Cross-ecosystem Resource Research Articles

Maintaining mechanisms of riparian invertebrate biodiversity: A review

Riparian zones, the critical ecological interfaces between terrestrial and aquatic ecosystems, are species rich habitats. However, riparian zones are seriously threatened by human activities in the world. Riparian invertebrates represent a large proportion of riparian biodiversity, perform various ecological functions, and provide an essential link between aquatic and terrestrial ecosystems. Although many studies have investigated the riparian invertebrate communities, there is lacking a comprehensive summary of maintaining mechanisms underlying riparian invertebrate diversity. This review discussed seven characteristics of riparian zones that might support high riparian invertebrate diversity: flood and drought, nutrient, microhabitat diversity, riparian vegetation, microclimate gradients, food resources and river spatial gradients. Further, we summarized the maintaining mechanisms of riparian invertebrate diversity. Disturbances of periodic flood and drought trigger the reproduction and migration of invertebrates, increase the turnover of invertebrate communities, and create suitable conditions for riparian invertebrates. Adequate nutrients support a high invertebrate diversity by increasing primary productivity of riparian habitats. Elevated microhabitat diversity provides a variety of niche space for specialist riparian invertebrates. Strong microclimate gradients provide complex and diverse habitats and thus facilitate the coexistence of aquatic and terrestrial invertebrates in riparian zones. Cross-ecosystem resource subsidies increase food availability and contribute unique food sources to riparian invertebrates. The differentiation of these factors along river longitudinal and lateral gradients provides conditions for the diversification of riparian invertebrates at a larger scale. Understanding the maintaining mechanisms of riparian invertebrate diversity is important for conservation of riparian biodiversity and integrated management of river ecosystems.

Mussels can both outweigh and interact with the effects of terrestrial to freshwater resource subsidies on littoral benthic communities

Litterfall is an important resource subsidy for lake ecosystems that primarily accumulates in littoral zones. Bivalves are abundant within littoral zones and may modify the effects of terrestrial resource subsidies through trophic interactions and engineering their surrounding habitat. Leaf inputs to lakes and freshwater mussel abundances are changing throughout the boreal ecoregion so we set out to investigate how the co-occurring benthic community might respond.We set up an in situ mesocosm experiment in Ramsey Lake, Sudbury, ON, Canada. Mesocosms contained sediments of either 5% or 35% terrestrial organic matter (tOM), into which we placed mussels (Elliptio complanata) at differing densities (0, 0.4 and 2musselsm−2, with a sham mussel treatment at 0.4musselsm−2). Over one month we recorded the sediment chemistry (dissolved organic carbon, nitrogen and phosphorus), littoral organisms (benthic algae and zooplankton) and mussel growth.At high mussel densities we recorded a 90%, 80%, 45% and 40% reduction in phosphorus, dissolved organic carbon, nitrogen and benthic diatoms, respectively, whereas at low mussel densities we observed a 3-fold increase in zooplankton. We discuss that these results were caused by a combination of bioturbation and trophic interactions. Benthic diatom concentrations were also reduced by 20% in sediments of 35% tOM, likely due to shading and competition with bacteria.Mussel growth increased at high mussel densities but was offset at high tOM, likely due to the organic matter interfering with filter feeding.Our results suggest that mussels can alter the geochemical composition of sediments and abundances of associated littoral organisms, in some cases regardless of tOM quantity. Therefore, the dominant top-down control exerted by freshwater mussels may outweigh bottom-up effects of tOM additions. Generally, our study reveals the importance of considering dominant species when studying the effects of cross-ecosystem resource fluxes.

Terrestrial support of lake food webs: Synthesis reveals controls over cross-ecosystem resource use

Widespread evidence that organic matter exported from terrestrial into aquatic ecosystems supports recipient food webs remains controversial. A pressing question is not only whether high terrestrial support is possible but also what the general conditions are under which it arises. We assemble the largest data set, to date, of the isotopic composition (δ2H, δ13C, and δ15N) of lake zooplankton and the resources at the base of their associated food webs. In total, our data set spans 559 observations across 147 lakes from the boreal to subtropics. By predicting terrestrial resource support from within-lake and catchment-level characteristics, we found that half of all consumer observations that is, the median were composed of at least 42% terrestrially derived material. In general, terrestrial support of zooplankton was greatest in lakes with large physical and hydrological connections to catchments that were rich in aboveground and belowground organic matter. However, some consumers responded less strongly to terrestrial resources where within-lake production was elevated. Our study shows that multiple mechanisms drive widespread cross-ecosystem support of aquatic consumers across Northern Hemisphere lakes and suggests that changes in terrestrial landscapes will influence ecosystem processes well beyond their boundaries.

Impacts of cross-habitat resource subsidies on ecosystems: A review.

The flux of matter, energy and nutrients across ecosystems, i.e., resource subsidy, is a fundamental attribute of ecosystems, as well as one of basic research questions in ecology. Common subsidies include leaf litter and terrestrial insects that fall into waters, the adults of aquatic insects, spawning salmon. The allocthonous input of resource subsidy can influence individual organisms, populations, communities, biodiversity and ecosystem functioning, such as enhancing individual growth, increasing species abundance and diversity, affecting community structure, enhancing secondary productivity, influencing food-chain length and food web. Due to increased human impacts on environments, especially at aspects of land use, climate change and invasive species, the influence of anthropogenic disturbance on cross-ecosystem resource subsidies will be intensified at both spacial and temporary scales, so that ecosystems will face severer threats. Accordingly, future ecological researches in this field should emphasize the following aspects: impacts of single and multiple stressors on subsidies and ecosystems, implementation of dynamic resource subsidies on ecosystem restoration and management, the dark sides of subsidy relating with pollutants, and basic ecological research on cross-ecosystem resource subsidy in tropics and sub-tropics, as well in China.

Getting quantitative about consequences of cross-ecosystem resource subsidies on recipient consumers

Most studies of cross-ecosystem resource subsidies have demonstrated positive effects on recipient consumer populations, often with very large effect sizes. However, it is important to move beyond these initial addition–exclusion experiments to consider the quantitative consequences for populations across gradients in the rates and quality of resource inputs. In our introduction to this special issue, we describe at least four potential models that describe functional relationships between subsidy input rates and consumer responses, most of them asymptotic. Here we aim to advance our quantitative understanding of how subsidy inputs influence recipient consumers and their communities. In the papers following, fish were either the recipient consumers or the subsidy as carcasses of anadromous species. Advancing general, predictive models will enable us to further consider what other factors are potentially co-limiting (e.g., nutrients, other population interactions, physical habitat, etc.) and better integrate resource subsidies into consumer–resource, biophysical dynamics models.

Invasive European bird cherry (Prunus padus) reduces terrestrial prey subsidies to urban Alaskan salmon streams

The spread of invasive species in riparian forests has the potential to affect both terrestrial and aquatic organisms linked through cross-ecosystem resource subsidies. However, this potential had not been explored in regards to terrestrial prey subsidies for stream fishes. To address this, we examined the effects of an invasive riparian tree, European bird cherry (EBC, Prunus padus), spreading along urban Alaskan salmon streams, by collecting terrestrial invertebrates present on the foliage of riparian trees, their subsidies to streams, and their consumption by juvenile coho salmon (Oncorhynchus kisutch). Riparian EBC supported four to six times less terrestrial invertebrate biomass on its foliage and contributed two to three times lower subsidies relative to native deciduous trees. This reduction in terrestrial invertebrate biomass was consistent between two watersheds over 2 years. In spite of this reduction in terrestrial prey resource input, juvenile coho salmon consumed similar levels of terrestrial invertebrates in stream reaches bordered by EBC. Although we did not see ecological effects extending to stream salmonids, reduced terrestrial prey subsidies to streams are likely to have negative consequences as EBC continues to spread.

How wide is a stream? Spatial extent of the potential “stream signature” in terrestrial food webs using meta‐analysis

The magnitude of cross-ecosystem resource subsidies is increasingly well recognized; however, less is known about the distance these subsidies travel into the recipient landscape. In streams and rivers, this distance can delimit the "biological stream width," complementary to hydro-geomorphic measures (e.g., channel banks) that have typically defined stream ecosystem boundaries. In this study we used meta-analysis to define a "stream signature" on land that relates the stream-to-land subsidy to distance. The 50% stream signature, for example, identifies the point on the landscape where subsidy resources are still at half of their maximum (in- or near-stream) level. The decay curve for these data was best fit by a negative power function in which the 50% stream signature was concentrated near stream banks (1.5 m), but a non-trivial (10%) portion of the maximum subsidy level was still found > 0.5 km from the water's edge. The meta-analysis also identified explanatory variables that affect the stream signature. This improves our understanding of ecosystem conditions that permit spatially extensive subsidy transmission, such as in highly productive, middle-order streams and rivers. Resultant multivariate models from this analysis may be useful to managers implementing buffer rules and conservation strategies for stream and riparian function, as they facilitate prediction of the extent of subsidies. Our results stress that much of the subsidy remains near the stream, but also that subsidies (and aquatic organisms) are capable of long-distance dispersal into adjacent environments, and that the effective "biological stream width" of stream and river ecosystems is often much larger than has been defined by hydro-geomorphic metrics alone. Limited data available from marine and lake sources overlap well with the stream signature data, indicating that the "signature" approach may also be applicable to subsidy spatial dynamics across other ecosystems.

Increasing donor ecosystem productivity decreases terrestrial consumer reliance on a stream resource subsidy

Because nutrient enrichment can increase ecosystem productivity, it may enhance resource flows to adjacent ecosystems as organisms cross ecosystem boundaries and subsidize predators in recipient ecosystems. Here, we quantified the biomass and abundance of aquatic emergence and terrestrial spiders in a reference and treatment stream that had been continuously enriched with nitrogen and phosphorus for 5 years. Because we previously showed that enrichment increased secondary production of stream consumers, we predicted that aquatic emergence flux would be higher in the treatment stream, subsequently increasing the biomass and abundance of terrestrial spiders. Those increases were predicted to be greatest for spiders specializing on aquatic emergence subsidies (e.g., Tetragnathidae). By adding a (15)N stable isotope tracer to both streams, we also quantified nitrogen flow from the stream into the riparian community. Emergence biomass, but not abundance, was higher in the treatment stream. The average body size of emerging adult insects and the relative dominance of Trichoptera adults were also greater in the treatment stream. However, spider biomass did not differ between streams. Spiders also exhibited substantially lower reliance on aquatic emergence nitrogen in the treatment stream. This reduced reliance likely resulted from shifts in the body size distributions and community composition of insect emergence that may have altered predator consumption efficiency in the treatment stream. Despite nutrient enrichment approximately doubling stream productivity and associated cross-ecosystem resource flows, the response of terrestrial predators depended more on the resource subsidy's characteristics that affected the predator's ability to capitalize on such increases.

Crab regulation of cross-ecosystem resource transfer by marine foraging fire ants

Permeability of boundaries in biological systems is regulated by biotic and/or abiotic factors. Despite this knowledge, the role of biotic factors in regulating resource transfer across ecosystem boundaries has received little study. Additionally, little is known about how cross-ecosystem resource transfer affects source populations. We used experiments, observations and stable isotopes, to evaluate: (1) the proportion of intertidal-foraging black fire ant (Solenopsis richteri) diet derived from marine sources, (2) how black fire ant cross-ecosystem resource transfer is altered by the dominant bioengineer in the intertidal, a burrowing crab (Neohelice granulata), (3) the top-down impact of these terrestrial ants on a marine resource, and (4) the effect of marine resources on recipient black fire ants. We found that more than 85% of the black fire ant diet is derived from marine sources, the number of intertidal foraging ants doubles in the absence of crab burrows, and that ants cause a 50% reduction in intertidal polychaetes. Also, ant mound density is three times greater adjacent to marine systems. This study reveals that cross-ecosystem foraging terrestrial ants can clearly have strong impacts on marine resources. Furthermore, ecosystem engineers that modify and occupy habitat in these ecosystem boundaries can strongly regulate the degree of cross-ecosystem resource transfer and resultant top down impacts.

Variation in active and passive resource inputs to experimental pools: mechanisms and possible consequences for food webs

1. Cross-ecosystem movements of resources, including detritus, nutrients and living prey, can strongly influence food web dynamics in recipient habitats. Variation in resource inputs is thought to be driven by factors external to the recipient habitat (e.g. donor habitat productivity and boundary conditions). However, inputs of or by ‘active’ living resources may be strongly influenced by recipient habitat quality when organisms exhibit behavioural habitat selection when crossing ecosystem boundaries. 2. To examine whether behavioural responses to recipient habitat quality alter the relative inputs of ‘active’ living and ‘passive’ detrital resources to recipient food webs, we manipulated the presence of caged predatory fish and measured biomass, energy and organic content of inputs to outdoor experimental pools of adult aquatic insects, frog eggs, terrestrial plant matter and terrestrial arthropods. 3. Caged fish reduced the biomass, energy and organic matter donated to pools by tree frog eggs by ∼70%, but did not alter insect colonisation or passive allochthonous inputs of terrestrial arthropods and plant material. Terrestrial plant matter and adult aquatic insects provided the most energy and organic matter inputs to the pools (40–50%), while terrestrial arthropods provided the least (7%). Inputs of frog egg were relatively small but varied considerably among pools and over time (3%, range = 0–20%). Absolute and proportional amounts varied by input type. 4. Aquatic predators can strongly affect the magnitude of active, but not passive, inputs and that the effect of recipient habitat quality on active inputs is variable. Furthermore, some active inputs (i.e. aquatic insect colonists) can provide similar amounts of energy and organic matter as passive inputs of terrestrial plant matter, which are well known to be important. Because inputs differ in quality and the trophic level they subsidise, proportional changes in input type could have strong effects on recipient food webs. 5. Cross-ecosystem resource inputs have previously been characterised as donor-controlled. However, control by the recipient food web could lead to greater feedback between resource flow and consumer dynamics than has been appreciated so far.

FLOODING IMPACTS ON RESPONSES OF A RIPARIAN CONSUMER TO CROSS-ECOSYSTEM SUBSIDIES

Landscape-driven processes impact the magnitude and direction of cross-ecosystem resource subsidies, but they may also control consumers' numerical and functional responses by altering habitat availability. We investigated effects of the interaction between habitat availability and subsidy level on populations of a riparian fishing spider, Dolomedes aquaticus, using a flood disturbance gradient in the Waimakariri River catchment, New Zealand. D. aquaticus predominantly eat aquatic prey as they hunt from the water surface. However, D. aquaticus biomass peaked at rivers with intermediate flood disturbance, rather than at less flood-prone rivers where the biomass of aquatic insect prey was markedly higher. Flooding positively influenced spider habitat quality, and an experimental manipulation at stable rivers indicated that unembedded cobbles, preferred D. aquaticus habitat, were a limiting factor, preventing response to the increased prey resource at stable sites. Potential terrestrial prey abundance was low, did not vary across the disturbance gradient, and is likely to have been a much smaller component of the fishing spiders' diet than aquatic insect prey. Thus landscape-driven factors not only controlled the magnitude of resource subsidies, but also influenced the ability of consumers to respond to them by altering the physical nature of the ecosystem boundary.