In partially migratory species, such as Oncorhynchus mykiss, the emergence of life history phenotypes is often attributed to fitness trade-offs associated with growth and survival. Fitness trade-offs can be linked to reproductive tactics that vary between the sexes, as well as the influence of environmental conditions. We found that O. mykiss outmigrants are more likely to be female in nine populations throughout western North America (grand mean 65% female), in support of the hypothesis that anadromy is more likely to benefit females. This bias was not related to migration distance or freshwater productivity, as indicated by latitude. Within one O. mykiss population we also measured the resident sex ratio and did not observe a male bias, despite a high female bias among outmigrants in that system. We provide a simulation to demonstrate the relationship between sex ratios and the proportion of anadromy and show how sex ratios could be a valuable tool for predicting the prevalence of life history types in a population.
We consider the phenomenon of partial migration which is exhibited by populations in which some individuals migrate between habitats during their lifetime, but others do not. First, using an adaptive dynamics approach, we show that partial migration can be explained on the basis of negative density dependence in the per capita fertilities alone, provided that this density dependence is attenuated for increasing abundances of the subtypes that make up the population. We present an exact formula for the optimal proportion of migrants which is expressed in terms of the vital rates of migrant and non-migrant subtypes only. We show that this allocation strategy is both an evolutionary stable strategy (ESS) as well as a convergence stable strategy (CSS). To establish the former, we generalize the classical notion of an ESS because it is based on invasion exponents obtained from linearization arguments, which fail to capture the stabilizing effects of the nonlinear density dependence. These results clarify precisely when the notion of a "weak ESS", as proposed in Lundberg (2013) for a related model, is a genuine ESS. Secondly, we use an evolutionary game theory approach, and confirm, once again, that partial migration can be attributed to negative density dependence alone. In this context, the result holds even when density dependence is not attenuated. In this case, the optimal allocation strategy towards migrants is the same as the ESS stemming from the analysis based on the adaptive dynamics. The key feature of the population models considered here is that they are monotone dynamical systems, which enables a rather comprehensive mathematical analysis.
Ecological traps can be caused when partial restoration leads organisms to make maladaptive habitat choices. One example of this is fishways (e.g., fish ladders) that provide upstream passage at dams, but are not paired with adequate downstream passage. We tested the hypothesis that attracting anadromous fishes to spawn above a dam, but blocking downstream passage of their offspring leads to an ecological trap. Using passive integrated transponder (PIT) tags, we monitored the movements of steelhead (Oncorhynchus mykiss) at a dam and fishway on the central California coast. We found that downstream passage for juveniles and kelts was limited by four factors: migration delay, loss in the reservoir, avoidance of the downstream bypass, and water depths on the spillway. Based on the spillway-passage depth-thresholds, we estimated that the ability for fish to pass downstream was limited to only half of the migration season in 55% of the past 20 years (2002-2021). Our results support the ecological trap hypothesis, which may explain why restoration using fishways has failed to produce recovery gains in this population and elsewhere.
Partial migration, the phenomenon in which animal populations are composed of both migratory and nonmigratory individuals, is widespread among migrating animals. The proportion of migrants in these populations has direct influences on population genetics and dynamics, ecosystem dynamics, mating systems, evolution, and responses to environmental change, yet there are very few studies that measure the proportion of migrants. This is because existing methods to estimate the proportion of migrants are time-consuming and expensive. In this paper, we demonstrate a new method for estimating the proportion of migrants in a population based on sex ratio measurements. Many partially migratory taxa exhibit sex-biased migration or residency, and in these cases, the sex ratios of migrants and nonmigrants are fundamentally related to the proportion of migrants in the population. We define this relationship quantitatively and show how it can be used to infer the proportion of migrants in a population through a process we term “sex-ratio balancing”. We obtain Bayesian estimates of proportion of migrants and quantify the uncertainty in these estimates with highest posterior density intervals. Lastly, we validate the sex-ratio balancing approach with a Chinook salmon (Oncorhynchus tshawytscha Walbaum in Artedi, 1792) data set. Sex-ratio balancing holds promise as a tool for quantifying partial migration and filling a key data gap about partially migratory taxa.
Abstract. Populations exhibiting partial migration consist of two groups of individuals: Those that migrate between habitats, and those that remain fixed in a single habitat. We propose several discrete-time population models to investigate the coexistence of migrants and residents. The first class of models is linear, and we distinguish two scenarios. In the first, there is a single egg pool to which both populations contribute. A fraction of the eggs is destined to become migrants, and the remainder become residents. In a second model, there are two distinct egg pools to which the two types contribute, one corresponding to residents and another to migrants. The asymptotic growth or decline in these models can be phrased in terms of the value of the basic reproduction number being larger or less than one respectively. A second class of models incorporates density dependence effects. It is assumed that increased densities in the various life history stages adversely affect the success of transitioning of individuals to subsequent stages. Here too we consider models with one or two egg pools. Although these are nonlinear models, their asymptotic dynamics can still be classified in terms of the value of a locally defined basic reproduction number: If it is less than one, then the entire population goes extinct, whereas it settles at a unique fixed point consisting of a mixture of residents and migrants, when it is larger than one. Thus, the value of the basic reproduction number can be used to predict the stable coexistence or collapse of populations exhibiting partial migration.
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