Global patterns of allochthony in stream-riparian meta-ecosystems.

Ecosystems that are coupled by reciprocal flows of energy and nutrient subsidies can be viewed as a single "meta-ecosystem." Despite these connections, the reciprocal flow of subsidies is greatly asymmetrical and seasonally pulsed. Here, we synthesize existing literature on stream-riparian meta-ecosystems to quantify global patterns of the amount of subsidy consumption by organisms, known as "allochthony." These resource flows are important since they can comprise a large portion of consumer diets, but can be disrupted by human modification of streams and riparian zones. Despite asymmetrical subsidy flows, we found stream and riparian consumer allochthony to be equivalent. Although both fish and stream invertebrates rely on seasonally pulsed allochthonous resources, we find allochthony varies seasonally only for fish, being nearly three times greater during the summer and fall than during the winter and spring. We also find that consumer allochthony varies with feeding traits for aquatic invertebrates, fish, and terrestrial arthropods, but not for terrestrial vertebrates. Finally, we find that allochthony varies by climate for aquatic invertebrates, being nearly twice as great in arid climates than in tropical climates, but not for fish. These findings are critical to understanding the consequences of global change, as ecosystem connections are being increasingly disrupted.

Connectivity mediates the spatial ecological impacts of a glyphosate-based herbicide in experimental metaecosystems.

Metacommunity ecology has shown that connectivity is important for the persistence of a species locally and across connected ecosystems, however we do not know if ecological effects in freshwater ecosystems exposed to biocides leaking from agriculture depend on metaecosystem connectivity. We experimentally replicated metaecosystems in the laboratory using gradostats as a model system. We tested the effects of connectivity, in terms of node distance from the pollutant-source, flow rate, and a glyphosate-based herbicide, on phytoplankton productivity, diversity and stability. Gradostats were composed of interconnected equally spaced nodes where resources and phytoplankton move directionally along a gradient of increasing distance from the source of the polluting herbicide. We hypothesised that ecological effects would be stronger in the node situated closer to the point of herbicide input, but that flow would suppress phytoplankton populations in distant nodes. Overall, RoundUp impacted phytoplankton productivity and stability by reducing algal biomass and abundances. This occurred especially in the node closest to the diluted herbicide point-source and under high flow, where species abundances were heavily suppressed by the effects of the rapidly flowing herbicide. At low flow on the other hand, distant nodes where buffered from the effects of the slow-moving herbicide. No differences in beta and gamma diversity among replicate metaecosystems was found; however, a significant loss of alpha diversity in all metaecosystems occurred through time until the end of the experiment. Together, these results point to the importance of considering aquatic connectivity in management plans for monitoring and mitigating unintended ecological consequences of agrochemical runoff.

Quality matters: Stoichiometry of resources modulates spatial feedbacks in aquatic-terrestrial meta-ecosystems.

Species dispersal and resource spatial flows greatly affect the dynamics of connected ecosystems. So far, research on meta-ecosystems has mainly focused on the quantitative effect of subsidy flows. Yet, resource exchanges at heterotrophic-autotrophic (e.g. aquatic-terrestrial) ecotones display a stoichiometric asymmetry that likely matters for functioning. Here, we joined ecological stoichiometry and the meta-ecosystem framework to understand how subsidy stoichiometry mediates the response of the meta-ecosystem to subsidy flows. Our model results demonstrate that resource flows between ecosystems can induce a positive spatial feedback loop, leading to higher production at the meta-ecosystem scale by relaxing local ecosystem limitations ('spatial complementarity'). Furthermore, we show that spatial flows can also have an unexpected negative impact on production when accentuating the stoichiometric mismatch between local resources and basal species needs. This study paves the way for studies on the interdependency of ecosystems at the landscape extent.

The significance of partial migration for food web and ecosystem dynamics.

Migration is ubiquitous and can strongly shape food webs and ecosystems. Less familiar, however, is that the majority of life cycle, seasonal and diel migrations in nature are partial migrations: only a fraction of the population migrates while the other individuals remain in their resident ecosystem. Here, we demonstrate different impacts of partial migration rendering it fundamental to our understanding of the significance of migration for food web and ecosystem dynamics. First, partial migration affects the spatiotemporal distribution of individuals and the food web and ecosystem-level processes they drive differently than expected under full migration. Second, whether an individual migrates or not is regularly correlated with morphological, physiological, and/or behavioural traits that shape its food-web and ecosystem-level impacts. Third, food web and ecosystem dynamics can drive the fraction of the population migrating, enabling the potential for feedbacks between the causes and consequences of migration within and across ecosystems. These impacts, individually and in combination, can yield unintuitive effects of migration and drive the dynamics, diversity and functions of ecosystems. By presenting the first full integration of partial migration and trophic (meta-)community and (meta-)ecosystem ecology, we provide a roadmap for studying how migration affects and is affected by ecosystem dynamics in a changing world.

Fresh perspectives on the River Continuum Concept require trophic ecology approaches focussed on food web structure and energy mobilisation routes.

Stream-dwelling communities are expected to show a gradual replacement of the dominant feeding types following the type of resources found along the river continuum. Yet, the underlying longitudinal gradients in food web structure and energy flow-paths remain poorly understood. Here, I synthesise novel research on the River Continuum Concept (RCC) and identify promising areas for future research linked to longitudinal changes in food-chain length and energy mobilisation routes. For example, feeding links and connectance should reach maximum values in mid-order rivers and then decrease to river mouths following uncovered longitudinal diversity patterns. Regarding energy mobilisation routes, a gradual replacement in the food web fuelling between allochthonous (leaf litter) and autochthonous (periphyton) resources should be expected. Beyond longitudinal changes in primary basal resource to consumer paths, other allochthonous (e.g. riparian arthropod inputs) and autochthonous (e.g. fish prey) inputs subsidising higher level consumers may show longitudinal changes, that is, terrestrial invertebrates decreasing but piscivory increasing downstream. However, the role of these inputs, that can alter predator niche variation and have indirect community-based effects, on both food web structure and energy flow-paths along the river continuum is not clear yet. Incorporating energy mobilisation and food web structure into RCC principles is necessary for a broad understanding of ecosystem functioning and trophic diversity in riverine systems, driving the emergence of novel insights. How function and structure of riverine food webs adapt to longitudinal changes in physical and biological environments represent a challenge for next generation of stream ecologists.

Effect of human disturbances and hydrologic elements on the distribution of plant diversity within the Shamu watershed, Mt. Yuntai Nature Reserve, China.

This paper explores how human disturbance and hydrologic elements affect the spatial distribution pattern of plant diversity in the watershed, taking Shamu watershed in the World Natural Heritage Site as a case study. Spatial analysis of multisource remote sensing and plant diversity plots data were conducted using linear mixed effects models and structural equation models. Results revealed that the distribution of plant diversity in the watershed is mainly affected by human disturbance. However, under similar human disturbance levels, hydrologic elements also affect the plant diversity within the watershed. The topographic undulation and surface runoff significantly promote plant diversity, while the river network density, the watershed shape factor, the river longitudinal gradient do not. The influence of topographic undulation is more obvious than that of runoff on plant diversity, but the effect of topographic undulation and runoff on plant diversity is getting weaker from upstream to downstream within the watershed. In addition, the impact of hydrologic elements on plant diversity is mainly regulated by environmental factors Pre and Tem. The findings clarify how human disturbance and hydrologic elements affect plant diversity distribution within the watershed, optimizing the conservation theory of plant diversity resources and scientifically guiding the region's sustainable development.

A methodological roadmap to quantify animal-vectored spatial ecosystem subsidies.

Energy, nutrients and organisms move over landscapes, connecting ecosystems across space and time. Meta-ecosystem theory investigates the emerging properties of local ecosystems coupled spatially by these movements of organisms and matter, by explicitly tracking exchanges of multiple substances across ecosystem borders. To date, meta-ecosystem research has focused mostly on abiotic flows-neglecting biotic nutrient flows. However, recent work has indicated animals act as spatial nutrient vectors when they transport nutrients across landscapes in the form of excreta, egesta and their own bodies. Partly due to its high level of abstraction, there are few empirical tests of meta-ecosystem theory. Furthermore, while animals may be viewed as important mediators of ecosystem functions, better integration of tools is needed to develop predictive insights of their relative roles and impacts on diverse ecosystems. We present a methodological roadmap that explains how to do such integration by discussing how to combine insights from movement, foraging and ecosystem ecology to develop a coherent understanding of animal-vectored nutrient transport on meta-ecosystems processes. We discuss how the slate of newly developed technologies and methods-tracking devices, mechanistic movement models, diet reconstruction techniques and remote sensing-that when integrated have the potential to advance the quantification of animal-vectored nutrient flows and increase the predictive power of meta-ecosystem theory. We demonstrate that by integrating novel and established tools of animal ecology, ecosystem ecology and remote sensing, we can begin to identify and quantify animal-mediated nutrient translocation by large animals. We also provide conceptual examples that show how our proposed integration of methodologies can help investigate ecosystem impacts of large animal movement. We conclude by describing practical advancements to understanding cross-ecosystem contributions of animals on the move. Understanding the mechanisms by which animals shape ecosystem dynamics is important for ongoing conservation, rewilding and restoration initiatives around the world, and for developing more accurate models of ecosystem nutrient budgets. Our roadmap will enable ecologists to better qualify and quantify animal-mediated nutrient translocation for animals on the move.

Understanding the connectivity of ecosystems in the Anthropocene.

In focus: Nash, L. N., Antiqueira, P. A. P., Romero, G. Q., de Omena, P. M., and Kratina, P. (2021). Warming of aquatic ecosystems disrupts aquatic-terrestrial linkages in the tropics. Journal of Animal Ecology. Meta-ecosystem ecology offers a holistic framework linking populations, communities and ecosystems in heterogeneous landscapes. This perspective is particularly relevant as anthropogenic drivers of environmental change proliferate, with the potential for impacts to propagate to spatially connected habitats. In aquatic ecosystems, reciprocal exchanges of energy, materials and organisms that form strong connections with adjacent terrestrial habitats can be disrupted by human impacts. Nash et al. (2021) demonstrate how a warming environment alters aquatic-terrestrial linkages by measuring rates of aquatic insect emergence and decomposition in a tropical context. While decomposition predictably increased with warming, insect emergence was greatly reduced. Using metabolic theory, I highlight how these results deviate from previous studies and help underscore the need for comparative research in different biomes. I conclude by exploring where progress can be made in quantifying, predicting and utilising the connectivity of ecosystems to fully realise the potential of a meta-ecosystem perspective and help address the dual crises of biodiversity loss and climate change.

Cross-ecosystem bottlenecks alter reciprocal subsidies within meta-ecosystems.

Reciprocal subsidies link ecosystems into meta-ecosystems, but energy transfer to organisms that do not cross boundaries may create sinks, reducing reciprocal subsidy transfer. We investigated how the type of subsidy and top predator presence influenced reciprocal flows of energy, by manipulating the addition of terrestrial leaf and terrestrial insect subsidies to experimental freshwater pond mesocosms with and without predatory fish. Over 18 months, fortnightly addition of subsidies (terrestrial beetle larvae) to top-predators was crossed with monthly addition of subsidies (willow leaves) to primary consumers in mesocosms with and without top predators (upland bullies) in a 2 × 2 × 2 factorial design in four replicate blocks. Terrestrial insect subsidies increased reciprocal flows, measured as the emergence of aquatic insects out of mesocosms, but leaf subsidies dampened those effects. However, the presence of fish and snails, consumers with no terrestrial life stage, usurped and retained the energy within in the aquatic ecosystem, creating a cross-ecosystem bottleneck to energy flow. Thus, changes in species composition of donor or recipient food webs within a meta-ecosystems can alter reciprocal subsidies through cross-ecosystem bottlenecks.

Habitat patchiness, ecological connectivity and the uneven recovery of boreal stream ecosystems from an experimental drought.

Ongoing climate change is increasing the occurrence and intensity of drought episodes worldwide, including in boreal regions not previously regarded as drought prone, and where the impacts of drought remain poorly understood. Ecological connectivity is one factor that might influence community structure and ecosystem functioning post-drought, by facilitating the recovery of sensitive species via dispersal at both local (e.g. a nearby habitat patch) and regional (from other systems within the same region) scales. In an outdoor mesocosm experiment, we investigated how impacts of drought on boreal stream ecosystems are altered by the spatial arrangement of local habitat patches within stream channels, and variation in ecological connectivity with a regional species pool. We measured basal ecosystem processes underlying carbon and nutrient cycling: (a) algal biomass accrual; (b) microbial respiration; and (c) decomposition of organic matter, and sampled communities of aquatic fungi and benthic invertebrates. An 8-day drought event had strong impacts on both community structure and ecosystem functioning, including algal accrual, leaf decomposition and microbial respiration, with many of these impacts persisting even after water levels had been restored for 3.5 weeks. Enhanced connectivity with the regional species pool and increased aggregation of habitat patches also affected multiple response variables, especially those associated with microbes, and in some cases reduced the effects of drought to a small extent. This indicates that spatial processes might play a role in the resilience of communities and ecosystem functioning, given enough time. These effects were however insufficient to facilitate significant recovery in algal growth before seasonal dieback began in autumn. The limited resilience of ecosystem functioning in our experiment suggests that even short-term droughts can have extended consequences for stream ecosystems in the world's vast boreal region, and especially on the ecosystem processes and services mediated by algal biofilms.

The strength of ecological subsidies across ecosystems: a latitudinal gradient of direct and indirect impacts on food webs.

Material and energy flows among ecosystems can directly and indirectly drive ecosystem functions. Yet, how populations of consumers respond to allochthonous inputs at a macroecological scale is still unclear. Using a meta-analysis spanning several biomes, we show that the abundance of recipient populations is 36-57% larger with increased allochthonous inputs. The strength of direct effects on the recipients of these inputs as well as the indirect effects on the consumers of these recipients (i.e. ascending indirect effects) are constant across a latitudinal gradient spanning subtropical, arid, temperate, boreal and arctic ecosystems. However, indirect effect on the in situ resources of the input recipient (i.e. descending indirect effects) decreases with latitude. Our results suggest that the influence of allochthonous inputs can vary across large-scale gradients of ecosystem productivity and may be driven by the types of trophic interactions within recipient food webs.

Maximization of fertility transfers from rangeland to cropland: The contribution of control theory.

In traditional mixed farming systems, soil fertility in cropland relies on the transfer of fertility from rangeland through the transfer of manure produced by livestock that grazes in rangeland. In this work, we introduce a simple meta-ecosystem model in which the mixed farming system is represented by a cropland sub-system connected to a rangeland sub-system by nutrient fluxes. The livestock plays the role of nutrient-pump from the rangeland sub-system to the cropland sub-system. We use this model to study how spatial organization and practices of livestock management such as the control of grazing pressure and night corralling can help optimize both nutrient transfers and crop production. We argue that addressing the optimization of crop production requires different methods, depending on whether the agricultural practice in focus is constant or variable over time. We first used classical optimization methods at equilibrium to address optimization when the grazing pressure was assumed to be constant over time. Second, we address optimization for a more realistic configuration of our model, where grazing pressure was assumed to vary over the course of a year. In this case, we used methods developed in the field of the control theory. Classical methods showed the existence of an optimal level of constant grazing pressure that maximizes the transfers from rangeland to cropland, leading to the maximization of crop production. Control methods showed that by varying the grazing pressure adequately an additional gain of production is possible, with higher crop production and lower nutrient transfer from rangeland to cropland. This additional gain arises from the fact that the requirement of nutrient by crops is variable along the year. Consequently, a constant adjustment of the grazing pressure allows a better match between nutrient transfer and nutrient requirement over time, leading to a substantial gain of crop biomass. Our results provide new insights for a "smarter" management of fertility transfers leading to higher crop production with less rangeland surface.

Dispersal syndromes can impact ecosystem functioning in spatially structured freshwater populations.

Dispersal can strongly influence ecological and evolutionary dynamics. Besides the direct contribution of dispersal to population dynamics, dispersers often differ in their phenotypic attributes from non-dispersers, which leads to dispersal syndromes. The consequences of such dispersal syndromes have been widely explored at the population and community level; however, to date, ecosystem-level effects remain unclear. Here, we examine whether dispersing and resident individuals of two different aquatic keystone invertebrate species have different contributions to detrital processing, a key function in freshwater ecosystems. Using experimental two-patch systems, we found no difference in leaf consumption rates with dispersal status of the common native species Gammarus fossarum. In Dikerogammarus villosus, however, a Ponto-Caspian species now expanding throughout Europe, dispersers consumed leaf litter at roughly three times the rate of non-dispersers. Furthermore, this put the contribution of dispersing D. villosus to leaf litter processing on par with native G. fossarum, after adjusting for differences in organismal size. Given that leaf litter decomposition is a key function in aquatic ecosystems, and the rapid species turnover in freshwater habitats with range expansions of non-native species, this finding suggests that dispersal syndromes may have important consequences for ecosystem functioning.

Stoichiometric distribution models: ecological stoichiometry at the landscape extent.

Human activities are altering the fundamental geography of biogeochemicals. Yet we lack an understanding of how the spatial patterns in organismal stoichiometry affect biogeochemical processes and the tools to predict the impacts of global changes on biogeochemical processes. In this contribution we develop stoichiometric distribution models (StDMs), which allow us to map spatial structure in resource elemental composition across a landscape and evaluate spatial responses of consumers. We parameterise StDMs for a consumer-resource (moose-white birch) system and demonstrate that we can develop predictive models of resource stoichiometry across a landscape and that such models could improve our predictions of consumer space use. With results from our study system application, we argue that explicit consideration of the spatial patterns in organismal elemental composition may uncover emergent individual, population, community and ecosystem properties that are not revealed at the local extents routinely used in ecological stoichiometry. We discuss perspectives for further developments and application of StDMs to advance three emerging frameworks for spatial ecosystem ecology in an era of global change; meta-ecosystem theory, macroecological stoichiometry and remotely sensed biogeochemistry. Progress on these emerging frameworks will allow for the integration of ecological stoichiometry and individual space use and fitness.

Ecosystem flux and biotic modification as drivers of metaecosystem dynamics.

The fluxes of energy, matter, and organisms are important structuring forces of metaecosystems. Such ecosystem fluxes likely interact with environmental heterogeneity and differentially affect the diversity of multiple communities. In an aquatic mesocosm experiment, we tested how ecosystem flux and patch heterogeneity affected the diversity of bacteria, phytoplankton, and zooplankton metacommunities, and the structure and functioning of metaecosystems. We built metaecosystems consisting of three mesocosms that were either connected by flux of living organisms, organic material, and nutrients (alive ecosystem flux) or only by flux of organic material and nutrients (dead ecosystem flux). The three patches of each metaecosystem were either homogeneous or heterogeneous in nutrient loading. We found that the three groups of organisms responded differently to our treatments: flux of living organisms increased bacterial diversity irrespective of nutrient heterogeneity, while flux effects on phytoplankton diversity depended on nutrient heterogeneity, potentially indicating source-sink effects. Although zooplankton diversity was largely unaffected by our manipulations, subtle changes of community composition in response to ecosystem flux had strong effects on lower trophic levels, highlighting the importance of indirect flux effects via alterations in trophic interactions. Furthermore, differential effects of communities on the mean and spatial variability of local abiotic environments influenced the development of metaecosystem heterogeneity through time. Despite identical nutrient loading at the scale of the metaecosystem, abiotic conditions diverged between homogeneous and heterogeneous metaecosystems. For example, concentrations in dissolved organic carbon (DOC) were higher in homogeneous than heterogeneous metaecosystems, possibly because of differential responses of the algal community to local environmental conditions. Similarly, we found that flux effects on organisms translated into effects on DOC concentrations at the patch level, suggesting that flux-mediated changes in abundances of species can alter abiotic conditions. Our study shows that the dynamics of biotic and abiotic compartments of spatially structured ecosystems are intricately linked, highlighting the importance of integrating metacommunity and metaecosystem perspectives.

Emergence of nutrient co-limitation through movement in stoichiometric meta-ecosystems.

Evidence that ecosystems and primary producers are limited in their productivity by multiple nutrients has caused the traditional nutrient limitation framework to include multiple limiting nutrients. The models built to mimic these responses have invoked local mechanisms at the level of the primary producers. In this paper, we explore an alternative explanation for the emergence of co-limitation by developing a simple, stoichiometrically explicit meta-ecosystem model with two limiting nutrients, autotrophs and herbivores. Our results show that differences in movement rates for the nutrients, autotrophs and herbivores can allow for nutrient co-limitation in biomass response to emerge despite no local mechanisms of nutrient co-limitation. Furthermore, our results provide an explanation to why autotrophs show positive growth responses to nutrients despite 'nominal' top-down control by herbivores. These results suggest that spatial processes can be mechanisms for nutrient co-limitation at local and regional scales, and can help explain anomalous results in the co-limitation literature.

General relationships between consumer dispersal, resource dispersal and metacommunity diversity.

One of the central questions of metacommunity theory is how dispersal of organisms affects species diversity. Here, we show that the diversity-dispersal relationship should not be studied in isolation of other abiotic and biotic flows in the metacommunity. We study a mechanistic metacommunity model in which consumer species compete for an abiotic or biotic resource. We consider both consumer species specialised to a habitat patch, and generalist species capable of using the resource throughout the metacommunity. We present analytical results for different limiting values of consumer dispersal and resource dispersal, and complement these results with simulations for intermediate dispersal values. Our analysis reveals generic patterns for the combined effects of consumer and resource dispersal on the metacommunity diversity of consumer species, and shows that hump-shaped relationships between local diversity and dispersal are not universal. Diversity-dispersal relationships can also be monotonically increasing or multimodal. Our work is a new step towards a general theory of metacommunity diversity integrating dispersal at multiple trophic levels.

Effects of local negative feedbacks on the evolution of species within metacommunities.

Local negative feedbacks occur when the occupation of a site by a species decreases the subsequent fitness of related individuals compared to potential competitors. Such negative feedbacks can enhance diversity by changing the spatial structure of the environment. The conditions, however, involve dispersive, environmental and evolutionary processes in complex interactive ways. We introduce a model that accounts for four mechanisms: colonisation-competition-extinction ecological dynamics, evolutionary dynamics, local negative feedbacks and environmental averaging. Three qualitatively distinct dynamics are possible, one dominated by specialists, another dominated by generalists and an intermediate situation exhibiting taxon cycles. We discuss how metacommunity diversity, macro-ecological patterns and environmental patterning are linked to the three qualitative dynamics. The model provides classical shapes for morph-abundance distributions, or diversity-area relationships. Diversity can be high when specialists dominate or when taxon cycles happen. Finally, local negative feedbacks often yield fine-grain environments for taxon cycle dynamics and coarse-grain environments when generalists dominate.

Meta-ecosystem dynamics and functioning on finite spatial networks.

The addition of spatial structure to ecological concepts and theories has spurred integration between sub-disciplines within ecology, including community and ecosystem ecology. However, the complexity of spatial models limits their implementation to idealized, regular landscapes. We present a model meta-ecosystem with finite and irregular spatial structure consisting of local nutrient-autotrophs-herbivores ecosystems connected through spatial flows of materials and organisms. We study the effect of spatial flows on stability and ecosystem functions, and provide simple metrics of connectivity that can predict these effects. Our results show that high rates of nutrient and herbivore movement can destabilize local ecosystem dynamics, leading to spatially heterogeneous equilibria or oscillations across the meta-ecosystem, with generally increased meta-ecosystem primary and secondary production. However, the onset and the spatial scale of these emergent dynamics depend heavily on the spatial structure of the meta-ecosystem and on the relative movement rate of the autotrophs. We show how this strong dependence on finite spatial structure eludes commonly used metrics of connectivity, but can be predicted by the eigenvalues and eigenvectors of the connectivity matrix that describe the spatial structure and scale. Our study indicates the need to consider finite-size ecosystems in meta-ecosystem theory.

Experimental addition of marine-derived nutrients affects wildflower traits in a coastal meta-ecosystem.

Organismal movement can bring individuals, resources and novel interactions across ecosystem boundaries and into recipient habitats, thereby forming meta-ecosystems. For example, Pacific salmon ecosystems receive large marine-derived nitrogen subsidies during annual spawning events, which can have a wide range of effects on aquatic and terrestrial plant species and communities. In this study, we evaluate the effects of cross-ecosystem nutrient subsidies on terrestrial plant growth and reproduction. We conducted a large-scale field experiment with four treatments: (i) addition of a pink salmon (Oncorhynchus gorbuscha) carcass, (ii) addition of the drift seaweed rockweed (Fucus distichus), (iii) addition of both salmon + rockweed, and (iv) a control. We examined treatment effects on leaf nitrogen and fitness-associated floral traits in four common estuarine wildflower species. We found elevated leaf ∂15N in all plant species and all sampling years in treatments with salmon carcass additions but did not observe any differences in leaf per cent nitrogen. We also observed larger leaf area in two species, a context-dependent increase in floral display area in two species, and a limited increase in plant seed set in response to both salmon carcass treatments. In sum, our study suggests that marine nutrients can affect terrestrial plant growth and reproduction.