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Mark S. Wipfli

Publications and source records attributed to Mark S. Wipfli.

At least 37 records · Page 2Linked to original sources

Riparian defoliation by the invasive green alder sawfly influences terrestrial prey subsidies to salmon streams

Invasive species in riparian forests are unique as their effects can transcend ecosystem boundaries via stream‐riparian linkages. The green alder sawfly ( Monsoma pulveratum ) is an invasive wasp whose larvae are defoliating riparian thin‐leaf alder ( Alnus tenuifolia ) stands across southcentral Alaska. To test the hypothesis that riparian defoliation by this invasive sawfly negatively affects the flow of terrestrial prey resources to stream fishes, we sampled terrestrial invertebrates on riparian alder foliage, their subsidies to streams and their consumption by juvenile coho salmon ( Oncorhynchus kisutch ). Invasive sawflies altered the composition of terrestrial invertebrates on riparian alder foliage and as terrestrial prey subsidies to streams. Community analyses supported these findings revealing that invasive sawflies shifted the community structure of terrestrial invertebrates between seasons and levels of energy flow (riparian foliage, streams and fish). Invasive sawfly biomass peaked mid‐summer, altering the timing and magnitude of terrestrial prey subsidies to streams. Contrary to our hypothesis, invasive sawflies had no effect on the biomass of native taxa on riparian alder foliage, as terrestrial prey subsidies, or in juvenile coho salmon diets. Juvenile coho salmon consumed invasive sawflies when most abundant, but relied more on other prey types selecting against sawflies relative to their availability. Although we did not find effects of invasive sawflies extending to juvenile coho salmon in this study, these results could change as the distribution of invasive sawflies expands or as defoliation intensifies. Nevertheless, riparian defoliation by these invasive sawflies is likely having other ecological effects that merits further investigation.

Alaska

Generalist feeding strategies in Arctic freshwater fish: A mechanism for dealing with extreme environments

Generalist feeding strategies are favoured in stressful or variable environments where flexibility in ecological traits is beneficial. Species that feed across multiple habitat types and trophic levels may impart stability on food webs through the use of readily available, alternative energy pools. In lakes, generalist fish species may take advantage of spatially and temporally variable prey by consuming both benthic and pelagic prey to meet their energy demands. Using stomach content and stable isotope analyses, we examined the feeding habits of fish species in Alaska's Arctic Coastal Plain (ACP) lakes to determine the prevalence of generalist feeding strategies as a mechanism for persistence in extreme environments (e.g. low productivity, extreme cold and short growing season). Generalist and flexible feeding strategies were evident in five common fish species. Fish fed on benthic and pelagic (or nektonic) prey and across trophic levels. Three species were clearly omnivorous, feeding on fish and their shared invertebrate prey. Dietary differences based on stomach content analysis often exceeded 70%, and overlap in dietary niches based on shared isotopic space varied from zero to 40%. Metrics of community‐wide trophic structure varied with the number and identity of species involved and on the dietary overlap and niche size of individual fishes. Accumulation of energy from shared carbon sources by Arctic fishes creates redundancy in food webs, increasing likely resistance to perturbations or stochastic events. Therefore, the generalist and omnivorous feeding strategies employed by ACP fish may maintain energy flow and food web stability in extreme environments.

Ecology of Freshwater Fish

Trophic pathways supporting Arctic grayling in a small stream on the Arctic Coastal Plain, Alaska

Beaded streams are prominent across the Arctic Coastal Plain (ACP) of Alaska, yet prey flow and food web dynamics supporting fish inhabiting these streams are poorly understood. Arctic grayling ( Thymallus arcticus ) are a widely distributed upper-level consumer on the ACP and migrate into beaded streams to forage during the short 3-month open-water season. We investigated energy pathways and key prey resources that support grayling in a representative beaded stream, Crea Creek. We measured terrestrial invertebrates entering the stream from predominant riparian vegetation types, prey types supporting a range of fish size classes, and how riparian plants and fish size influenced foraging habits. We found that riparian plants influenced the quantity of terrestrial invertebrates entering Crea Creek; however, these differences were not reflected in fish diets. Prey type and size ingested varied with grayling size and season. Small grayling (<15 cm fork length (FL)) consumed mostly aquatic invertebrates early in the summer, and terrestrial invertebrates later in summer, while larger fish (>15 cm FL) foraged most heavily on ninespine stickleback ( Pungitius pungitius ) throughout the summer, indicating that grayling can be insectivorous and piscivorous, depending on size. These findings underscore the potential importance of small streams in Arctic ecosystems as key summer foraging habitats for fish. Understanding trophic pathways supporting stream fishes in these systems will help interpret whether and how petroleum development and climate change may affect energy flow and stream productivity, terrestrial–aquatic linkages and fishes in Arctic ecosystems.

Alaska

Future of Pacific salmon in the face of environmental change: Lessons from one of the world's remaining productive salmon regions

Pacific salmon Oncorhynchus spp. face serious challenges from climate and landscape change, particularly in the southern portion of their native range. Conversely, climate warming appears to be allowing salmon to expand northwards into the Arctic. Between these geographic extremes, in the Gulf of Alaska region, salmon are at historically high abundances but face an uncertain future due to rapid environmental change. We examined changes in climate, hydrology, land cover, salmon populations, and fisheries over the past 30–70 years in this region. We focused on the Kenai River, which supports world-famous fisheries but where Chinook Salmon O. tshawytscha populations have declined, raising concerns about their future resilience. The region is warming and experiencing drier summers and wetter autumns. The landscape is also changing, with melting glaciers, wetland loss, wildfires, and human development. This environmental transformation will likely harm some salmon populations while benefiting others. Lowland salmon streams are especially vulnerable, but retreating glaciers may allow production gains in other streams. Some fishing communities harvest a diverse portfolio of fluctuating resources, whereas others have specialized over time, potentially limiting their resilience. Maintaining diverse habitats and salmon runs may allow ecosystems and fisheries to continue to thrive amidst these changes.

Alaska

Top-down control of invertebrates by Ninespine Stickleback in Arctic ponds

Despite their widespread presence in northern-latitude ecosystems, the ecological role of Ninespine Stickleback Pungitius pungitius is not well understood. Ninespine Stickleback can occupy both top and intermediate trophic levels in freshwater ecosystems, so their role in food webs as a predator on invertebrates and as a forage fish for upper level consumers probably is substantial. We introduced Ninespine Sticklebacks to fishless ponds to elucidate their potential effects as a predator on invertebrate communities in Arctic lentic freshwaters. We hypothesized that Ninespine Stickleback would affect freshwater invertebrate communities in a top-down manner. We predicted that the addition of Ninespine Sticklebacks to fishless ponds would: 1) reduce invertebrate taxonomic richness, 2) decrease overall invertebrate abundance, 3) reduce invertebrate biomass, and 4) decrease average invertebrate body size. We tested our hypothesis at 2 locations by adding Ninespine Stickleback to isolated ponds and compared invertebrate communities over time between fish-addition and fishless control ponds. Ninespine Sticklebacks exerted strong top-down pressure on invertebrate communities mainly by changing invertebrate taxonomic richness and biomass and, to a lesser extent, abundance and average invertebrate size. Our results supported the hypothesis that Ninespine Stickleback may help shape lentic food webs in the Arctic.

Alaska

Measuring fish and their physical habitats: Versatile 2D and 3D video techniques with user-friendly software

Applications of video in fisheries research range from simple biodiversity surveys to three-dimensional (3D) measurement of complex swimming, schooling, feeding, and territorial behaviors. However, researchers lack a transparently developed, easy-to-use, general purpose tool for 3D video measurement and event logging. Thus, we developed a new measurement system, with freely available, user-friendly software, easily obtained hardware, and flexible underlying mathematical methods capable of high precision and accuracy. The software, VidSync, allows users to efficiently record, organize, and navigate complex 2D or 3D measurements of fish and their physical habitats. Laboratory tests showed submillimetre accuracy in length measurements of 50.8 mm targets at close range, with increasing errors (mostly <1%) at longer range and for longer targets. A field test on juvenile Chinook salmon ( Oncorhynchus tshawytscha ) feeding behavior in Alaska streams found that individuals within aggregations avoided the immediate proximity of their competitors, out to a distance of 1.0 to 2.9 body lengths. This system makes 3D video measurement a practical tool for laboratory and field studies of aquatic or terrestrial animal behavior and ecology.

Canadian Journal of Fisheries and Aquatic Sciences

A science of integration: frameworks, processes, and products in a place-based, integrative study

Integrative research is increasingly a priority within the scientific community and is a central goal for the evolving field of sustainability science. While it is conceptually attractive, its successful implementation has been challenging and recent work suggests that the move towards interdisciplinarity and transdisciplinarity in sustainability science is being only partially realized. To address this from the perspective of social-ecological systems (SES) research, we examine the process of conducting a science of integration within the Southcentral Alaska Test Case (SCTC) of Alaska-EPSCoR as a test-bed for this approach. The SCTC is part of a large, 5 year, interdisciplinary study investigating changing environments and adaptations to those changes in Alaska. In this paper, we review progress toward a science of integration and present our efforts to confront the practical issues of applying proposed integration frameworks. We: (1) define our integration framework; (2) describe the collaborative processes, including the co-development of science through stakeholder engagement and partnerships; and (3) illustrate potential products of integrative, social-ecological systems research. The approaches we use can also be applied outside of this particular framework. We highlight challenges and propose improvements for integration in sustainability science by addressing the need for common frameworks and improved contextual understanding. These insights may be useful for capacity-building for interdisciplinary projects that address complex real-world social and environmental problems.

Sustainability Science

Prey partitioning and use of insects by juvenile sockeye salmon and a potential competitor, threespine stickleback, in Afognak Lake, Alaska

Freshwater growth of juvenile sockeye salmon ( Oncorhynchus nerka ) depends upon the quality and quantity of prey and interactions with potential competitors in the foraging environment. To a large extent, knowledge about the ecology of lake-rearing juvenile sockeye salmon has emerged from studies of commercially important runs returning to deep nursery lakes, yet information from shallow nursery lakes (mean depth ≤ 10 m) is limited. We examined seasonal and ontogenetic variation in diets of juvenile sockeye salmon ( N = 219, 30–85 mm) and an abundant potential competitor, threespine stickleback ( Gasterosteus aculeatus ; N = 198, 42–67 mm), to understand their foraging ecology and potential trophic interactions in a shallow Alaska lake. This study revealed that adult insects made up 74% of all sockeye salmon diets by weight and were present in 98% of all stomachs in Afognak Lake during the summer of 2013. Diets varied temporally for all fishes, but small sockeye salmon (<60 mm) showed a distinct shift in consumption from zooplankton in early summer to adult insects in late summer. We found significant differences in diet composition between sockeye salmon and threespine stickleback and the origin of their prey indicated that they also separated their use of habitat on a fine scale; however, the two species showed overlap in size selectivity of zooplankton prey. Considering that aquatic insects can be a primary resource for juvenile sockeye salmon in Afognak Lake, we encourage the development of nursery lake carrying capacity models that include aquatic insects as a prey source for sockeye salmon.

Alaska

Surface water connectivity drives richness and composition of Arctic lake fish assemblages

Surface water connectivity can influence the richness and composition of fish assemblages, particularly in harsh environments where colonisation factors and access to seasonal refugia are required for species persistence. Studies regarding influence of connectivity on Arctic fish distributions are limited and are rarely applied to whole assemblage patterns. To increase our understanding of how surface water connectivity and related hydrologic variables influence assemblage patterns, we investigated species richness and composition of Arctic lake fishes over a large region, 8500 km 2 , of the central Arctic Coastal Plain, Alaska. We collected fish presence/non-detection data from 102 lakes and used a hierarchical multispecies occupancy framework to derive species richness and inform species composition patterns. Our mean estimate of regional richness was 12.3 (SD 0.5) species. Presence of a permanent channel connection was an overriding factor affecting species richness (mean 3.6, 95% CI 3.1&ndash;4.9), presumably driving lake colonisation potential. In lakes without a permanent channel connection, data suggest richness (mean 2.0, 95% CI 1.7&ndash;3.3) increased with the availability of in-lake winter refugia and with the potential of ephemeral connections during spring floods. Fish species functional traits and environmental faunal filters contributed to patterns of richness and assemblage composition. Composition corresponded with richness in a coherent manner, where each successive level of richness contained several discrete assemblages that showed similar responses to the environment. Lakes with permanent channel connections contained both widespread and restricted species, while the species-poor lakes that lacked a connection contained mainly widespread species. This work provides useful baseline information on the processes that drive the relations between patch connectivity and fish species richness and assemblage composition. The environmental processes that organise fish assemblages in Arctic lakes are likely to change in a warming climate.

Freshwater Biology

Production of resident fish benefits from experimental salmon subsidies via direct and indirect pathways across stream-riparian boundaries

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 bottom-up, through aquatic primary and secondary production, with little consideration of reciprocal aquatic-terrestrial 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&ndash;8 fold increases in streambed biofilms in the 2&ndash;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&ndash;140% and 44&ndash;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&ndash;24.0 g&middot;m &minus;2 &middot;yr &minus;1 of salmon carcass and <1&ndash;11.0 g&middot;m &minus;2 &middot;yr &minus;1 of analog material, which resulted in 1.2&ndash;2.9 g&middot;m &minus;2 &middot;yr &minus;1 and 0.03&ndash;1.4 g&middot;m &minus;2 &middot;yr &minus;1 of tissue produced. In addition, a feedback flux of terrestrial maggots to streams contributed 0.0&ndash;2.0 g&middot;m &minus;2 &middot;yr &minus;1 to trout production. Overall, treatments increased annual trout production by 2&ndash;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.

Ecosphere

Invasive European bird cherry disrupts stream-riparian linkages: effects on terrestrial invertebrate prey subsidies for juvenile coho salmon

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.

Canadian Journal of Fisheries and Aquatic Sciences

Trophic pathways supporting juvenile Chinook and Coho salmon in the glacial Susitna River, Alaska: patterns of freshwater, marine, and terrestrial resource use across a seasonally dynamic habitat mosaic

Contributions of terrestrial-, freshwater-, and marine-derived prey resources to stream fishes vary over time and space, altering the energy pathways that regulate production. In this study, we determined large-scale use of these resources by juvenile Chinook and coho salmon ( Oncorhynchus tshawytscha and Oncorhynchus kisutch , respectively) in the glacial Susitna River, Alaska. We resolved spatial and temporal trophic patterns among multiple macrohabitat types along a 97 km segment of the river corridor via stable isotope and stomach content analyses. Juvenile salmon were supported primarily by freshwater-derived resources and secondarily by marine and terrestrial sources. The relative contribution of marine-derived prey to rearing salmon was greatest in the fall within off-channel macrohabitats, whereas the contributions of terrestrial invertebrate prey were generally greatest during midsummer, across all macrohabitats. No longitudinal (upstream–downstream) diet pattern was discernable. These results highlight large-scale spatial and seasonal patterns of energy flow and the dynamic interplay of pulsed marine and terrestrial prey subsidies to juvenile Chinook and coho salmon in a large, complex, and relatively pristine glacial river.

Alaska

Seasonal cues of Arctic grayling movement in a small Arctic stream: the importance of surface water connectivity

In Arctic ecosystems, freshwater fish migrate seasonally between productive shallow water habitats that freeze in winter and deep overwinter refuge in rivers and lakes. How these movements relate to seasonal hydrology is not well understood. We used passive integrated transponder tags and stream wide antennae to track 1035 Arctic grayling in Crea Creek, a seasonally flowing beaded stream on the Arctic Coastal Plain, Alaska. Migration of juvenile and adult fish into Crea Creek peaked in June immediately after ice break-up in the stream. Fish that entered the stream during periods of high flow and cold stream temperature traveled farther upstream than those entering during periods of lower flow and warmer temperature. We used generalized linear models to relate migration of adult and juvenile fish out of Crea Creek to hydrology. Most adults migrated in late June &ndash; early July, and there was best support (Akaike weight = 0.46; w i ) for a model indicating that the rate of migration increased with decreasing discharge. Juvenile migration occurred in two peaks; the early peak consisted of larger juveniles and coincided with adult migration, while the later peak occurred shortly before freeze-up in September and included smaller juveniles. A model that included discharge, minimum stream temperature, year, season, and mean size of potential migrants was most strongly supported ( w i = 0.86). Juvenile migration rate increased sharply as daily minimum stream temperature decreased, suggesting fish respond to impending freeze-up. We found fish movements to be intimately tied to the strong seasonality of discharge and temperature, and demonstrate the importance of small stream connectivity for migratory Arctic grayling during the entire open-water period. The ongoing and anticipated effects of climate change and petroleum development on Arctic hydrology (e.g. reduced stream connectivity, earlier peak flows, increased evapotranspiration) have important implications for Arctic freshwater ecosystems.

Alaska

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.

Environmental Management

Climate change implications in the northern coastal temperate rainforest of North America

We synthesized an expert review of climate change implications for hydroecological and terrestrial ecological systems in the northern coastal temperate rainforest of North America. Our synthesis is based on an analysis of projected temperature, precipitation, and snowfall stratified by eight biogeoclimatic provinces and three vegetation zones. Five IPCC CMIP5 global climate models (GCMs) and two representative concentration pathways (RCPs) are the basis for projections of mean annual temperature increasing from a current average (1961&ndash;1990) of 3.2 &deg;C to 4.9&ndash;6.9 &deg;C (5 GCM range; RCP4.5 scenario) or 6.4&ndash;8.7 &deg;C (RCP8.5), mean annual precipitation increasing from 3130 mm to 3210&ndash;3400 mm (3&ndash;9 % increase) or 3320&ndash;3690 mm (6&ndash;18 % increase), and total precipitation as snow decreasing from 1200 mm to 940&ndash;720 mm (22&ndash;40 % decrease) or 720&ndash;500 mm (40&ndash;58 % decrease) by the 2080s (2071&ndash;2100; 30-year normal period). These projected changes are anticipated to result in a cascade of ecosystem-level effects including: increased frequency of flooding and rain-on-snow events; an elevated snowline and reduced snowpack; changes in the timing and magnitude of stream flow, freshwater thermal regimes, and riverine nutrient exports; shrinking alpine habitats; altitudinal and latitudinal expansion of lowland and subalpine forest types; shifts in suitable habitat boundaries for vegetation and wildlife communities; adverse effects on species with rare ecological niches or limited dispersibility; and shifts in anadromous salmon distribution and productivity. Our collaborative synthesis of potential impacts highlights the coupling of social and ecological systems that characterize the region as well as a number of major information gaps to help guide assessments of future conditions and adaptive capacity.

Alaska, British Columbia

Management and the conservation of freshwater ecosystems

Riparian areas are the terrestrial environment adjacent to water that both influences and is influenced by the aquatic feature (Gregory et al ., 1991; Naiman et al ., 2010). Riparian areas along streams provide shade, sources of wood and organic matter, contribute to bank stability, filter sediments, take up excess nutrients from groundwater inputs, and other key processes that protect freshwaters (e.g. Naiman et al ., 2010; Richardson & Danehy, 2007; Figure 9.1). Riparian areas also increase biodiversity through habitat complexity and close juxtaposition of aquatic and terrestrial environments (Quinn et al ., 2004; Naiman et al ., 2010). Alterations to riparian areas, despite their small area relative to the landscape, have disproportionate effects on habitats and fish communities (Naiman et al ., 2010; Wipfli & Baxter, 2010). Key habitat losses and alterations are derived from modification of riparian areas by reducing instream habitat complexity (Bilby & Ward, 1989; Fausch & Northcote, 1992; Naiman et al ., 2010), diminishing the productive basis of freshwater food webs (Belsky et al ., 1999; Quinn et al ., 2004), increasing nutrient, contaminant and sediment intrusion (Muscutt et al ., 1993; Daniels & Gilliam, 1996; Nguyen et al ., 1998; Waters, 1999). Riparian and freshwater ecosystems are typically tightly coupled, especially in their natural states, and the linkages that couple them frequently exert strong influence on their associated invertebrate and fish fauna (e.g. Gregory et al ., 1991; Naiman et al ., 2010). Riparian habitats, and the condition of these habitats, further plays a key role in the ecology of these fresh waters, influencing critical processes such as water, nutrient and sediment delivery and dynamics; prey resources for fish and other consumers, and other organic materials exchanged between aquatic and terrestrial habitats (Nakano et al ., 1999; Naiman et al ., 2010); light and water temperature dynamics that in turn affect food web processes and fish metabolism and growth; aquatic physical habitat (wood); and terrestrial consumers that prey upon fishes (Bisson & Bilby, 1998; Naiman et al ., 2010; Wipfli & Baxter, 2010). These processes in turn directly or indirectly influence fishes in freshwater systems (Wang et al ., 2001; Pusey & Arthington, 2003; Allan, 2004; Richardson et al ., 2010a).

Book chapter

Body size and condition influence migration timing of juvenile Arctic grayling

Freshwater fishes utilising seasonally available habitats within annual migratory circuits time movements out of such habitats with changing hydrology, although individual attributes of fish may also mediate the behavioural response to environmental conditions. We tagged juvenile Arctic grayling in a seasonally flowing stream on the Arctic Coastal Plain in Alaska and recorded migration timing towards overwintering habitat. We examined the relationship between individual migration date, and fork length (FL) and body condition index (BCI) for fish tagged in June, July and August in three separate models. Larger fish migrated earlier; however, only the August model suggested a significant relationship with BCI. In this model, 42% of variability in migration timing was explained by FL and BCI, and fish in better condition were predicted to migrate earlier than those in poor condition. Here, the majority (33%) of variability was captured by FL with an additional 9% attributable to BCI. We also noted strong seasonal trends in BCI reflecting overwinter mass loss and subsequent growth within the study area. These results are interpreted in the context of size and energetic state-specific risks of overwinter starvation and mortality (which can be very high in the Arctic), which may influence individuals at greater risk to extend summer foraging in a risky, yet prey rich, habitat. Our research provides further evidence that heterogeneity among individuals within a population can influence migratory behaviour and identifies potential risks to late season migrants in Arctic beaded stream habitats influenced by climate change and petroleum development.

Alaska