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.
Abstract. Artificial additions of nutrients of differing forms such as salmon carcasses and analog pellets (i.e. pasteurized fishmeal) have been proposed as a means of stimulating aquatic productivity and enhancing populations of anadromous and resident fishes. Nutrient mitigation to enhance fish production in stream ecosystems assumes that the central pathway by which effects occur is bottomup, through aquatic primary and secondary production, with little consideration of reciprocal aquaticterrestrial pathways. The net outcome (i.e. bottom-up vs. top-down) of adding salmon-derived materials to streams depend on whether or not these subsidies indirectly intensify predation on in situ prey via increases in a shared predator or alleviate such predation pressure. We conducted a 3-year experiment across nine tributaries of the N. Fork Boise River, Idaho, USA, consisting of 500-m stream reaches treated with salmon carcasses (n = 3), salmon carcass analog (n = 3), and untreated control reaches (n = 3). We observed 2-8 fold increases in streambed biofilms in the 2-6 weeks following additions of both salmon subsidy treatments in years 1 and 2 and a 1.5-fold increase in standing crop biomass of aquatic invertebrates to carcass additions in the second year of our experiment. The consumption of benthic invertebrates by stream fishes increased 110-140% and 44-66% in carcass and analog streams in the same time frame, which may have masked invertebrate standing crop responses in years 3 and 4. Resident trout directly consumed 10.0-24.0 g·m −2 ·yr −1 of salmon carcass and <1-11.0 g·m −2 ·yr −1 of analog material, which resulted in 1.2-2.9 g·m −2 ·yr −1 and 0.03-1.4 g·m −2 ·yr −1 of tissue produced. In addition, a feedback flux of terrestrial maggots to streams contributed 0.0-2.0 g·m −2 ·yr −1 to trout production. Overall, treatments increased annual trout production by 2-3 fold, though density and biomass were unaffected. Our results indicate the strength of bottom-up and top-down responses to subsidy additions was asymmetrical, with top-down forces masking bottom-up effects that required multiple years to manifest. The findings also highlight the need for nutrient mitigation programs to consider multiple pathways of energy and nutrient flow to account for the complex effects of salmon subsidies in stream-riparian ecosystems.
We critically evaluate some of the key ecological assumptions underpinning the use of nutrient replacement as a means of recovering salmon populations and a range of other organisms thought to be linked to productive salmon runs. These assumptions include: (1) nutrient mitigation mimics the ecological roles of salmon, (2) mitigation is needed to replace salmon-derived nutrients and stimulate primary and invertebrate production in streams, and (3) food resources in rearing habitats limit populations of salmon and resident fishes. First, we call into question assumption one because an array of evidence points to the multi-faceted role played by spawning salmon, including disturbance via redd-building, nutrient recycling by live fish, and consumption by terrestrial consumers. Second, we show that assumption two may require qualification based upon a more complete understanding of nutrient cycling and productivity in streams. Third, we evaluate the empirical evidence supporting food limitation of fish populations and conclude it has been only weakly tested. On the basis of this assessment, we urge caution in the application of nutrient mitigation as a management tool. Although applications of nutrients and other materials intended to mitigate for lost or diminished runs of Pacific salmon may trigger ecological responses within treated ecosystems, contributions of these activities toward actual mitigation may be limited.
Bighead carp (Hypophthalmichthys nobilis) are an invasive planktivore that can greatly deplete planktonic resources. Due to the inefficient conversion of food into fish tissue, large portions of consumed materials are egested and shunted to benthic habitats. We explored how bighead carp alter pools of organic matter between planktonic and benthic habitats, and across ecosystem boundaries. Here, we report evidence from a manipulative experiment demonstrating that bighead carp greatly reapportion pools of organic matter from planktonic to benthic habitats to such a degree that additional effects propagated across ecological boundaries into terrestrial ecosystems. Strong direct consumption by bighead carp reduced filamentous algae, biomass and production of zooplankton, and production of a native planktivorous fish within planktonic habitats. Reduced herbivory indirectly increased phytoplankton (chlorophyll a). Direct consumption of organic matter by bighead carp supported high carp production and concomitant losses of materials due to egestion. Perhaps in response to organic matter subsidies provided by fish egestion, ponds having bighead carp had higher standing crop biomass of Chironomidae larvae, as well as cross-boundary fluxes of their adult life stage. In contrast, we detected reduced cross-boundary fluxes of adult Chaoboridae midges in ponds having bighead carp. Consideration of bighead carp as mediators of organic matter exchanges provides a clearer framework for predicting the direct and extended impacts of these invasive planktivores in freshwater ecosystems. The perception of bighead carp must evolve beyond competitors for planktonic resources, to mediators and processors of nutrients and energy within and across ecosystems.
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