Female mating decisions in the Trinidadian guppy, Poecilia reticulata
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Contrary to traditional belief, the decisions that females make before, during and after mating shape its outcome and ultimately fitness. The aim of this thesis was to examine how females modify and adjust their mating decisions in line with social and environmental variability and how these directly and indirectly affect mating benefits. To address this aim I have formulated four main questions that correspond to chapters 2 to 6 in this thesis. Firstly I asked whether there was evidence for female choice being driven by mating benefits. More precisely, in chapter 2, I reviewed the literature in search of evidence for direct and indirect benefits in female choice among freshwater fish species. Direct mating benefits were defined as an increase in female’s reproductive success (number of offspring). Conversely, increases in offspring reproductive success were considered to be indirect benefits. The results showed that despite the multiple suggestions and the great amount of information available, to date there is still no evidence for both direct (increase of F1) or indirect (increase in F2) mating benefits, nor their influence in female mating decisions if freshwater fishes species. Furthermore, although polyandry occurred in more than 60% of the species reviewed, I was unable to confirm that polyandry was maintained because of indirect benefits. These findings justified the need to experimentally investigate the drivers of female mating decisions in freshwater fish species and lead to the questions addressed on chapters 5 and 6. For the experimental chapters 3-6 I used the Trinidadian guppy, Poecilia reticulata, as a model species. Their unique and remarkable ecological and biological characteristics coupled with easy maintenance in laboratory conditions make guppies an ideal species for behavioural studies. But in particular, the fact that guppies live in a promiscuous mating system where females despite being constantly harassed by males may be able to control paternity, makes guppies the ideal species to address my aims. The second question was how much were females in control of their mating decisions, and how social environment could limit these decisions. In particular, in chapter 3 I looked at how females adapt and adjust their mating decisions in line with extreme differences in population sex ratio. In chapter 4, I examined to what extent male sexual harassment affects female reproductive behaviours. Results from these two chapters indicated that female guppies have a remarkable reproductive plasticity that enable them to control their mating decisions. When faced with extreme differences in population sex ratio, female mating decisions were made in an optimal way that maximized the ratio of female reproductive benefits per investment. This translated into producing bigger offspring when in a strong female biased environment, than when in a male biased environment. Further, chapter 4 illustrated that female guppies can, despite high levels of male sexual harassment, be in control of their mating decisions. These two chapters demonstrated and have reinforced previous findings of the remarkable reproductive adaptation of female guppies to differences in the social environment. The third question I addressed was: do multiply mated females have greater direct or indirect benefits than single mated females? To answer this question I followed for the first time reproductive success of females over two generations. I measured fitness directly (number of F1 and F2) and took as well as multiple indirect measures of fitness components for two generations in search for evidence of direct and indirect mating benefits in explaining the maintenance of female multiple mating. The results of chapter 5 revealed that female guppies do not have a higher number of F1 and F2 from polyandry or either from mating with males possessing allegedly good quality traits. I, therefore, stressed the idea that potential differences in sexual selection pressure between laboratory and wild populations may influence the expression and intensity of mating benefits between thus explaining the difficulty of finding mating benefits. In my fourth and last question, I used a novel statistical approach based on the analysis of the dispersion in phenotypes, to look for potential alternative explanations for the prevalence of polyandry. The results of this analysis show offspring from multiple mated mothers were phenotypically more diverse than offspring from single mated mothers. Given the direct relationship between phenotypic diversity and potential fitness gains in stochastic systems, female guppies are likely to get greater benefits from mating with males with different phenotypes than with males with a particular sexual trait. This result provides an alternative explanation for the maintenance of polyandry in resource free systems. Overall the results of this thesis reinforce previous suggestions that female guppies are active participants in the mating process, and not necessarily limited to post-copulatory mechanisms of selection of sperm. It also showed the remarkable ability of females to adjust their reproductive investment in line with changes in the social conditions. Interestingly, my results contradict the commonly accepted assertion that females’ mating preference converges towards unique male sexual traits. This result stresses the need to look at alternative explanations to justify female mating decisions.
Thesis, PhD Doctor of Philosophy
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