Michael P. Beakes

Steelhead Life History on California's Central Coast: Insights from a State-Dependent Model

Abstract Steelhead Oncorhynchus mykiss display a dizzying array of life history variation (including the purely resident form, rainbow trout). We developed a model for female steelhead in coastal California (close to the southern boundary of their range) in small coastal streams. We combined proximate (physiological) and ultimate (expected reproductive success) considerations to generalize the notion of a threshold size for emigration or maturity through the development of a state-dependent life history theory. The model involves strategies that depend on age, size or condition, and recent rates of change in size or condition during specific periods (decision windows) in advance of the actual smolting or spawning event. This is the first study in which such a model is fully parameterized based on data collected entirely from California steelhead populations, the majority of data coming from two watersheds the mouths of whose rivers are separated by less than 8 km along the coast of Santa Cruz County. We p...

State-dependent life history models in a changing (and regulated) environment: steelhead in the California Central Valley

We use a state dependent life history model to predict the life history strategies of female steelhead trout (Oncorhynchus mykiss) in altered environments. As a case study of a broadly applicable approach, we applied this model to the American and Mokelumne Rivers in central California, where steelhead are listed as threatened. Both rivers have been drastically altered, with highly regulated flows and translocations that may have diluted local adaptation. Nevertheless, evolutionary optimization models could successfully predict the life history displayed by fish on the American River (all anadromous, with young smolts) and on the Mokelumne River (a mix of anadromy and residency). The similar fitness of the two strategies for the Mokelumne suggested that a mixed strategy could be favored in a variable environment. We advance the management utility of this framework by explicitly modeling growth as a function of environmental conditions and using sensitivity analyses to predict likely evolutionary endpoints under changed environments. We conclude that the greatest management concern with respect to preserving anadromy is reduced survival of emigrating smolts, although large changes in freshwater survival or growth rates are potentially also important. We also demonstrate the importance of considering asymptotic size along with maximum growth rate.

Smolt Transformation in Two California Steelhead Populations: Effects of Temporal Variability in Growth

Abstract We tested the effect of temporal patterns in food supply on life history decisions in coastal steelhead Oncorhynchus mykiss irideus from a Central California coastal (CCC) population (Scott Creek) and a Northern California Central Valley (NCCV) population (upper Sacramento River basin). We manipulated growth through feeding experiments conducted from May to the following March using warm (2006 cohort) and cool (2007 cohort) temperature regimes. Survival in seawater challenges just before the time of typical juvenile emigration provided an index of steelhead smolt versus nonsmolt life history pathways. Survival varied significantly with fish size (with larger fish being more likely to survive than smaller fish) and by source population (with CCC steelhead being more likely to survive than NCCV steelhead of the same size). The timing of increased food supply (treatment group) did not significantly affect seawater survival rates in either NCCV or CCC steelhead. For both strains, the eventual survivo...

Contrasts in Habitat Characteristics and Life History Patterns of Oncorhynchus mykiss in California's Central Coast and Central Valley

Abstract Oncorhynchus mykiss exhibit high plasticity in their life history patterns. Individual life history decisions are hypothesized to result from genetic thresholds shaped by local adaptation, with variation in environmental factors influencing the trajectories of growth and condition (e.g., Fultons K, lipid content). We compared growth rates and life history patterns in two coastal creeks (Scott and Soquel) and two Central Valley (CV) rivers (American and Mokelumne) in California. The two regions differed markedly in habitat and physical factors, including hydrograph timing and amplitude, temperature regime, and food availability (measured as drift). Growth rates of coastal age-0 fish averaged 0.1 mm/d in summer–fall and 0.2 mm/d in winter–spring. Growth rates of CV fish were up to 10 times faster than those of fish on the coast and had the opposite seasonal pattern, in which growth in summer–fall was faster than that in winter–spring. Fish growth also differed between CV rivers; the mean growth ra...

Separating Intrinsic and Environmental Contributions to Growth and Their Population Consequences

Among-individual heterogeneity in growth is a commonly observed phenomenon that has clear consequences for population and community dynamics yet has proved difficult to quantify in practice. In particular, observed among-individual variation in growth can be difficult to link to any given mechanism. Here, we develop a Bayesian state-space framework for modeling growth that bridges the complexity of bioenergetic models and the statistical simplicity of phenomenological growth models. The model allows for intrinsic individual variation in traits, a shared environment, process stochasticity, and measurement error. We apply the model to two populations of steelhead trout (Oncorhynchus mykiss) grown under common but temporally varying food conditions. Models allowing for individual variation match available data better than models that assume a single shared trait for all individuals. Estimated individual variation translated into a roughly twofold range in realized growth rates within populations. Comparisons between populations showed strong differences in trait means, trait variability, and responses to a shared environment. Together, individual- and population-level variation have substantial implications for variation in size and growth rates among and within populations. State-dependent life-history models predict that this variation can lead to differences in individual life-history expression, lifetime reproductive output, and population life-history diversity.

Emergent stability in a large, free‐flowing watershed

While it is widely recognized that financial stock portfolios can be stabilized through diverse investments, it is also possible that certain habitats can function as natural portfolios that stabilize ecosystem processes. Here we propose and examine the hypothesis that free-flowing river networks act as such portfolios and confer stability through their integration of upstream geological, hydrological, and biological diversity. We compiled a spatially (142 sites) and temporally (1980-present) extensive data set on fisheries, water flows, and temperatures, from sites within one of the largest watersheds in the world that remains without dams on its mainstem, the Fraser River, British Columbia, Canada. We found that larger catchments had more stable fisheries catches, water flows, and water temperatures than smaller catchments. These data provide evidence that free-flowing river networks function as hierarchically nested portfolios with stability as an emergent property. Thus, free-flowing river networks can represent a natural system for buffering variation and extreme events.

Wildfire and the effects of shifting stream temperature on salmonids

The frequency and magnitude of wildfires in North America have increased by four-fold over the last two decades. However, the impacts of wildfires on the thermal environments of freshwaters, and potential effects on coldwater fishes are incompletely understood. We examined the short-term effects of a wildfire on temperatures and Steelhead/Rainbow Trout (Oncorhynchus mykiss) bioenergetics and distribution in a California coastal stream. One year after the wildfire, mean daily stream temperatures were elevated by up to 0.6°C in burned compared to unburned pools. Among burned pools, light flux explained over 85% of the variation in altered stream temperatures, and 76% of the variation in light flux was explained by an index of burn severity based on proximity of the pool to burned streamside. We estimated that salmonids of variable sizes inhabiting burned pools had to consume between 0.3–264.3 mg of additional prey over 48 days to offset the 0.01–6.04 kJ increase in metabolic demand during the first post-fire summer. However, stomach content analysis showed that fish in the burned region were consuming relatively little prey and significantly less than fish in the reference region. Presumably due to starvation, mortality, or emigration, we found a significant negative relationship between the change in total salmonid biomass over the post-fire summer and the average energy costs (kJ·g−1·day−1) within a burned pool. This study demonstrates that wildfire can generate thermal heterogeneity in aquatic ecosystems and drive short-term increases in stream temperature, exacerbating bioenergetically stressful seasons for coldwater fishes.

Using scale characteristics and water temperature to reconstruct growth rates of juvenile steelhead Oncorhynchus mykiss

Juvenile steelhead Oncorhynchus mykiss from a northern California Central Valley population were reared in a controlled laboratory experiment. Significantly different rates of growth were observed among fish reared under two ration treatments and three temperature treatments (8, 14 and 20°C). Wider circulus spacing and faster deposition was associated with faster growth. For the same growth rate, however, circulus spacing was two-fold wider and deposited 36% less frequently in the cold compared to the hot temperature treatment. In a multiple linear regression, median circulus spacing and water temperature accounted for 68% of the variation in observed O. mykiss growth. These results corroborate previous research on scale characteristics and growth, while providing novel evidence that highlights the importance of water temperature in these relationships. Thus, this study establishes the utility of using scale analysis as a relatively non-invasive method for inferring growth in salmonids.