Null models confirm nest site fidelity by male smallmouth bass, Micropterus dolomieu.

BACKGROUND: Many animals appear to preferentially renest in proximity to a site they previously occupied. Evidence of nest fidelity is often inferred from a right skewed distribution of distances between the nests of individuals that breed in two consecutive reproduction episodes, where many individuals nest some arbitrarily close distance to their prior nest and others, in the extended right tail of the distribution, nest far from the nest they previously occupied. Because right skewed distributions of inter-nest distances can arise even when individuals choose nest locations randomly, however, such inferences are prone to error. The importance of null models-used to generate patterns of individual inter-nest distances by processes that do not involve site attachment-for inferences about site fidelity has been known for decades but is still often unappreciated or ignored. METHODS: The right skewed distributions of inter-nest distances observed in two earlier studies of male smallmouth bass (Micropterus dolomieu) suggest prima facie that males exhibit nest site fidelity between annual reproduction episodes, but patterns of inter-nest distances have yet to be compared to an adequate null model. Here, we evaluate the nest site fidelity of marked male M. dolomieu in a decade-long dataset, where we apply a randomization procedure based on the rencontre probability problem to generate null models. Eight observed distributions of individual, annual inter-nest distances are compared to a year-specific null model to determine whether random processes are sufficient to explain the observed distributions of inter-nest distances. RESULTS: Through contrasts between observed annual inter-nest distances and results derived from null models that imposed realistic constraints on behavior, we show that some males were undoubtedly nest-site faithful. To reinforce the utility of null models and to make these kinds of models more accessible, we also provide a supplemental tutorial. The tutorial illustrates how random site choices, subject to common ecological and behavioral constraints, and even how distance is measured, can produce patterns of inter-nest distances that falsely imply nest site fidelity, or a lack of fidelity. The R code needed to reproduce these null models is included. The inference errors evident in our examples generalize to other forms of site fidelity, such as the apparent patch fidelity of certain sea bird foragers. CONCLUSIONS: The comparisons of observed distributions of inter-nest distances with those generated by null models imply that, as suggested in prior studies, male M. dolomieu indeed exhibit annual nest site fidelity. Procedures like those we apply are necessary first steps in analyses when distributions of distances between the nests of individuals in consecutive reproduction episodes are used to infer nest-site fidelity. Why male M. dolomieu are site faithful is a question yet to be answered.

Negative maternal or paternal effects on tactic inheritance under a conditional strategy.

Alternative behavioral and life-history tactics are common in animal populations. The conditional strategy model provides a powerful explanation for the evolution and persistence of such tactics, as it allows alternative tactics to be perpetuated even if there is tactic inheritance and tactics yield unequal mean fitness. In many biological systems negative maternal or paternal effects complicate the inheritance of condition and, hence, the inheritance of alternative tactics. Indeed, the inheritance of condition may result in the alternation of tactics across generations. In this paper, we show that the conditional strategy is robust to these effects on progeny condition. There is a unique and stable proportion of tactics under standard inheritance and unequal tactic fitness, and these two important properties of the conditional strategy hold even if negative maternal or paternal effects on progeny condition cause tactics to alternate across generations. However, the dynamics of tactic proportions pursuant to a perturbation of the equilibrium tactic proportions depend on the form of tactic inheritance. An application of our theoretical results to data from a population of smallmouth bass (Micropterus dolomieu) in which negative paternal effects dictate progeny condition reveals that age at first reproduction in males alternates regularly across generations. Furthermore, the model indicates that the population would return rapidly to equilibrium if the proportions of males that mature early or late in life were perturbed from the equilibrium within the system. This example shows how the model of the conditional strategy can be used to gain insight into tactic dynamics in situations where some of the model parameters are difficult or impossible to measure empirically.

SEXUAL SELECTION AND FITNESS VARIATION IN A POPULATION OF SMALLMOUTH BASS, MICROPTERUS DOLOMIEUI (PISCES: CENTRARCHIDAE).

Monogamy is often presumed to constrain mating variance and restrict the action of sexual selection. We examined the reproductive patterns of a monogamous population of smallmouth bass (Micropterus dolomieui), and attempted to identify sources of within-season fitness variation among females and known-age males. Many males did not acquire a nest site, and many territorial males were unsuccessful in acquiring a mate. The likelihood that territorial males mated depended on several aspects of nest sites. Mated males of age three were larger than the average size of age-three males in the population. The mean sizes of age-four and age-five mated males were not different from the average of same-age males in the population. Thus, selection resulting from the acquisition of a mate favored large size among only age-three males. Timing of nest construction and breeding among territorial males was negatively related to male size and did not depend on male age after taking male size into account. Indirect evidence (numbers of eggs deposited in nests) suggests that the timing of spawning among females was also negatively related to female size. Fertility selection favored early reproduction within the season by males of all ages, but large male size was favored among only age-four males. The combined early breeding of fecund females and female mate choice of large males may explain the positive correlation between the size of age-four males and the number of eggs acquired. Despite large differences of female fecundity, however, the variance of relative mate number contributed about two times more than the variance of relative fertility among females to the total variance of relative fitness within each sex.