Broad Whitefish (Coregonus nasus) isotopic niches: Stable isotopes reveal diverse foraging strategies and habitat use in Arctic Alaska.

Understanding the ecological niche of some fishes is complicated by their frequent use of a broad range of food resources and habitats across space and time. Little is known about Broad Whitefish (Coregonus nasus) ecological niches in Arctic landscapes even though they are an important subsistence species for Alaska's Indigenous communities. We investigated the foraging ecology and habitat use of Broad Whitefish via stable isotope analyses of muscle and liver tissue and otoliths from mature fish migrating in the Colville River within Arctic Alaska. The range of δ13C (-31.8- -21.9‰) and δ15N (6.6-13.1‰) across tissue types and among individuals overlapped with isotope values previously observed in Arctic lakes and rivers, estuaries, and nearshore marine habitat. The large range of δ18O (4.5-10.9‰) and δD (-237.6- -158.9‰) suggests fish utilized a broad spectrum of habitats across elevational and latitudinal gradients. Cluster analysis of muscle δ13C', δ15N, δ18O, and δD indicated that Broad Whitefish occupied four different foraging niches that relied on marine and land-based (i.e., freshwater and terrestrial) food sources to varying degrees. Most individuals had isotopic signatures representative of coastal freshwater habitat (Group 3; 25%) or coastal lagoon and delta habitat (Group 1; 57%), while individuals that mainly utilized inland freshwater (Group 4; 4%) and nearshore marine habitats (Group 2; 14%) represented smaller proportions. Otolith microchemistry confirmed that individuals with more enriched muscle tissue δ13C', δD, and δ18O tended to use marine habitats, while individuals that mainly used freshwater habitats had values that were less enriched. The isotopic niches identified here represent important foraging habitats utilized by Broad Whitefish. To preserve access to these diverse habitats it will be important to limit barriers along nearshore areas and reduce impacts like roads and climate change on natural flow regimes. Maintaining these diverse connected habitats will facilitate long-term population stability, buffering populations from future environmental and anthropogenic perturbations.

Strontium isotopes reveal diverse life history variations, migration patterns, and habitat use for Broad Whitefish (Coregonus nasus) in Arctic, Alaska.

Conservation of Arctic fish species is challenging partly due to our limited ability to track fish through time and space, which constrains our understanding of life history diversity and lifelong habitat use. Broad Whitefish (Coregonus nasus) is an important subsistence species for Alaska's Arctic Indigenous communities, yet little is known about life history diversity, migration patterns, and freshwater habitat use. Using laser ablation Sr isotope otolith microchemistry, we analyzed Colville River Broad Whitefish 87Sr/86Sr chronologies (n = 61) to reconstruct movements and habitat use across the lives of individual fish. We found evidence of at least six life history types, including three anadromous types, one semi-anadromous type, and two nonanadromous types. Anadromous life history types comprised a large proportion of individuals sampled (collectively, 59%) and most of these (59%) migrated to sea between ages 0-2 and spent varying durations at sea. The semi-anadromous life history type comprised 28% of samples and entered marine habitat as larvae. Nonanadromous life history types comprised the remainder (collectively, 13%). Otolith 87Sr/86Sr data from juvenile and adult freshwater stages suggest that habitat use changed in association with age, seasons, and life history strategies. This information on Broad Whitefish life histories and habitat use across time and space will help managers and conservation planners better understand the risks of anthropogenic impacts and help conserve this vital subsistence resource.

Reverberating effects of resource exchanges in stream-riparian food webs.

Fluxes of materials or organisms across ecological boundaries, often termed "resource subsidies," directly affect recipient food webs. Few studies have addressed how such direct responses in one ecosystem may, in turn, influence the fluxes of materials or organisms to other habitats or the potential for feedback relationships to occur among ecosystems. As part of a large-scale, multi-year experiment, we evaluated the hypothesis that the input of a marine-derived subsidy results in a complex array of resource exchanges (i.e., inputs, outputs, feedbacks) between stream and riparian ecosystems as responses disperse across ecological boundaries. Moreover, we evaluated how the physical properties of resource subsidies mediated complex responses by contrasting carcasses with a pelletized salmon treatment. We found that salmon carcasses altered stream-riparian food webs by directly subsidizing multiple aquatic and terrestrial organisms (e.g., benthic insect larvae, fishes, and terrestrial flies). Such responses further influenced food webs along indirect pathways, some of which spanned land and water (e.g., subsidized fishes reduced aquatic insect emergence, with consequences for spiders and bats). Subsidy-mediated feedbacks manifested when carcasses were removed to riparian habitats where they were colonized by carrion flies, some of which fell into the stream and acted as another prey subsidy for fishes. As the effects of salmon subsidies propagated through the stream-riparian food web, the sign of consumer responses was not always positive and appeared to be determined by the outcome of trophic interactions, such that localized trophic interactions within one ecosystem mediated the export of organisms to others.

A Critical Assessment of the Ecological Assumptions Underpinning Compensatory Mitigation of Salmon-Derived Nutrients.

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.

Marine-derived nitrogen and carbon in freshwater-riparian food webs of the Copper River Delta, southcentral Alaska.

After rearing to adulthood at sea, coho salmon (Oncorhynchus kisutch) return to freshwater to spawn once and then die on or near their spawning grounds. We tested the hypothesis that spawning coho salmon return marine N and C to beaver (Castor canadensis) ponds of the Copper River Delta (CRD), Cordova, southcentral Alaska, thereby enhancing productivity of the aquatic food webs that support juvenile coho salmon. We sampled three types of pond treatment: (1) natural enrichment by spawning salmon, (2) artificial enrichment via addition of salmon carcasses and eggs, and (3) ponds with no salmon enrichment. All ponds supported juvenile coho salmon. Seasonal samples of stable isotopes revealed that juvenile coho salmon, threespine sticklebacks (Gasterosteus aculeatus), caddisfly larvae, leeches, and chironomid midge larvae were enriched with marine N and C. The aquatic vascular plants bur reed (Sparganium hyperboreum), pondweed (Potamogeton gramineus), and mare's tail (Hippuris vulgaris) were enriched with marine N only. Riparian vegetation (Sitka alder Alnus viridis ssp. sinuata and willow Salix spp.) did not show enrichment. Artificial additions of adult carcasses and eggs of coho salmon increased the delta15N and delta13C values of juvenile coho salmon. In this dynamic and hydrologically complex coastal environment, spawning coho salmon contributed marine N and C comprising 10-50% of the dietary needs of juvenile coho salmon through direct consumption of eggs and carcass material. Invertebrates that have assimilated marine N and C yield a further indirect contribution. This perennial subsidy maintains the productivity of the ecosystem of the coho salmon on the CRD.