A multidecade experiment shows that fertilization by salmon carcasses enhanced tree growth in the riparian zone.

As they return to spawn and die in their natal streams, anadromous, semelparous fishes such as Pacific salmon import marine-derived nutrients to otherwise nutrient-poor freshwater and riparian ecosystems. Diverse organisms exploit this resource, and previous studies have indicated that riparian tree growth may be enhanced by such marine-derived nutrients. However, these studies were largely inferential and did not account for all factors affecting tree growth. As an experimental test of the contribution of carcasses to tree growth, for 20 yr, we systematically deposited all sockeye salmon (Oncorhynchus nerka) carcasses (217,055 individual salmon) in the riparian zone on one bank of a 2-km-long stream in southwestern Alaska, reducing carcass accumulation on one bank and enhancing it on the other. After accounting for partial consumption and movement of carcasses by brown bears (Ursus arctos) and variation in salmon abundance and body size, we estimated that 267,620 kg of salmon were deposited on the enhanced bank and 45,200 kg on the depleted bank over the 20 yr, for a 5.9-fold difference in total mass. In 2016, we sampled needles of 84 white spruce trees (Picea glauca) the dominant riparian tree species, for foliar nitrogen (N) content and stable isotope ratios (δ15 N), and took core samples for annual growth increments. Stable isotope analysis indicated that marine-derived N was incorporated into the new growth of the trees on the enhanced bank. Analysis of tree cores indicated that in the two decades prior to our enhancement experiment, trees on the south-facing (subsequently the depleted) bank grew faster than those on the north-facing (later enhanced) bank. This difference was reduced significantly during the two decades of fertilization, indicating an effect of the carcass transfer experiment against the background of other factors affecting tree growth.

Within-lake habitat heterogeneity mediates community response to warming trends.

Climate change is rapidly altering many aquatic systems, and life history traits and physiological diversity create differences in organism responses. In addition, habitat diversity may be expressed on small spatial scales, and it is therefore necessary to account for variation among both species and locations when evaluating climate impacts on biological communities. Here, we investigated the effects of temperature and spatial heterogeneity on long-term community composition in a large boreal lake. We used a five-decade time series of water temperature and relative abundance of fish species captured in the littoral zone throughout the summer at 10 discrete locations around the lake. We applied a spatial dynamic factor analysis (SDFA) model to this time series, which estimates the sensitivity of each species to changing water temperature while accounting for spatiotemporal variation. This analysis described the trend in community composition at each sampling location in the lake, given their different trends in temperature over time. The SDFA indicated different magnitude and direction of species responses to temperature; some species increased while others decreased in abundance. The model also identified five unique trends in species abundance across sites and time, indicating residual dynamics in abundance after accounting for temperature effects. Thus, different regions in the lake have experienced different trajectories in community change associated with different rates of temperature change. These results highlight the importance of considering habitat heterogeneity in explaining and predicting future species abundances, and our model provides a means of visualizing spatially-explicit temporal variation in species' dynamics.

Climate change alters the reproductive phenology and investment of a lacustrine fish, the three-spine stickleback.

High-latitude lakes are particularly sensitive to the effects of global climate change, demonstrating earlier ice breakup, longer ice-free seasons, and increased water temperatures. Such physical changes have implications for diverse life-history traits in taxa across entire lake food webs. Here, we use a five-decade time series from an Alaskan lake to explore effects of climate change on growth and reproduction of a widely distributed lacustrine fish, the three-spine stickleback (Gasterosteus aculeatus). We used multivariate autoregressive state-space (MARSS) models to describe trends in the mean length for multiple size classes and to explore the influence of physical (date of ice breakup, surface water temperature) and biological (density of con- and heterospecifics) factors. As predicted, mean size of age 1 and older fish at the end of the growing season increased across years with earlier ice breakup and warmer temperatures. In contrast, mean size of age 0 fish decreased over time. Overall, lower fish density and warmer water temperatures were associated with larger size for all cohorts. Earlier ice breakup was associated with larger size for age 1 and older fish but, paradoxically, with smaller size of age 0 fish. To explore this latter result, we used mixing models on age 0 size distributions, which revealed an additional cohort in years with early ice breakup, lowering the mean size of age 0 fish. Moreover, early ice breakup was associated with earlier breeding, evidenced by earlier capture of age 0 fish. Our results suggest that early ice breakup altered both timing and frequency of breeding; three-spine stickleback spawned earlier and more often in response to earlier ice breakup date. While previous studies have shown the influence of changing conditions in northern lakes on breeding timing and growth, this is the first to document increased breeding frequency, highlighting another pathway by which climate change can alter the ecology of northern lakes.