Forty years of solitude: life-history divergence and behavioural isolation between laboratory lines of Drosophila melanogaster.

The study of the early stages of speciation can benefit from examination of differences between populations of known history that have been separated for a short time, such as a few thousands of generations. We asked whether two lines of Drosophila melanogaster that were isolated more than 40 years ago have evolved differences in life-history characters, or have begun to evolve behavioural or postzygotic isolation. One line, which is resistant to DDT, showed lower egg production and a shorter lifespan than a susceptible line. These differences are not a pleiotropic effect of resistance because they are not attributable to the chromosome that contains the resistance factors. The two lines have begun to become behaviourally isolated. Again, the isolation is not attributable to genes on the chromosome that contains resistance factors. The lines show only prezygotic isolation; there is no evidence of reduced fitness of F1 or F2 hybrids. These lines and others like them, should be excellent subjects for analyses of genetic changes that could lead to speciation.

Inheritance of behavioural differences between two interfertile, sympatric species, Drosophila silvestris and D. heteroneura.

The Hawaiian fly species, Drosophila silvestris and D. heteroneura, are sympatric and interfertile but show strong behavioural isolation and major differences in male aggressive behaviour and the associated morphology. As a first step in elucidating the genetic control of the differences between these species, we examined the mating and aggressive behaviour of their reciprocal F1 hybrids. The latency to the first wing vibration and the latency to copulate did not differ significantly between the parental species. However, D. heteroneura females had a very low tendency to copulate with D. silvestris males, rarely mating during the observation period. The duration of copulation also differed significantly; same-species pairs of D. silvestris had copulations that lasted about 50% longer than those of same-species pairs of D. heteroneura. The hybrids were intermediate between the parental species for both the tendency to copulate with D. silvestris males and the duration of copulation, suggesting codominance or polygenic inheritance for those traits. The aggression traits that we scored were the leg posture and wing extension during early aggression, and the leg posture and head position during escalated aggression. The parental species showed clear differences for each of these traits. The F1 hybrids resembled one parent or the other, without showing intermediate values, suggesting single-gene dominance or threshold expression of many genes for those traits. None of the courtship or aggressive traits showed X-chromosomal effects, although the head shape of hybrids is influenced by genes on the X chromosome. It is difficult to reconcile the patterns of inheritance of aggressive behaviour and the lack of an X-chromosomal effect with the hypothesis that these traits are influenced by a coadapted gene complex.

On the evolution of premating isolation after a founder event.

We present a new simple model for the evolution of premating reproductive isolation. Using this model we first analyze the level of genetic variability maintained by mutation in a large stable population. Then we consider the plausibility of the evolution of strong premating reproductive isolation after a founder event. We demonstrate that after a founder event a new adaptive combination of genes may rise to high frequencies in the presence of an old combination of genes. We compare the probabilities of speciation after a founder event with those in a stable population and with those when reproductive isolation is due to viability selection against hybrids. We argue that premating reproductive isolation is more efficient than postmating reproductive isolation in maintaining the integrity of sympatric species. This might have contributed to the pattern of stronger premating isolation than postmating isolation between closely related pairs of sympatric species.

Is sexual selection and species recognition a continuum? Mating behavior of the stalk-eyed fly Drosophila heteroneura.

If behavioral isolation between species can evolve as a consequence of sexual selection within a species, then traits that are both sexually selected and used as a criterion of species recognition by females should be identifiable. The broad male head of the Hawaiian picture-winged fly Drosophila heteroneura is a novel sexual dimorphism that may be sexually selected and involved in behavioral isolation from D. silvestris. We found that males with broad heads are more successful in sexual selection, both through female mate choice and through aggressive interactions. However, female D. heteroneura do not discriminate against hybrids on the basis of their head width. Thus, this novel trait is sexually selected but is not a major contributor to species recognition. Our methods should be applicable to other species in which behavioral isolation is a factor.

Sexual selection in relation to pest-management strategies.

The application of principles derived from the sexual selection literature can assist attempts to subvert the normal mating behavior of pests. Sexual selection encompasses both intermale competition for access to females and female choice of mates. It can operate during long-range attraction and short-range courtship, as well as after copulation. We review the major aspects of sexual selection and illustrate their application to pheromonal and SIT pest-management programs. Pheromones are important both in long-range attraction and in close-range mate choice; parapheromones may be very useful in pest management because of their influence on male mating success. Sexual selection theory provides a scheme for studying the normal mating behavior of a pest species and thus determining which attributes of the mass-reared sterile males are critical to their success with wild females. We hope that our review will suggest novel ways of attacking pests as well as encourage behavioral ecologists to study pest species.

Optimality and evolutionary genetics: complementary procedures for evolutionary analysis in behavioural ecology.

The concepts and tools of optimality and game theory are a major component of research in behavioural ecology. In contrast, the theory and practice of evolutionary, ecological, population and quantitative genetics have made less of an impact on those studying the evolution of animal behaviour. A more complete understanding of the evolution of behaviour can be achieved by pursuing research that combines optimality and genetics, thereby overcoming some of the limitations inherent in a single approach.

Coevolution of senders and receivers of sexual signals: Genetic coupling and genetic correlations.

The parallel evolution of senders and receivers of sexual signals has been a topic of research in both neuroethology and evolutionary quantitative genetics. Neuroethologists have debated whether the same physiological mechanism underlies both production and reception of a signal, and whether the same genes affect the physiology of communication in each sex. Quantitative geneticists have discussed the possibility that particular types of signals, and preferences for those types, are inherited together. Studies of communication by a variety of insect species do not provide strong support for a common physiological mechanism, but do not rule out the genetic effect. The neuroethological perspective may be of assistance in understanding the evolution of sexual communication because it offers a way to subdivide communication into units for genetic analysis.

Sexual selection and speciation: Issues raised by Hawaiian Drosophila.

The Drosophila fauna of Hawaii is extraordinarily diverse, representing about 25% of the world's described species. The most notable characteristics which differentiate the species in Hawaii are morphological and behavioral ones used in courtship and mating. These flies are excellent model species for investigating the evolution of sexually selected traits. Hypotheses regarding the associations between species formation and mating behaviour have been formulated as a result of work on this group, leading to further empirical and theoretical research.

Genetic consequences of mate choice: a quantitative genetic method for testing sexual selection theory.

To investigate whether female mate choice could be directed at male genetic quality, male chemical signals and progeny fitness were studied in the red flour beetle (Tribolium castaneum). Differences among males in the attractiveness of their pheromone to females were statistically significant. Developmental time of progeny was significantly heritable, indicating that some males have "good genes" for this trait. There was no statistically significant correlation between progeny fitness and male attractiveness. These results do not support the hypothesis that in this species the evolution of female preferences for male pheromone is adaptive.

Aggressive signal in "courtship" chirps of a gregarious cricket.

Unlike other known species of crickets, Amphiacusta maya in Central America mates in groups. Experimentally silenced males experience reduced mating success, not owing to decreased receptivity by females, but owing to increased time spent fighting with other males that persistently interrupt silent courtships. Thus, the data indicate that "courtship" chirping functions as a warning to other males, rather than as a signal to females.