Mate choice opportunity leads to shorter offspring development time in a desert insect.

We describe indirect genetic benefits of mate choice in two allopatric populations of cactophilic Drosophila mojavensis. By manipulating mate choice opportunity, we show that greater mate choice among sexually mature adults leads to shorter offspring egg-to-adult development times; the extent of this reduction was influenced by population origin and by host plant environment. We performed multiple-choice mating trials with individually marked flies to investigate whether differential male mating success was a consequence of female choice, male interaction, or both. We demonstrate that male copulation frequency was not random and instead, was determined by female choice. Virgin females in these trials were no less discriminating than females that had been previously exposed to males. These results suggest that there are indirect benefits of female mate choice that are population and environment specific, consistent with the hypothesis of ecologically influenced 'good genes' sexual selection.

Quantitative genetics of sex ratio traits in the parasitic wasp, Nasonia vitripennis.

We detected significant parent-offspring regressions for the first sex ratio (the sex ratio produced by a female in a fresh host) and the second sex ratio (the sex ratio produced by a female in a previously parasitized host) in the parasitic wasp, Nasonia vitripennis. For both traits, estimates of the narrow-sense heritability range from approximately 0.05 to approximately 0.15 (depending on how the data are analyzed). The study population was derived from isofemale strains created from wasps captured in a single bird nest. The same population exhibited no significant parent-offspring regression for the brood sizes associated with the first and second sex ratios. There may be a significant negative parent-offspring regression for diapause proportion in the first sex ratio broods. The estimates of the genetic correlations between first and second sex ratios are positive although almost all are not significantly different from 0.0. To our knowledge, this study is the first "fine-scale" analysis of genetic variation for sex ratio traits in any species of insect. Such studies are an essential part of the assessment of the validity of claims that sex ratio traits are locally optimal.

How (not) to test an optimality model.

The controversy over the use of optimality models in the investigation of adaptation is long-standing. Nonetheless, little or no attention has been paid in this debate to the most important question to be asked about such models: how should the test of an optimality model be structured if the local optimality of the trait is to be assessed? Here we answer this question and describe how such a test can contribute to a test of adaptationism.

The comparative biology of genetic variation for conditional sex ratio behavior in a parasitic wasp, Nasonia vitripennis.

Using genetic markers, we tracked the sex ratio behavior of individual females of the parasitic wasp, Nasonia vitripennis, in foundress groups of size 1, 2, 4, 8 and 16. Comparison of 12 isofemale strains extracted from a natural population reveals significant between-strain heterogeneity of sex ratios produced in all sizes of foundress group. Under simple assumptions about population structure, this heterogeneity results in heterogeneity of fitnesses. The strains differ in their conditional sex ratio behavior (the sex ratio response of a female to foundress groups of different sizes). Females of some strains produce more males as foundress group size increases (up to size eight). Females of another strain produce more males when not alone but do not respond differentially to group size otherwise. Females of two other strains show no conditional sex ratio behavior. Females of only two strains behave differently in foundress groups of size 8 and 16. Correlation and regression analyses indicate that the strains differ significantly in their fit to the predictions of an evolutionarily stable strategy (ESS) model of conditional sex ratio behavior. Such heterogeneity contradicts the notion that females of this species possess conditonal sex ratio behavior that is optimal in the ESS sense. The results imply that this ESS model is useful but not sufficient for understanding the causal basis of the evolution of this behavior in this species. This is the first report on the sex ratio behavior of individual females in multiple foundress groups in any species of parasitic wasp. Data of this type (and not foundress group or "patch" sex ratios) are essential for testing evolutionary models that predict the sex ratio behaviors of individuals. We suggest that a test for an ESS model include the answers to two important questions: 1) is the model quantitatively accurate? and 2) is there reasonable evidence to indicate that natural selection has caused individuals to manifest the ESS behavior?

Genetic variation for sex ratio traits within a natural population of a parasitic wasp, Nasonia vitripennis.

By analyzing isofemale strains extracted from a natural population of Nasonia vitripennis, we detected variation for the sex ratios produced in fresh hosts (first sex ratios) and in previously parasitized hosts (second sex ratios). Under simple assumptions of population structure, this between-strain heterogeneity of first sex ratios results in heterogeneity of fitnesses. There is approximately ten percent difference in average fitnesses between the strains. (The fitnesses of second sex ratios are analyzed in the accompanying paper.) Average first and average second sex ratios are uncorrelated. There is significant between-female heterogeneity within some strains for first sex ratios but not for second sex ratios. In addition, the average direct-developing and diapause first sex ratios (but not second sex ratios) are significantly correlated. There are significant correlations between the direct-developing and diapause sex ratios produced by the same female. The strains differ in their effects on the sex ratio and size of another female's brood in the same host. Data on these types of variation for sex ratio traits are essential for further progress in the study of sex ratio evolution.

The comparative biology of second sex ratio evolution within a natural population of a parasitic wasp, Nasonia vitripennis.

Correlation and regression analyses indicate that isofemale strains extracted from a population of the parasitic wasp, Nasonia vitripennis, differ in the fit of their second sex ratios (those produced in previously parasitized hosts) to the predictions of the theory of optimal facultative sex ratio adjustment. Under the theory's simple assumptions about population structure, there is significant heterogeneity of fitnesses among the isofemale strains. The reasons underlying these types of heterogeneity must be understood before we can make statements about the nature of sex ratio evolution in this species. These results suggest that comparative analyses are essential for testing the qualitative predictions of optimality models.

Extinction dynamics of age-structured populations in a fluctuating environment.

We model density-independent growth of an age- (or stage-) structured population, assuming that mortality and reproductive rates fluctuate as stationary time series. Analytical formulas are derived for the distribution of time to extinction and the cumulative probability of extinction before a certain time, which are determined by the initial age distribution, and by the infinitesimal mean and variance, mu and sigma 2, of a diffusion approximation for the logarithm of total population size. These parameters can be estimated from the average life history and the pattern of environmental fluctuations in the vital rates. We also show that the distribution of time to extinction (conditional on the event) depends on the magnitude but not the sign of mu. When the environmental fluctuations in vital rates are small or moderate, the diffusion approximation gives accurate estimates of cumulative extinction probabilities obtained from computer simulations.

Genetic variation for the sex ratio in Nasonia vitripennis.

We detected genetic variation for the sex ratio in the parasitoid wasp Nasonia vitripennis by analysis of inbred lines and with an artificial selection experiment. Sex ratios differed significantly among five independently isolated lines. Furthermore, sex ratio in broods produced by single females in single hosts shifted from 80-90% female to 50-55% female in 13 to 15 generations in each of two replicate selection lines. The final sex ratios of both selection lines were significantly lower than any of the inbred line sex ratios. Backcrosses revealed that the selection response was due to nuclear genes acting through the female parent. In light of known facultative sex ratio behavior and major genes affecting sex ratio in Nasonia, our results suggest that population and individual sex ratios in this species are molded by processes at both genetic and behavioral levels.