Speciation in colour

Martine Maan
University of Groningen Groningen institute for evolutionary life sciences

Geographical isolation & divergent selection (allo/sym)

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