Extraordinarily precise nematode sex ratios: Adaptive responses to vanishingly rare mating options

Sex ratio theory predicts both mean sex ratio and variance under a range of population structures. Here, we compare two genera of phoretic nematodes (Parasitodiplogaster and Ficophagus spp.) associated with twelve fig-pollinating wasp species in Panama. The host wasps exhibit classic Local Mate Competition: only inseminated females disperse from natal figs, and their offspring form mating pools that consist of scores of the adult offspring contributed by one or a few foundress mothers. In contrast, in both nematode genera, only sexually undifferentiated juveniles disperse, and their mating pools routinely consist of eight or fewer adults. Across all mating pool sizes, the sex ratios observed in both nematode genera are consistently female-biased (~0.34 males), which is markedly less female-biased than is often observed in the host wasps (~0.10 males). In further contrast with their hosts, variances in nematode sex ratios are also consistently precise (significantly less than binomial). The constraints associated with predictably small mating pools within highly subdivided populations appear to select for precise sex ratios that contribute both to the reproductive success of individual nematodes, and to the evolutionary persistence of nematode species. We suggest that some form of environmental sex determination underlies these precise sex ratios.

Using Meta-analysis to Test Ecological and Evolutionary Theory

This chapter describes nine case studies that illustrate how meta-analysis has contributed to theoretical developments in basic research in ecology and evolution. The main research topics cover are maintenance of biodiversity (Case 1); sexual selection (mate choice/fighting behavior) (cases 2, 8, 9); sex ratio theory (Case 3); allometric scaling (Case 4); the invasiveness of exotic plants (Case 5); seed size and plant abundance (Case 6); and the role of competition and predation in structuring communities (Case 7). It is hoped that these case studies will resonate with the reader and provide “templates” for ways to conduct comparable tests on analogous controversies in their own fields of research.

Measuring mating competition correctly: available evidence supports operational sex ratio theory

Central to sexual selection theory is the question of when individuals should compete for mates. Theory predicts that the sex ratio of ready-to-mate individuals (operational sex ratio; OSR) affects male and female mating competition. In accordance with this, the strength of mating competition, measured by agonistic behaviors and courtship displays, has been found to co-vary with the OSR in field populations of several species. However, laboratory experiments have often produced results that seemingly contradict OSR theory, especially for courtship behavior. We argue that this may be because experiments typically measure frequencies of competitive behaviors. Frequencies of courtship and agonistic behavior are not only affected by the level of mating competition, but also by the number of potential mates or competitors encountered. In contrast, the propensity to behave competitively at a given encounter represents a behavioral response, and thus directly reflects mating competition. We show in 2 simple models that 1) courtship frequency can be expected to respond differently from courtship propensity to changes in OSR and 2) an increase in frequency of agonistic behaviors could occur even if the propensity is not affected by the OSR. In a meta-analysis of studies on courtship competition, we show that frequency measures produced largely opposite results to propensity measures, as predicted by our model. Moreover, courtship propensity increased when the OSR became more biased toward competitors. This presents strong evidence that the OSR affects competition, in the form of courtship, as predicted by OSR theory.

Male gametocyte fecundity and sex ratio of a malaria parasite, Plasmodium mexicanum

SUMMARYEvolutionary theory predicts that the sex ratio of Plasmodium gametocytes will be determined by the number of gametes produced per male gametocyte (male fecundity), parasite clonal diversity and any factor that reduces male gametes' ability to find and combine with female gametes. Despite the importance of male gametocyte fecundity for sex ratio theory as applied to malaria parasites, few data are available on gamete production by male gametocytes. In this study, exflagellating gametes, a measure of male fecundity, were counted for 866 gametocytes from 26 natural infections of the lizard malaria parasite, Plasmodium mexicanum. The maximum male fecundity observed was 8, but most gametocytes produced 2–3 gametes, a value consistent with the typical sex ratio observed for P. mexicanum. Male gametocytes in infections with higher gametocytaemia had lower fecundity. Male fecundity was not correlated with gametocyte size, but differed among infections, suggesting genetic variation for fecundity. Fecundity and sex ratio were correlated (more female gametocytes with higher fecundity) as predicted by theory. Results agree with evolutionary theory, but also suggest a possible tradeoff between production time and fecundity, which could explain the low fecundity of this species, the variation among infections, and the correlation with gametocytaemia.