Phenological shifts and mismatch with marine productivity vary among Pacific salmon species and populations.

Global climate change is shifting the timing of life-cycle events, sometimes resulting in phenological mismatches between predators and prey. Phenological shifts and subsequent mismatches may be consistent across populations, or they could vary unpredictably across populations within the same species. For anadromous Pacific salmon (Oncorhynchus spp.), juveniles from thousands of locally adapted populations migrate from diverse freshwater habitats to the Pacific Ocean every year. Both the timing of freshwater migration and ocean arrival, relative to nearshore prey (phenological match/mismatch), can control marine survival and population dynamics. Here we examined phenological change of 66 populations across six anadromous Pacific salmon species throughout their range in western North America with the longest time series spanning 1951-2019. We show that different salmon species have different rates of phenological change but that there was substantial within-species variation that was not correlated with changing environmental conditions or geographic patterns. Moreover, outmigration phenologies have not tracked shifts in the timing of marine primary productivity, potentially increasing the frequency of future phenological mismatches. Understanding population responses to mismatches with prey are an important part of characterizing overall population-specific climate vulnerability.

Stream and ocean hydrodynamics mediate partial migration strategies in an amphidromous Hawaiian goby.

Partial migration strategies, in which some individuals migrate but others do not, are widely observed in populations of migratory animals. Such patterns could arise via variation in migratory behaviors made by individual animals, via genetic variation in migratory predisposition, or simply by variation in migration opportunities mediated by environmental conditions. Here we use spatiotemporal variation in partial migration across populations of an amphidromous Hawaiian goby to test whether stream or ocean conditions favor completing its life cycle entirely within freshwater streams rather than undergoing an oceanic larval migration. Across 35 watersheds, microchemical analysis of otoliths revealed that most adult Awaous stamineus were freshwater residents (62% of n = 316 in 2009, 83% of n = 274 in 2011), but we found considerable variation among watersheds. We then tested the hypothesis that the prevalence of freshwater residency increases with the stability of stream flows and decreases with the availability of dispersal pathways arising from ocean hydrodynamics. We found that streams with low variation of daily discharge were home to a higher incidence of freshwater residents in each survey year. The magnitude of the shift in freshwater residency between survey years was positively associated with predicted interannual variability in the success of larval settlement in streams on each island based on passive drift in ocean currents. We built on these findings by developing a theoretical model of goby life history to further evaluate whether mediation of migration outcomes by stream and ocean hydrodynamics could be sufficient to explain the range of partial migration frequency observed across populations. The model illustrates that the proportion of larvae entering the ocean and differential survival of freshwater-resident versus ocean-going larvae are plausible mechanisms for range-wide shifts in migration strategies. Thus, we propose that hydrologic variation in both ocean and stream environments contributes to spatiotemporal variation in the prevalence of migration phenotypes in A. stamineus. Our empirical and theoretical results suggest that the capacity for partial migration could enhance the persistence of metapopulations of diadromous fish when confronted with variable ocean and stream conditions.

Invasion and rapid adaptation of guppies (Poecilia reticulata) across the Hawaiian Archipelago.

How much does natural selection, as opposed to genetic drift, admixture, and gene flow, contribute to the evolution of invasive species following introduction to a new environment? Here we assess how evolution can shape biological invasions by examining population genomic variation in non-native guppies (Poecilia reticulata) introduced to the Hawaiian Islands approximately a century ago. By examining 18 invasive populations from four Hawaiian islands and four populations from the native range in northern South America, we reconstructed the history of introductions and evaluated population structure as well as the extent of ongoing gene flow across watersheds and among islands. Patterns of differentiation indicate that guppies have developed significant population structure, with little natural or human-mediated gene flow having occurred among populations following introduction. Demographic modeling and admixture graph analyses together suggest that guppies were initially introduced to O'ahu and Maui and then translocated to Hawai'i and Kaua'i. We detected evidence for only one introduction event from the native range, implying that any adaptive evolution in introduced populations likely utilized the genetic variation present in the founding population. Environmental association tests accounting for population structure identified loci exhibiting signatures of adaptive variation related to predators and landscape characteristics but not nutrient regimes. When paired with high estimates of effective population sizes and detectable population structure, the presence of environment-associated loci supports the role of natural selection in shaping contemporary evolution of Hawaiian guppy populations. Our findings indicate that local adaptation may engender invasion success, particularly in species with life histories that facilitate rapid evolution. Finally, evidence of low gene flow between populations suggests that removal could be an effective approach to control invasive guppies across the Hawaiian archipelago.

Adaptive capacity at the northern front: sockeye salmon behaviourally thermoregulate during novel exposure to warm temperatures.

As climate change increases maximal water temperatures, behavioural thermoregulation may be crucial for the persistence of coldwater fishes, such as salmonids. Although myriad studies have documented behavioural thermoregulation in southern populations of salmonids, few if any have explored this phenomenon in northern populations, which are less likely to have an evolutionary history of heat stress, yet are predicted to experience substantial warming. Here, we treated a rare heat wave as a natural experiment to test whether wild sockeye salmon (Oncorhynchus nerka) at the northern extent of their primary range (60° latitude) can thermoregulate in response to abnormally high thermal conditions. We tagged adult sockeye salmon with temperature loggers as they staged in a lake epilimnion prior to spawning in small cold streams (n = 40 recovered loggers). As lake surface temperatures warmed to physiologically suboptimal levels (15-20°C), sockeye salmon thermoregulated by moving to tributary plumes or the lake metalimnion. A regression of fish body temperature against lake surface temperature indicated that fish moved to cooler waters when the epilimnion temperature exceeded ~12°C. A bioenergetics model suggested that the observed behaviour reduced daily metabolic costs by as much as ~50% during the warmest conditions (18-20°C). These results provide rare evidence of cool-seeking thermoregulation at the poleward extent of a species range, emphasizing the potential ubiquity of maximal temperature constraints and the functional significance of thermal heterogeneity for buffering poikilotherms from climate change.

Riding the crimson tide: mobile terrestrial consumers track phenological variation in spawning of an anadromous fish.

When resources are spatially and temporally variable, consumers can increase their foraging success by moving to track ephemeral feeding opportunities as these shift across the landscape; the best examples derive from herbivore-plant systems, where grazers migrate to capitalize on the seasonal waves of vegetation growth. We evaluated whether analogous processes occur in watersheds supporting spawning sockeye salmon (Oncorhynchus nerka), asking whether seasonal activities of predators and scavengers shift spatial distributions to capitalize on asynchronous spawning among populations of salmon. Both glaucous-winged gulls and coastal brown bears showed distinct shifts in their spatial distributions over the course of the summer, reflecting the shifting distribution of spawning sockeye salmon, which was associated with variation in water temperature among spawning sites. By tracking the spatial and temporal variation in the phenology of their principal prey, consumers substantially extended their foraging opportunity on a superabundant, yet locally ephemeral, resource. Ecosystem-based fishery management efforts that seek to balance trade-offs between fisheries and ecosystem processes supported by salmon should, therefore, assess the importance of life-history variation, particularly in phenological traits, for maintaining important ecosystem functions, such as providing marine-derived resources for terrestrial predators and scavengers.

Centennial-scale fluctuations and regional complexity characterize Pacific salmon population dynamics over the past five centuries.

Observational data from the past century have highlighted the importance of interdecadal modes of variability in fish population dynamics, but how these patterns of variation fit into a broader temporal and spatial context remains largely unknown. We analyzed time series of stable nitrogen isotopes from the sediments of 20 sockeye salmon nursery lakes across western Alaska to characterize temporal and spatial patterns in salmon abundance over the past ∼500 y. Although some stocks varied on interdecadal time scales (30- to 80-y cycles), centennial-scale variation, undetectable in modern-day catch records and survey data, has dominated salmon population dynamics over the past 500 y. Before 1900, variation in abundance was clearly not synchronous among stocks, and the only temporal signal common to lake sediment records from this region was the onset of commercial fishing in the late 1800s. Thus, historical changes in climate did not synchronize stock dynamics over centennial time scales, emphasizing that ecosystem complexity can produce a diversity of ecological responses to regional climate forcing. Our results show that marine fish populations may alternate between naturally driven periods of high and low abundance over time scales of decades to centuries and suggest that management models that assume time-invariant productivity or carrying capacity parameters may be poor representations of the biological reality in these systems.

A coherent signature of anthropogenic nitrogen deposition to remote watersheds of the Northern Hemisphere.

Humans have more than doubled the amount of reactive nitrogen (Nr) added to the biosphere, yet most of what is known about its accumulation and ecological effects is derived from studies of heavily populated regions. Nitrogen (N) stable isotope ratios ((15)N:(14)N) in dated sediments from 25 remote Northern Hemisphere lakes show a coherent signal of an isotopically distinct source of N to ecosystems beginning in 1895 ± 10 years (±1 standard deviation). Initial shifts in N isotope composition recorded in lake sediments coincide with anthropogenic CO(2) emissions but accelerate with widespread industrial Nr production during the past half century. Although current atmospheric Nr deposition rates in remote regions are relatively low, anthropogenic N has probably influenced watershed N budgets across the Northern Hemisphere for over a century.