Tidal gradients, fine-scale homing and a potential cryptic ecotype of wild spawning pink salmon (Oncorhynchus gorbuscha).

The homing behaviour of salmon is a remarkable natural phenomenon, critical for shaping the ecology and evolution of populations yet the spatial scale at which it occurs is poorly understood. This study investigated the spatial scale and mechanisms driving homing as depicted by spawning site-choice behaviour in pink salmon (Oncorhynchus gorbuscha) in Prince William Sound, Alaska. Molecular pedigree analyses of over 30,000 adult spawners in four streams revealed that pink salmon exhibit fine-scale site fidelity within a stream, returning to within <100 m of their parents. Homing behaviours were driven in part by a salinity gradient between intertidal and freshwater environments, with individuals incubated in freshwater environments more than twice as likely to spawn upstream of tidal influence than those incubated in the intertidal. Our findings challenge the traditional view that pink salmon populations are genetically and phenotypically homogenous due to their short freshwater residency as juveniles and high rates of dispersal as returning adults (i.e. straying). This study has important implications for rates of inbreeding, local adaptation and gene flow within populations, and is particularly relevant to the management of salmon hatcheries, given the high incidence of hatchery-origin pink salmon, reared in freshwater hatchery environments, that stray into wild populations of Prince William Sound.

Species distributions and the recognition of risk in restoration planning: A case study of salmonid fishes.

One of the risks faced by habitat restoration practitioners is whether habitats included in restoration planning will be used by the target species or, conversely, whether habitats excluded from restoration planning would have benefited the target species. With the goal of providing a quantitative decision-making approach that represented varying levels of risk tolerance, we used multiple probability decision thresholds (PDT) to predict the range of occurrence for three anadromous fishes (Oncorhynchus spp.) in a watershed in southwestern Washington, USA. For each species, we compared the predicted range of occurrence to the distribution used for restoration planning and quantified the amount of habitat blocked by anthropogenic barriers. Coho salmon (O. kisutch) had the broadest predicted range of occurrence (3061.6-6357.9 km; 0.75-0.25 PDT), followed by steelhead trout (O. mykiss; 1828.8-2836.8 km) and chum salmon (O. keta; 1373.9-1629.1 km). For each species, the predicted range of occurrence was similar or greater than the distribution used for restoration planning, suggesting that the current plan may exclude habitats that would benefit each species. Coho salmon had the greatest percentage of habitat blocked by anthropogenic barriers, followed by steelhead trout and chum salmon, respectively. Modeling species distributions at multiple risk-tolerance scenarios acknowledges uncertainty in restoration planning and allows practitioners to weigh the ecological benefits and budgetary constraints when considering locations for restoration. To effectively communicate restoration science to support practitioners in decision-making, we developed an R Shiny application online user interface available at: https://shiny.wdfw-fish.us/ChehalisRiverBasinSalmonidRangeOfOccurence/.

Collective navigation can facilitate passage through human-made barriers by homeward migrating Pacific salmon.

The mass migration of animals is one of the great wonders of the natural world. Although there are multiple benefits for individuals migrating in groups, an increasingly recognized benefit is collective navigation, whereby social interactions improve animals' ability to find their way. Despite substantial evidence from theory and laboratory-based experiments, empirical evidence of collective navigation in nature remains sparse. Here we used a unique large-scale radiotelemetry dataset to analyse the movements of adult Pacific salmon (Oncorhynchus sp.) in the Columbia River Basin, USA. These salmon face substantial migratory challenges approaching, entering and transiting fishways at multiple large-scale hydroelectric mainstem dams. We assess the potential role of collective navigation in overcoming these challenges and show that Chinook salmon (O. tshawytscha), but not sockeye salmon (O. nerka) locate fishways faster and pass in fewer attempts at higher densities, consistent with collective navigation. The magnitude of the density effects were comparable to major established drivers such as water temperature, and model simulations predicted that major fluctuations in population density can have substantial impacts on key quantities including mean passage time and fraction of fish with very long passage times. The magnitude of these effects indicates the importance of incorporating conspecific density and social dynamics into models of the migration process. Density effects on both ability to locate fishways and number of passage attempts have the potential to enrich our understanding of migratory energetics and success of migrating anadromous salmonids. More broadly, our work reveals a potential role of collective navigation, in at least one species, to mitigate the effects of anthropogenic barriers to animals on the move.

Signals of large scale climate drivers, hatchery enhancement, and marine factors in Yukon River Chinook salmon survival revealed with a Bayesian life history model.

Understanding how species might respond to climate change involves disentangling the influence of co-occurring environmental factors on population dynamics, and is especially problematic for migratory species like Pacific salmon that move between ecosystems. To date, debate surrounding the causes of recent declines in Yukon River Chinook salmon (Oncorhynchus tshawytscha) abundance has centered on whether factors in freshwater or marine environments control variation in survival, and how these populations at the northern extremity of the species range will respond to climate change. To estimate the effect of factors in marine and freshwater environments on Chinook salmon survival, we constructed a stage-structured assessment model that incorporates the best available data, estimates incidental marine bycatch mortality in trawl fisheries, and uses Bayesian model selection methods to quantify support for alternative hypotheses. Models fitted to two index populations of Yukon River Chinook salmon indicate that processes in the nearshore and marine environments are the most important determinants of survival. Specifically, survival declines when ice leaves the Yukon River later in the spring, increases with wintertime temperature in the Bering Sea, and declines with the abundance of globally enhanced salmon species consistent with competition at sea. In addition, we found support for density-dependent survival limitations in freshwater but not marine portions of the life cycle, increasing average survival with ocean age, and age-specific selectivity of bycatch mortality in the Bering Sea. This study underscores the utility of flexible estimation models capable of fitting multiple data types and evaluating mortality from both natural and anthropogenic sources in multiple habitats. Overall, these analyses suggest that mortality at sea is the primary driver of population dynamics, yet under warming climate Chinook salmon populations at the northern extent of the species' range may be expected to fare better than southern populations, but are influenced by foreign salmon production.

Thermal adaptation and phenotypic plasticity in a warming world: Insights from common garden experiments on Alaskan sockeye salmon.

An important unresolved question is how populations of coldwater-dependent fishes will respond to rapidly warming water temperatures. For example, the culturally and economically important group, Pacific salmon (Oncorhynchus spp.), experience site-specific thermal regimes during early development that could be disrupted by warming. To test for thermal local adaptation and heritable phenotypic plasticity in Pacific salmon embryos, we measured the developmental rate, survival, and body size at hatching in two populations of sockeye salmon (Oncorhynchus nerka) that overlap in timing of spawning but incubate in contrasting natural thermal regimes. Using a split half-sibling design, we exposed embryos of 10 families from each of two populations to variable and constant thermal regimes. These represented both experienced temperatures by each population, and predicted temperatures under plausible future conditions based on a warming scenario from the downscaled global climate model (MIROC A1B scenario). We did not find evidence of thermal local adaptation during the embryonic stage for developmental rate or survival. Within treatments, populations hatched within 1 day of each other, on average, and among treatments, did not differ in survival in response to temperature. We did detect plasticity to temperature; embryos developed 2.5 times longer (189 days) in the coolest regime compared to the warmest regime (74 days). We also detected variation in developmental rates among families within and among temperature regimes, indicating heritable plasticity. Families exhibited a strong positive relationship between thermal variability and phenotypic variability in developmental rate but body length and mass at hatching were largely insensitive to temperature. Overall, our results indicated a lack of thermal local adaptation, but a presence of plasticity in populations experiencing contrasting conditions, as well as family-specific heritable plasticity that could facilitate adaptive change.

Legacy introductions and climatic variation explain spatiotemporal patterns of invasive hybridization in a native trout.

Hybridization between invasive and native species, a significant threat to worldwide biodiversity, is predicted to increase due to climate-induced expansions of invasive species. Long-term research and monitoring are crucial for understanding the ecological and evolutionary processes that modulate the effects of invasive species. Using a large, multidecade genetics dataset (N = 582 sites, 12,878 individuals) with high-resolution climate predictions and extensive stocking records, we evaluate the spatiotemporal dynamics of hybridization between native cutthroat trout and invasive rainbow trout, the world's most widely introduced invasive fish, across the Northern Rocky Mountains of the United States. Historical effects of stocking and contemporary patterns of climatic variation were strongly related to the spread of hybridization across space and time. The probability of occurrence, extent of, and temporal changes in hybridization increased at sites in close proximity to historical stocking locations with greater rainbow trout propagule pressure, warmer water temperatures, and lower spring precipitation. Although locations with warmer water temperatures were more prone to hybridization, cold sites were not protected from invasion; 58% of hybridized sites had cold mean summer water temperatures (<11°C). Despite cessation of stocking over 40 years ago, hybridization increased over time at half (50%) of the locations with long-term data, the vast majority of which (74%) were initially nonhybridized, emphasizing the chronic, negative impacts of human-mediated hybridization. These results show that effects of climate change on biodiversity must be analyzed in the context of historical human impacts that set ecological and evolutionary trajectories.

Signals of climate, conspecific density, and watershed features in patterns of homing and dispersal by Pacific salmon.

It is widely assumed that rates of dispersal in animal populations are plastic in response to intrinsic and extrinsic cues, yet the factors influencing this plasticity are rarely known. This knowledge gap is surprising given the important role of dispersal in facilitating range shifts that may allow populations to persist in a rapidly changing global climate. We used two decades of tagging and recapture data from 19 hatchery populations of Oncorhynchus tshawytscha (Chinook salmon) in the Columbia River, USA, to quantify the effects of regional and local climate conditions, density dependence, watershed features such as area and position on the landscape, and direct anthropogenic influence on dispersal rates by adult salmon during the breeding season. We found that the probability of dispersal, termed "straying" in salmon, is plastic in'response to multiple factors and that populations showed varied responses that were largely idiosyncratic. A regional climate index (Pacific Decadal Oscillation), water temperatures in the mainstem Columbia River that was commonly experience by populations during migration, water temperatures in local subbasins unique to each population during the breeding season, migration distance, and density dependence had the strongest effects on dispersal. Patterns of dispersal plasticity in response to commonly experienced conditions were consistent with gene by environment interactions, though we are tentative about this interpretation given the domesticated history of these populations. Overall, our results warn against attempts to predict future range shifts of migratory species without considering population-specific dispersal plasticity, and also caution against the use of few populations to infer species-level patterns. Ultimately, our results provide evidence that analyses that examine the response of dispersal to single factors may be misleading.

Salmon hatchery strays can demographically boost wild populations at the cost of diversity: quantitative genetic modelling of Alaska pink salmon.

Hatcheries are vital to many salmon fisheries, with inherent risks and rewards. While hatcheries can increase the returns of adult fish, the demographic and evolutionary consequences for natural populations interacting with hatchery fish on spawning grounds remain unclear. This study examined the impacts of stray hatchery-origin pink salmon on natural population productivity and resilience. We explored temporal assortative mating dynamics using a quantitative genetic model that assumed the only difference between hatchery- and natural-origin adults was their return timing to natural spawning grounds. This model was parameterized with empirical data from an intensive multi-generational study of hatchery-wild interactions in the world's largest pink salmon fisheries enhancement program located in Prince William Sound, Alaska. Across scenarios of increasing hatchery fish presence on spawning grounds, our findings underscore a trade-off between demographic enhancement and preservation of natural population diversity. While enhancement bolstered natural population sizes towards local carrying capacities, hatchery introgression reduced variation in adult return timing by up to 20%. Results indicated that hatchery-origin alleles can rapidly assimilate into natural populations, despite the reduced fitness of hatchery fish attributable to phenotypic mismatches. These findings elucidate the potential for long-term demographic and evolutionary consequences arising from specific hatchery-wild interactions, emphasizing the need for management strategies that balance demographic enhancement with the conservation of natural diversity.

Fine-scale local adaptation in an invasive freshwater fish has evolved in contemporary time.

Adaptive evolutionary change in only a few generations can increase the ability of non-native invasive species to spread, and yet adaptive divergence is rarely assessed in recently established populations. In this study, we experimentally test for evidence of fine-scale local adaptation in juvenile survival and growth among three populations of an invasive freshwater fish with reciprocal transplants and common-garden experiments. Despite intrinsic differences in habitat quality, in two of three populations we detected evidence of increased survival in 'home' versus 'away' environments with a Bayesian occupancy model fitted to mark-recapture data. We found support for the 'local' versus 'foreign' criterion of local adaptation as 14 of 15 pairwise comparisons of performance were consistent with local adaptation (p < 0.001). Patterns in growth were less clear, though we detected evidence of location- and population-level effects. Although the agents of divergent ecological selection are not known in this system, our results combine to indicate that adaptive divergence--reflected by higher relative survival of local individuals--can occur in a small number of generations and only a few kilometres apart on the landscape.

Hybridization between genetically modified Atlantic salmon and wild brown trout reveals novel ecological interactions.

Interspecific hybridization is a route for transgenes from genetically modified (GM) animals to invade wild populations, yet the ecological effects and potential risks that may emerge from such hybridization are unknown. Through experimental crosses, we demonstrate transmission of a growth hormone transgene via hybridization between a candidate for commercial aquaculture production, GM Atlantic salmon (Salmo salar) and closely related wild brown trout (Salmo trutta). Transgenic hybrids were viable and grew more rapidly than transgenic salmon and other non-transgenic crosses in hatchery-like conditions. In stream mesocosms designed to more closely emulate natural conditions, transgenic hybrids appeared to express competitive dominance and suppressed the growth of transgenic and non-transgenic (wild-type) salmon by 82 and 54 per cent, respectively. To the best of our knowledge, this is the first demonstration of environmental impacts of hybridization between a GM animal and a closely related species. These results provide empirical evidence of the first steps towards introgression of foreign transgenes into the genomes of new species and contribute to the growing evidence that transgenic animals have complex and context-specific interactions with wild populations. We suggest that interspecific hybridization be explicitly considered when assessing the environmental consequences should transgenic animals escape to nature.

Genomics reveal the origins and current structure of a genetically depauperate freshwater species in its introduced Alaskan range.

Invasive species are a major threat to global biodiversity, yet also represent large-scale unplanned ecological and evolutionary experiments to address fundamental questions in nature. Here we analyzed both native and invasive populations of predatory northern pike (Esox lucius) to characterize landscape genetic variation, determine the most likely origins of introduced populations, and investigate a presumably postglacial population from Southeast Alaska of unclear provenance. Using a set of 4329 SNPs from 351 individual Alaskan northern pike representing the most widespread geographic sampling to date, our results confirm low levels of genetic diversity in native populations (average 𝝅 of 3.18 × 10-4) and even less in invasive populations (average 𝝅 of 2.68 × 10-4) consistent with bottleneck effects. Our analyses indicate that invasive northern pike likely came from multiple introductions from different native Alaskan populations and subsequently dispersed from original introduction sites. At the broadest scale, invasive populations appear to have been founded from two distinct regions of Alaska, indicative of two independent introduction events. Genetic admixture resulting from introductions from multiple source populations may have mitigated the negative effects associated with genetic bottlenecks in this species with naturally low levels of genetic diversity. Genomic signatures strongly suggest an excess of rare, population-specific alleles, pointing to a small number of founding individuals in both native and introduced populations consistent with a species' life history of limited dispersal and gene flow. Lastly, the results strongly suggest that a small isolated population of pike, located in Southeast Alaska, is native in origin rather than stemming from a contemporary introduction event. Although theory predicts that lack of genetic variation may limit colonization success of novel environments, we detected no evidence that a lack of standing variation limited the success of this genetically depauperate apex predator.

Acute Toxicity of Copper to Three Species of Pacific Salmon Fry in Water with Low Hardness and Low Dissolved Organic Carbon.

Proposed development of a mine within Alaska's Bristol Bay watershed (USA) has raised concerns about the potential impact of copper (Cu) on Pacific salmon (Oncorhynchus spp.). We conducted 96-h flow-through bioassays using low-hardness and low dissolved organic carbon water to determine the acute lethal toxicity of Cu to sockeye (Oncorhynchus nerka), Chinook (Oncorhynchus tshawytscha), and coho salmon (Oncorhynchus kisutch) fry. We aimed to determine Cu toxicity under field-relevant water quality conditions and to assess three methods of calculating ambient Cu criteria: the biotic ligand model (BLM), a multiple linear regression model endorsed by the US Environmental Protection Agency, and the hardness-based model currently used by the State of Alaska. The criteria generated by all models were below 20% lethal Cu concentrations by factors ranging from 2.2 to 54.3, indicating that all criteria would be protective against mortality. The multiple linear regression-based criteria were the most conservative and were comparable to BLM-based criteria. The median lethal concentrations (LC50s) for sockeye, Chinook, and coho were 35.2, 23.9, and 6.3 µg Cu/L, respectively. We also used the BLM to predict LC50s for each species. Model predictions differed from empirical LC50s by factors of 0.7 for sockeye and Chinook salmon, and 1.1 for coho salmon. These differences fell within the acceptable range of ±2, indicating the model's accuracy. We calculated critical lethal Cu accumulation values for each species to account for differing water chemistry in each bioassay; the present study revealed that coho salmon were most sensitive to Cu, followed by sockeye and Chinook salmon. Our findings underscore the importance of considering site- and species-specific factors when modeling Cu toxicity. The empirical data we present may enhance Cu risk assessments for Pacific salmon. Environ Toxicol Chem 2023;42:2440-2452. © 2023 SETAC.

Hypoxia vulnerability in the salmon watersheds of Southeast Alaska.

The frequency of dissolved oxygen depletion events (hypoxia) in coastal aquatic ecosystems has risen dramatically since the late 20th century, yet the causes and consequences of hypoxia for some culturally and economically important species remain poorly understood. In rivers, oxygen depletion can be caused by high densities of spawning Pacific salmon (Oncorhynchus spp.) consuming oxygen faster than can be replaced by reaeration. This process may be exacerbated when salmon densities are artificially inflated, such as when hatchery-origin salmon stray into rivers instead of returning to hatcheries. In Southeast Alaska, hatchery salmon production has increased rapidly since the 1970s, with over 553 million chum salmon (O. keta) and 64 million pink salmon (O. gorbuscha) released in 2021 alone. Straying is pervasive in streams with outlets <25 km from nearshore marine hatchery release sites. Using a previously ground-truthed mechanistic model of dissolved oxygen dynamics, we examined how water temperature and low-flow channel hydraulics contribute to hypoxia vulnerability. We then applied the model to predict hypoxia vulnerability for watersheds within 25 km of hatchery salmon release points, where straying salmon spawner densities are expected to be higher and promote dissolved oxygen depletion. Our model predicted that low-gradient stream reaches, regardless of water temperature, are the most prone to hypoxia due to low reaeration rates. Our spatial analysis determined that nearly 17,000 km of anadromous-accessible stream reaches are vulnerable to high densities of hatchery-origin salmon based on 2021 release sites. To our knowledge, this study is the first to map the spatial variation of hypoxia vulnerability in anadromous watersheds, identify habitat conditions most likely to promote hypoxia, and provide a repeatable analytical approach to identify hypoxia-prone stream reaches that can be updated as empirical data sets improve.

Ecotoxicology of mercury in burbot (Lota lota) from interior Alaska and insights towards human health.

Fish consumption has many health benefits, but exposure to contaminants, such as mercury (Hg), in fish tissue can be detrimental to human health. The Tanana River drainage, Alaska, USA supports the largest recreational harvest of burbot (Lota lota) in the state, yet information to evaluate the potential risks of consumption by humans is lacking. To narrow this knowledge gap, we sought to (i) quantify the concentrations of total Hg ([THg]) in burbot muscle and liver tissue and the ratio between the two tissues, (ii) assess the effect of age, length, and sex on [THg] in muscle and liver tissue, (iii) evaluate if [THg] in muscle tissue varied based on trophic information, and (iv) compare observed [THg] to consumption guidelines and statewide baseline data. The mean [THg] was 268.2 ng/g ww for muscle tissue and 62.3 ng/g ww for liver tissue. Both muscle [THg] and liver [THg] values were positively associated with fish length. Trophic information (δ15N and δ13C) was not significantly related to measured [THg] in burbot muscle, which is inconsistent with typical patterns of biomagnification observed in other fishes. All burbot sampled were within the established categories for consumption recommendations determined by the State of Alaska for women of childbearing age and children. Our results provide the necessary first step towards informed risk assessment of burbot consumption in the Tanana drainage and offer parallels to fisheries and consumers throughout the subarctic and Arctic region.

What invasive species reveal about the rate and form of contemporary phenotypic change in nature.

Biological invasions are opportunities to gain insight into fundamental evolutionary questions, because reproductive isolation and sudden alterations in selection pressures are likely to lead to rapid evolutionary change. Here I investigate the role played by invasive species in revealing the rate and form of contemporary phenotypic change in wild populations by expanding a database of more than 5,500 rates of phenotypic change from 90 species of plants and animals. Invasive species are frequently used as model organisms and thus contribute disproportionately to available rates of phenotypic change. However, the preponderance of these rates is the consequence of extensive study in a small number of species. I found mixed evidence to support the hypothesis that phenotypic change is associated with time depending on the metric of choice (i.e., darwins or haldanes). Insights from both invasive and native species provide evidence for abrupt phenotypic change and suggest that the environment plays a potentially important role in driving trait change in wild populations, although the environmental influence on the observed trajectories remains unclear. Thus, future work should continue to seek an understanding of the mechanistic underpinnings--both genetic and environmental--of how phenotypic variation allows populations to adapt to rapidly changing global environments.

Can late stage marine mortality explain observed shifts in age structure of Chinook salmon?

Chinook salmon (Oncorhynchus tshawytscha) populations have experienced widespread declines in abundance and abrupt shifts toward younger and smaller adults returning to spawn in rivers. The causal agents underpinning these shifts are largely unknown. Here we investigate the potential role of late-stage marine mortality, defined as occurring after the first winter at sea, in driving this species' changing age structure. Simulations using a stage-based life cycle model that included additional mortality during after the first winter at sea better reflected observed changes in the age structure of a well-studied and representative population of Chinook salmon from the Yukon River drainage, compared with a model estimating environmentally-driven variation in age-specific survival alone. Although the specific agents of late-stage mortality are not known, our finding is consistent with work reporting predation by salmon sharks (Lamna ditropis) and marine mammals including killer whales (Orcinus orca). Taken as a whole, this work suggests that Pacific salmon mortality after the first winter at sea is likely to be higher than previously thought and highlights the need to investigate selective sources of mortality, such as predation, as major contributors to rapidly changing age structure of spawning adult Chinook salmon.

Vulnerability of Pacific salmon to invasion of northern pike (Esox lucius) in Southcentral Alaska.

The relentless role of invasive species in the extinction of native biota requires predictions of ecosystem vulnerability to inform proactive management strategies. The worldwide invasion and range expansion of predatory northern pike (Esox lucius) has been linked to the decline of native fishes and tools are needed to predict the vulnerability of habitats to invasion over broad geographic scales. To address this need, we coupled an intrinsic potential habitat modelling approach with a Bayesian network to evaluate the vulnerability of five culturally and economically vital species of Pacific salmon (Oncorhynchus spp.) to invasion by northern pike. This study was conducted along 22,875 stream km in the Southcentral region of Alaska, USA. Pink salmon (O. gorbuscha) were the most vulnerable species, with 15.2% (2,458 km) of their calculated extent identified as "highly" vulnerable, followed closely by chum salmon (O. keta, 14.8%; 2,557 km) and coho salmon (O. kisutch, 14.7%; 2,536 km). Moreover, all five Pacific salmon species were highly vulnerable in 1,001 stream km of shared habitat. This simple to implement, adaptable, and cost-effective framework will allow prioritizing habitats for early detection and monitoring of invading northern pike.

Natural habitat change, commercial fishing, climate, and dispersal interact to restructure an Alaskan fish metacommunity.

The metacommunity concept has recently been described to account for the roles of dispersal in regulating community structure. Despite its strong theoretical basis, there exist few large-scale and long-term examples of its applicability in aquatic ecosystems. In this study we used a long-term dataset (1961-2007) on the relative abundances of the dominant limnetic fishes from two interconnected lakes to investigate the synergistic effects of naturally declining lake volume (approximately 50% in 50 years), climate variation, fishery management, and dispersal on community composition. We found a marked shift in fish community composition and variability during a period of rapid natural habitat change; however, the change was most apparent in the downstream, more stable lake of the system rather than at the site of disturbance. Multivariate analysis suggested significant shifts in community composition and variability in the downstream lake. Results indicated that the community composition in both lakes was best explained by habitat loss in the upper watershed and the number of spawning adult sockeye salmon the previous year (reflecting both natural processes and commercial fishing). Furthermore, communities exhibited site-specific responses to climatic conditions (e.g., index of the Pacific Decadal Oscillation), whereby the upper lake responded to climate within a given year and with a 1-year time lag, whereas the downstream community responded only with a 1-year lag. We attribute this difference largely to downstream dispersal and recruitment of fish from the upper lake. Thus, we suggest that the interconnected nature of the communities in this system provides a useful and large-scale example of the metacommunity concept, whereby the effects of environmental disturbance on community structure ultimately depend on the effects of these disturbances on dispersal among ecosystems.

Documentation of en route mortality of summer chum salmon in the Koyukuk River, Alaska and its potential linkage to the heatwave of 2019.

This paper documents a mass en route mortality event of adult summer chum salmon (Oncorhynchus keta) returning to the Koyukuk River, Alaska in the Yukon River basin. In response to reports from local communities, a small team of researchers (including the author) surveyed ca. 275 km of river on July 26 and 27, 2019 and counted 1,364 dead salmon. Although the total magnitude of mortality is unknown, counts from the survey certainly represent only a small fraction of the true number of fish that died. We sampled 73 carcasses to confirm death occurred prematurely prior to complete maturation and spawning, and to quantify sex and length. Visual inspection revealed a substantial fraction exhibited patterns of fungal growth consistent with secondary infections of skin lesions caused by the ubiquitous natural bacterial pathogen Flavobacterium columnare. Water temperatures during the survey averaged 17.1°C and the water was approximately 85% saturated with oxygen (ca. 8.5 mg/L), which likely contributed to the stress for upstream migrants. Evidence suggests size-selective en route mortality as female migrants that died were 2% and male migrants 5% shorter than individuals that survived to their spawning grounds on Henshaw Creek. This translates to very strong estimates of natural selection using standardized selection differentials, yet it is unclear whether selection acts on body size directly or indirectly through correlated phenotypic traits such as run timing. The mortality event likely underpins the below average returns of summer chum salmon to the Koyukuk River in 2019, suggesting an impact on spawner abundance. The future consequences of this, or potentially increasingly frequent, en route mortality events for population productivity and the extent to which genetic adaptation or adaptive phenotypic plasticity of migration behavior may facilitate persistence of these populations is unknown.