacific Coho salmon (Oncorhynchus kisutch) along the Oregon Coast (OC). Our results show that a program aimed at increasing numbers of returning salmon can generate sizable benefits of up to $518 million/y for an extra 100,000 returning fish, even if the species is not officially declared recovered. Moreover, while conservation investment strategies expected to achieve relatively rapid results are likely to have higher up-front costs, our results show that the public attaches substantial additional value of up to $277 million/y for achieving conservation goals quickly. Our results and approach can be used to price natural capital investments that lead to gains in returning salmon, and as inputs to evaluations of the benefits and costs from alternative conservation strategies.

Supporting Spartina: Interdisciplinary perspective shows Spartina as a distinct solid genus.

In 2014, a DNA-based phylogenetic study confirming the paraphyly of the grass subtribe Sporobolinae proposed the creation of a large monophyletic genus Sporobolus, including (among others) species previously included in the genera Spartina, Calamovilfa, and Sporobolus. Spartina species have contributed substantially (and continue contributing) to our knowledge in multiple disciplines, including ecology, evolutionary biology, molecular biology, biogeography, experimental ecology, biological invasions, environmental management, restoration ecology, history, economics, and sociology. There is no rationale so compelling to subsume the name Spartina as a subgenus that could rival the striking, global iconic history and use of the name Spartina for over 200 yr. We do not agree with the subjective arguments underlying the proposal to change Spartina to Sporobolus. We understand the importance of both the objective phylogenetic insights and of the subjective formalized nomenclature and hope that by opening this debate we will encourage positive feedback that will strengthen taxonomic decisions with an interdisciplinary perspective. We consider that the strongly distinct, monophyletic clade Spartina should simply and efficiently be treated as the genus Spartina.

Regional processes are stronger determinants of rocky intertidal community dynamics than local biotic interactions.

Understanding the relative roles of species interactions and environmental factors in structuring communities has historically focused on local scales where manipulative experiments are possible. However, recent interest in predicting the effects of climate change and species invasions has spurred increasing attention to processes occurring at larger spatial and temporal scales. The "meta-ecosystem" approach is an ideal framework for integrating processes operating at multiple scales as it explicitly considers the influence of local biotic interactions and regional flows of energy, materials, and organisms on community structure. Using a comparative-experimental design, we asked (1) what is the relative importance of local biotic interactions and oceanic processes in determining rocky intertidal community structure in the low zone within the Northern California Current System, and (2) what factors are most important in regulating this structure and why? We focused on functional group interactions between macrophytes and sessile invertebrates and their consumers (grazers, predators), how these varied across spatial scales, and with ocean-driven conditions (upwelling, temperature) and ecological subsidies (nutrients, phytoplankton, sessile invertebrate recruits). Experiments were conducted at 13 sites divided across four capes in Oregon and northern California. Results showed that biotic interactions were variable in space and time but overall, sessile invertebrates had no effect on macrophytes while macrophytes had weakly negative effects on sessile invertebrates. Consumers, particularly predators, also had weakly negative effects on both functional groups. Overall, we found that 40-49% of the variance in community structure at the local scale was explained by external factors (e.g., spatial scale, time, upwelling, temperature, ecological subsidies) vs. 19-39% explained by functional group interactions. When individual functional group interaction strengths were used, only 2-3% of the variation was explained by any one functional group while 28-54% of the variation was explained by external factors. We conclude that community structure in the low intertidal zone is driven primarily by external factors at the regional scale with local biotic interactions playing a secondary role.

Generality in multispecies responses to ocean acidification revealed through multiple hypothesis testing.

Decades of research have demonstrated that many calcifying species are negatively affected by ocean acidification, a major anthropogenic threat in marine ecosystems. However, even closely related species may exhibit different responses to ocean acidification and less is known about the drivers that shape such variation in different species. Here, we examine the drivers of physiological performance under ocean acidification in a group of five species of turf-forming coralline algae. Specifically, quantitating the relative weight of evidence for each of ten hypotheses, we show that variation in coralline calcification and photosynthesis was best explained by allometric traits. Across ocean acidification conditions, larger individuals (measured as noncalcified mass) had higher net calcification and photosynthesis rates. Importantly, our approach was able to not only identify the aspect of size that drove the performance of coralline algae, but also determined that responses to ocean acidification were not dependent on species identity, evolutionary relatedness, habitat, shape, or structural composition. In fact, we found that failure to test multiple, alternative hypotheses would underestimate the generality of physiological performances, leading to the conclusion that each species had different baseline performance under ocean acidification. Testing among alternative hypotheses is an essential step toward determining the generalizability of experiments across taxa and identifying common drivers of species responses to global change.

Literature-based latitudinal distribution and possible range shifts of two US east coast dune grass species (Uniola paniculata and Ammophila breviligulata).

Previous work on the US Atlantic coast has generally shown that coastal foredunes are dominated by two dune grass species, Ammophila breviligulata (American beachgrass) and Uniola paniculata (sea oats). From Virginia northward, A. breviligulata dominates, while U. paniculata is the dominant grass south of Virginia. Previous work suggests that these grasses influence the shape of coastal foredunes in species-specific ways, and that they respond differently to environmental stressors; thus, it is important to know which species dominates a given dune system. The range boundaries of these two species remains unclear given the lack of comprehensive surveys. In an attempt to determine these boundaries, we conducted a literature survey of 98 studies that either stated the range limits and/or included field-based studies/observations of the two grass species. We then produced an interactive map that summarizes the locations of the surveyed papers and books. The literature review suggests that the current southern range limit for A. breviligulata is Cape Fear, NC, and the northern range limit for U. paniculata is Assateague Island, on the Maryland and Virginia border. Our data suggest a northward expansion of U. paniculata, possibly associated with warming trends observed near the northern range limit in Painter, VA. In contrast, the data regarding a range shift for A. breviligulata remain inconclusive. We also compare our literature-based map with geolocated records from the Global Biodiversity Information Facility and iNaturalist research grade crowd-sourced observations. We intend for our literature-based map to aid coastal researchers who are interested in the dynamics of these two species and the potential for their ranges to shift as a result of climate change.

Fundamental contradictions among observational and experimental estimates of non-trophic species interactions.

The difficulty of experimentally quantifying non-trophic species interactions has long troubled ecologists. Increasingly, a new application of the classic "checkerboard distribution" approach is used to infer interactions by examining the pairwise frequency at which species are found to spatially co-occur. However, the link between spatial associations, as estimated from observational co-occurrence, and species interactions has not been tested. Here we used nine common statistical methods to estimate associations from surveys of rocky intertidal communities in the Northeast Pacific Ocean. We compared those inferred associations with a new data set of experimentally determined net and direct species interactions. Although association methods generated networks with aggregate structure similar to previously published interaction networks, each method detected a different set of species associations from the same data set. Moreover, although association methods generally performed better than a random model, associations rarely matched empirical net or direct species interactions, with high rates of false positives and true positives, and many false negatives. Our findings cast doubt on studies that equate species co-occurrences to species interactions and highlight a persistent, unanswered question: how do we interpret spatial patterns in communities? We suggest future research directions to unify the observational and experimental study of species interactions, and discuss the need for community standards and best practices in association analysis.

Are large macroalgal blooms necessarily bad? Nutrient impacts on seagrass in upwelling-influenced estuaries.

Knowledge of nutrient pathways and their resulting ecological interactions can alleviate numerous environmental problems associated with nutrient increases in both natural and managed systems. Although not unique, coastal systems are particularly prone to complex ecological interactions resulting from nutrient inputs from both the land and sea. Nutrient inputs to coastal systems often spur ulvoid macroalgal blooms, with negative consequences for seagrasses, primarily through shading, as well as through changes in local biogeochemistry. We conducted complementary field and mesocosm experiments in an upwelling-influenced estuary, where marine-derived nutrients dominate, to understand the direct and indirect effects of nutrients on the macroalgal-eelgrass (Zostera marina L.) interaction. In the field experiment, we found weak evidence that nutrients and/or macroalgal treatments had a negative effect on eelgrass. However, in the mesocosm experiment, we found that a combination of nutrient and macroalgal treatments led to strongly negative eelgrass responses, primarily via indirect effects associated with macroalgal additions. Together, increased total light attenuation and decreased sediment oxygen levels were associated with larger effects on eelgrass than shading alone, which was evaluated using mimic algae treatments that did not alter sediment redox potential. Nutrient addition in the mesocosms directly affected seagrass density; biomass, and morphology, but not as strongly as macroalgae. We hypothesize that the contrary results from these parallel experiments are a consequence of differences in the hydrodynamics between field and mesocosm settings. We suggest that the high rates of water movement and tidal submersion of our intertidal field experiments alleviated the light reduction and negative biogeochemical changes in the sediment associated with macroalgal canopies, as well as the nutrient effects observed in the mesocosm experiments. Furthermore, adaptation of ulvoids and eelgrass to high, but variable, background nutrient concentrations in upwelling-influenced estuaries may partly explain the venue-specific results reported here. In order to manage critical seagrass habitats, nutrient criteria and macroalgal indicators must consider variability in marine-based nutrient delivery and local physical conditions among estuaries.

Coastal foredune evolution: the relative influence of vegetation and sand supply in the US Pacific Northwest.

Biophysical feedbacks between vegetation and sediment are important for forming and modifying landscape features and their ecosystem services. These feedbacks are especially important where landscape features differ in their provision of ecosystem services. For example, the shape of coastal foredunes, a product of both physical and biological forces, determines their ability to protect communities from rising seas and changing patterns of storminess. Here we assessed how sand supply and changes in vegetation over interannual (3 year) and decadal (21 year) scales influenced foredune shape along 100 km of coastline in the US Pacific Northwest. Across 21 years, vegetation switched from one congeneric non-native beachgrass to another (Ammophila arenaria to A. breviligulata) while sand supply rates were positive. At interannual timescales, sand supply rates explained the majority of change in foredune height (64-69%) and width (56-80%). However, at decadal scales, change in vegetation explained the majority of the change in foredune width (62-68%), whereas sand supply rates explained most of the change in foredune height (88-90%). In areas with lower shoreline change rates (±2 m yr(-1)), the change in vegetation explained the majority of decadal changes in foredune width (56-57%) and height (59-76%). Foredune shape directly impacts coastal protection, thus our findings are pertinent to coastal management given pressures of development and climate change.

Invasive congeners differ in successional impacts across space and time.

Invasive species can alter the succession of ecological communities because they are often adapted to the disturbed conditions that initiate succession. The extent to which this occurs may depend on how widely they are distributed across environmental gradients and how long they persist over the course of succession. We focus on plant communities of the USA Pacific Northwest coastal dunes, where disturbance is characterized by changes in sediment supply, and the plant community is dominated by two introduced grasses--the long-established Ammophila arenaria and the currently invading A. breviligulata. Previous studies showed that A. breviligulata has replaced A. arenaria and reduced community diversity. We hypothesize that this is largely due to A. breviligulata occupying a wider distribution across spatial environmental gradients and persisting in later-successional habitat than A. arenaria. We used multi-decadal chronosequences and a resurvey study spanning 2 decades to characterize distributions of both species across space and time, and investigated how these distributions were associated with changes in the plant community. The invading A. breviligulata persisted longer and occupied a wider spatial distribution across the dune, and this corresponded with a reduction in plant species richness and native cover. Furthermore, backdunes previously dominated by A. arenaria switched to being dominated by A. breviligulata, forest, or developed land over a 23-yr period. Ammophila breviligulata likely invades by displacing A. arenaria, and reduces plant diversity by maintaining its dominance into later successional backdunes. Our results suggest distinct roles in succession, with A. arenaria playing a more classically facilitative role and A. breviligulata a more inhibitory role. Differential abilities of closely-related invasive species to persist through time and occupy heterogeneous environments allows for distinct impacts on communities during succession.

Invasive grasses, climate change, and exposure to storm-wave overtopping in coastal dune ecosystems.

The world's coastal habitats are critical to human well-being, but are also highly sensitive to human habitat alterations and climate change. In particular, global climate is increasing sea levels and potentially altering storm intensities, which may result in increased risk of flooding in coastal areas. In the Pacific Northwest (USA), coastal dunes that protect the coast from flooding are largely the product of a grass introduced from Europe over a century ago (Ammophila arenaria). An introduced congener (A. breviligulata) is displacing A. arenaria and reducing dune height. Here we quantify the relative exposure to storm-wave induced dune overtopping posed by the A. breviligulata invasion in the face of projected multi-decadal changes in sea level and storm intensity. In our models, altered storm intensity was the largest driver of overtopping extent, however the invasion by A. breviligulata tripled the number of areas vulnerable to overtopping and posed a fourfold larger exposure than sea-level rise over multi-decadal time scales. Our work demonstrates the importance of a transdisciplinary approach that draws on insights from ecology, geomorphology, and civil engineering to assess the vulnerability of ecosystem services in light of global change.

Biophysical feedback mediates effects of invasive grasses on coastal dune shape.

Vegetation at the aquatic-terrestrial interface can alter landscape features through its growth and interactions with sediment and fluids. Even similar species may impart different effects due to variation in their interactions and feedbacks with the environment. Consequently, replacement of one engineering species by another can cause significant change in the physical environment. Here we investigate the species-specific ecological mechanisms influencing the geomorphology of U.S. Pacific Northwest coastal dunes. Over the last century, this system changed from open, shifting sand dunes with sparse vegetation (including native beach grass, Elymus mollis), to densely vegetated continuous foredune ridges resulting from the introduction and subsequent invasions of two nonnative grass species (Ammophila arenaria and Ammophila breviligulata), each of which is associated with different dune shapes and sediment supply rates along the coast. Here we propose a biophysical feedback responsible for differences in dune shape, and we investigate two, non-mutually exclusive ecological mechanisms for these differences: (1) species differ in their ability to capture sand and (2) species differ in their growth habit in response to sand deposition. To investigate sand capture, we used a moveable bed wind tunnel experiment and found that increasing tiller density increased sand capture efficiency and that, under different experimental densities, the native grass had higher sand capture efficiency compared to the Ammophila congeners. However, the greater densities of nonnative grasses under field conditions suggest that they have greater potential to capture more sand overall. We used a mesocosm experiment to look at plant growth responses to sand deposition and found that, in response to increasing sand supply rates, A. arenaria produced higher-density vertical tillers (characteristic of higher sand capture efficiency), while A. breviligulata and E. mollis responded with lower-density lateral tiller growth (characteristic of lower sand capture efficiency). Combined, these experiments provide evidence for a species-specific effect on coastal dune shape. Understanding how dominant ecosystem engineers, especially nonnative ones, differ in their interactions with abiotic factors is necessary to better parameterize coastal vulnerability models and inform management practices related to both coastal protection ecosystem services and ecosystem restoration.

Recruitment facilitation can promote coexistence and buffer population growth in metacommunities.

Although positive species interactions are ubiquitous in nature, theory has generally focused on the role of negative interactions to explain patterns of species diversity. Here, we incorporate recruitment facilitation, a positive interaction prevalent in marine and terrestrial systems, into a metacommunity framework to assess how the interplay between colonisation, competition and facilitation mediates coexistence. We show that when subordinate species facilitate the recruitment of dominant species, multi-species metacommunities can persist stably even if the colonisation rate of the dominant species is greater than that of the subordinate species. In addition, recruitment facilitation can buffer population growth from changes in colonisation rates, and thus explain the paradoxical mismatch between patterns of abundance and recruitment in marine systems. Overall, our results demonstrate that recruitment facilitation can have profound effects on the assembly, dissolution and regulation of metacommunities by mediating the relative influence of local and regional processes on population abundance and species diversity.

Ecosystem services as a common language for coastal ecosystem-based management.

Ecosystem-based management is logistically and politically challenging because ecosystems are inherently complex and management decisions affect a multitude of groups. Coastal ecosystems, which lie at the interface between marine and terrestrial ecosystems and provide an array of ecosystem services to different groups, aptly illustrate these challenges. Successful ecosystem-based management of coastal ecosystems requires incorporating scientific information and the knowledge and views of interested parties into the decision-making process. Estimating the provision of ecosystem services under alternative management schemes offers a systematic way to incorporate biogeophysical and socioeconomic information and the views of individuals and groups in the policy and management process. Employing ecosystem services as a common language to improve the process of ecosystem-based management presents both benefits and difficulties. Benefits include a transparent method for assessing trade-offs associated with management alternatives, a common set of facts and common currency on which to base negotiations, and improved communication among groups with competing interests or differing worldviews. Yet challenges to this approach remain, including predicting how human interventions will affect ecosystems, how such changes will affect the provision of ecosystem services, and how changes in service provision will affect the welfare of different groups in society. In a case study from Puget Sound, Washington, we illustrate the potential of applying ecosystem services as a common language for ecosystem-based management.

Current status of marine protected areas in latin america and the Caribbean.

Marine protected areas (MPAs), including no-take marine reserves (MRs), play an important role in the conservation of marine biodiversity. We document the status of MPAs and MRs in Latin America and the Caribbean, where little has been reported on the scope of such protection. Our survey of protected area databases, published and unpublished literature, and Internet searches yielded information from 30 countries and 12 overseas territories. At present more than 700 MPAs have been established, covering more than 300,000 km(2) or 1.5% of the coastal and shelf waters. We report on the status of 3 categories of protection: MPAs (limited take throughout the area), MRs (no-take throughout the area), and mixed-use (a limited-take MPA that contains an MR). The majority of protected areas in Latin America and the Caribbean are MPAs, which allow some or extensive extractive activities throughout the designated area. These 571 sites cover 51,505 km(2) or 0.3% of coastal and shelf waters. There are 98 MRs covering 16,862 km(2) or 0.1% of the coastal and shelf waters. Mixed-use MPAs are the fewest in number (87), but cover the largest area (236,853 km(2), 1.2%). Across Latin America and the Caribbean, many biogeographic provinces are underrepresented in these protected areas. Large coastal regions remain unprotected, in particular, the southern Pacific and southern Atlantic coasts of South America. Our analysis reveals multiple opportunities to strengthen marine conservation in Latin America and the Caribbean by improving implementation, management, and enforcement of existing MPAs; adding new MPAs and MRs strategically to enhance connectivity and sustainability of existing protection; and establishing new networks of MPAs and MRs or combinations thereof to enhance protection where little currently exists.

Coastal ecosystem-based management with nonlinear ecological functions and values.

A common assumption is that ecosystem services respond linearly to changes in habitat size. This assumption leads frequently to an "all or none" choice of either preserving coastal habitats or converting them to human use. However, our survey of wave attenuation data from field studies of mangroves, salt marshes, seagrass beds, nearshore coral reefs, and sand dunes reveals that these relationships are rarely linear. By incorporating nonlinear wave attenuation in estimating coastal protection values of mangroves in Thailand, we show that the optimal land use option may instead be the integration of development and conservation consistent with ecosystem-based management goals. This result suggests that reconciling competing demands on coastal habitats should not always result in stark preservation-versus-conversion choices.

Nitrogen effects on an interaction chain in a salt marsh community.

Nutrients can structure communities by influencing both plant interactions and plant herbivore interactions, though rarely do studies integrate these processes. In this study we examined how nitrogen fertilization influenced (1) the positive interaction between the marsh elder, Iva frutescens, and the black rush, Juncusgerardi, and (2) the quality of Iva as a host plant for the aphid, Uroleuconambrosiae. Previous studies have shown that by mitigating soil salt accumulation and hypoxia, Juncus is essential to the survival of Iva and its aphid herbivore at mid-marsh elevations. To address the effects of nitrogen on this interaction, we compared fertilized and unfertilized Iva plants subject to Juncus removal and control treatments in the field. Additionally, we measured the monthly population growth rates of aphids transplanted onto these Iva plants. Iva leaf biomass and flower number results indicated that fertilizing Iva eliminated its dependence upon Juncus, such that fertilized plants grown without Juncus were not different from unmanipulated plants. Aphid monthly population growth rates through mid-summer revealed that fertilization also eliminated the indirect dependency of aphids on Juncus, so that aphid growth rates on fertilized Iva without Juncus neighbors were similar to rates on unmanipulated Iva. Results also indicated that fertilizing Iva grown with Juncus increased Iva size, potentially enabling these plants to support larger aphid populations. Our results suggest that only under conditions of nitrogen limitation are the positive effects of Juncus essential to the mid-marsh persistence of Iva and its aphid herbivore. Furthermore, we found that nitrogen effects on aphid populations may arise not only from a direct effect of nutrients on Iva size but also through the indirect effects of nitrogen on the interaction between Juncus and Iva. We argue that studies integrating processes occurring both within and between trophic levels, are important to fully understanding the community-wide effects of nutrients.