Speciation ins sexually reproducing organisms Divergent ecological adaptation -> adaptation assortative mating (<- mate choice) -> reproductive isolation Sexually selected traits often differ between closely related species, not only influence reproductive success but also reproductive isolation between species Taxa with more pronounced sexual ornaments tend to have more species (Darwin) Meta-analysis of comparative evidence: significant but weak positive relationship between estimates of sexual selection and species richness (r=0.07-0.014) Sensory drive: sexual preferences influenced by sensory processes, sexual signals influenced by environmental heterogeneity in signal transmission Indicator mechanisms: sexual preferences: selection for direct or indirect benefits, sexual signals: signal costs Guppy colouration in different populations dependant on predation and dietary compounds (e.g. orange in carotenoid-deficient environment more influential) Ecology-associated variation in sexual communication E.g. bird beak morphology When does it become incipient speciation? Different ecological adaptations may effect mating cues and/or mating preferences Effect on preferences expected to be more powerful exert direct selection on cues while cues exert only indirect selection on preferences Divergent adaptation and assortative mating in Lake Victoria cichlid fish Cichlid: rapid speciation More than 500sp <15ky Colonised different habitats (depth, foraging niche) Diversity in ecologically important traits (foraging morphology, visual systems) Strong sexual selection (female-only parental care) Diversity in male colouration Species pair Pundamilia pundamilia & nyererei Habitat segregation by depth and diet Divergence in mating signals, different male nuptial colouration Divergence in mating preferences, colour-mediated assortative mating Within species sexual selection for conspicuous males Indicator mechanism: heterogeneity in signal value? female choice for locally adapted males Species difference in parasite infection (concept: males signal resistance to parasites through investment into e.g. colours, if different species had different parasites colouration may reflect that). Nyererei (red sp) has more of some that develop in open water more so greater exposure. Pundamilia have more nematodes, nemadotes exposed more as shallow water near edge where nematodes would enter water from bird faeces
Both sp, colourful males have lower parasite loads
Females can judge from male colour the parasite resistance for the correct parasite pressures havent tested heritability of resistance Sensory drive: heterogeneity in signal transmission, female choice for locally conspicuous males? Light frequency changed dependant on depth, red species habitat has not penetrated by blue wavelengths, blue species has more blue, both species trying to generate contrast with green background light, red must use red as the only wavelength available. No green species in lake so none would stand out. As water gets clearer (geographic variation), chroma of colour increases. Male colours diverge from environmental filtering Are there direct effects of divergent adaptation on preference evolution? Sensory drive: heterogeneity in sensory environment Species difference in colour vision, sequence difference in red sensitive pigment gene, red-shift in nyererei, visual pigment expression difference, red expresses more red (at cost of blue/green), behavioural tests: nyererey (red) more sensitive to red light (follow moving red object) Are visual differences adaptive, how do they affect mate choice? Manipulate visual perception -> Test consequences for ecological performance and mate choice Aspects of colour vision can be phenotypically plastic (rear under extreme light e.g. deprived of blue, no more blue cones) Alternative lighting conditions that mimic conditions of lake, short wavelength being there is difference Visual development: qPCR, deep-reared fish tend to express more red sensitive opsin and less blue sensitive opsin (LWS vs SWS) Ecological performance: both species do significantly better in their natural light environment, hybrids inbetween suggests visual environment has survival impact, even in lab, genetics avoid any plastic accommodation. Foraging efficiency (hypothesis) Mate choice: deep reared prefer red, shallow reared prefer blue. No effect of repeating test with/without short wavelength. Female that prefers blue always prefers blue. Conclusions: experimental manipulation of visual system properties seems to work, plasticity does not fully compensate for genetic differences: fish survive less well in wrong light environment (consistent with adaptive explanation for visual system differences, supporting ecology- driven speciation), visual manipulation during development affects mate choice (the idea that divergence in colour vision may pleiotropically cause reproductive isolation), manipulation of colour perception during mate choice has no effect Lack of ever seeing truly blue male before tests may explain choice Multiple mechs acting at once Sensory drive may be powerful because it can directly target both preferences and cues