Sex ratio theory predicts that malaria parasites should bias gametocyte production toward female cells in single-clone infections because they will experience complete inbreeding of parasite gametes within the vector. A higher proportion of male gametocytes is favoured under conditions that reduce success of male gametes at reaching females such as low gametocyte density or attack of the immune system later in the infection. Recent experimental studies reveal genetic variation for gametocyte sex ratio in single-clone infections. We examined these issues with a study of experimental single-clone infections for the lizard malaria parasite Plasmodium mexicanum in its natural host. Gametocyte sex ratios of replicate single-clone infections were determined over a period of 3–4 months. Sex ratios were generally female biased, but not as strongly as expected under simple sex ratio theory. Gametocyte density was not related to sex ratio, and male gametocytes did not become more common later in infections. The apparent surplus of male gametocytes could be explained if male fecundity is low in this parasite, or if rapid clotting of the lizard blood reduces male gamete mobility. There was also a significant clone effect on sex ratio, suggesting genetic variation for some life-history trait, possibly male fecundity.

Studies on mating ecology and sex allocation in fig-parasitizing wasps ovipositing from outside the fig have given valuable insights into known factors that are responsible for the theory of sex ratio. Similarly, internally ovipositing fig-parasitizing wasps and fig-pollinating wasps provide interesting models for comparative analysis. In addition to the fig-pollinating wasp Eupristina sp., we found that Ficus curtipes hosts two species of internally ovipositing fig-parasitizing wasps: D. yangi and Lipothymus sp. Eupristina sp. males showed less aggression. Eupristina sp. has wingless males that mate only within the natal patch, providing excellent examples of full local-mate competition. D. yangi males showed high levels of aggression and lethal combat. D. yangi has winged males but mate mostly within the natal patch. Only a few matings occur after male dispersal. Its sex ratio was lower than the prediction of partial local mate competition theory. Wingless male Lipothymus sp., which mate partly after dispersal, did not present fatal fight. Therefore, the mating behaviour of D. yangi and Lipothymus sp. did not follow predicted patterns, based on wing morph. The mating pattern of D. yangi and Lipothymus sp. should follow the partial local mate competition theory. Furthermore, there was not a significant correlation between the proportion of males and the proportion of fruit parasitized in both winged D. yangi males and wingless Lipothymus sp. males.

The frequency of recombination between unlike genotypes is central to understanding the generation of genetic diversity in natural populations of malaria. Here we suggest a way of investigating the problem which could complement conventional biochemical approaches to the population genetics of malaria. Sex allocation theory is one of the most successful areas of evolutionary biology. A well-supported prediction is that progressively less female-biased sex ratios are favoured with more outcrossing; equal numbers of males and females being evolutionarily stable in randomly mating outbred populations. We present a simple game theory model to support the idea that outcrossing rates in malaria will be correlated with the sex ratio of gametocytes in the peripheral blood of vertebrate hosts. Blood films from epidemiological surveys and culture-adapted isolates from Madang Province, Papua New Guinea, were used to estimate average gametocyte sex ratio of Plasmodium falciparum in the area. The geometric mean proportion of males in the population was 0.18 (95% confidence limits: 0.15–0.22). From our model, we estimate that, on average, 36% of zygotes are the result of outcrossing. This estimate assumes that most microgametes released following exflagellation are capable of fertilization. If, on average, fewer than about 70% of microgametes are capable of fertilization (as is the case in at least one other species of Plasmodium), the observed sex ratio would be consistent with between zero and 36% of zygotes being the result of outcrossing. These estimates suggest that there is usually a numerically dominant genotype in the gametocyte population in a blood meal, and that a considerable amount of selfing is occurring in P. falciparum populations in the Madang region, even though it is an area of intense year-round transmission.