Martine Maan | University of Groningen (original) (raw)

Papers by Martine Maan

Research paper thumbnail of Behavioural isolation may facilitate homoploid hybrid speciation in cichlid fish

Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental speci... more Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental species. In the absence of post-mating isolation due to structural changes in the genome, or temporal or spatial premating isolation, another form of reproductive isolation would be needed for homoploid hybrid speciation to occur. Here, we investigate the potential of behavioural mate choice to generate assortative mating among hybrids and parental species. We made three-first-generation hybrid crosses between different species of African cichlid fish. In three-way mate-choice experiments, we allowed hybrid and nonhybrid females to mate with either hybrid or non-hybrid males. We found that hybrids generally mated nonrandomly and that hybridization can lead to the expression of new combinations of traits and preferences that behaviourally isolate hybrids from both parental species. Specifically, we find that the phenotypic distinctiveness of hybrids predicts the symmetry and extent of their reproductive isolation. Our data suggest that behavioural mate choice among hybrids may facilitate the establishment of isolated hybrid populations, even in proximity to one or both parental species.

Research paper thumbnail of Species-Specific Relationships between Water Transparency and Male Coloration within and between Two Closely Related Lake Victoria Cichlid Species

Environmental variation in signalling conditions affects animal communication traits, with possib... more Environmental variation in signalling conditions affects animal communication traits, with possible consequences for sexual selection and reproductive isolation. Using spectrophotometry, we studied how male coloration within and between populations of two closely related Lake Victoria cichlid species (Pundamilia pundamilia and P. nyererei) covaries with water transparency. Focusing on coloration patches implicated in sexual selection, we predicted that in clear waters, with broad-spectrum light, (1) colours should become more saturated and (2) shift in hue away from the dominant ambient wavelengths, compared to more turbid waters. We found support for these predictions for the red and yellow coloration of P. nyererei but not the blue coloration of P. pundamilia. This may be explained by the species difference in depth distribution, which generates a steeper gradient in visual conditions for P. nyererei compared to P. pundamilia. Alternatively, the importance of male coloration in intraspecific sexual selection may differ between the species. We also found that anal fin spots, that is, the orange spots on male haplochromine anal fins that presumably mimic eggs, covaried with water transparency in a similar way for both species. This is in contrast to the other body regions studied and suggests that, while indeed functioning as signals, these spots may not play a role in species differentiation.

Research paper thumbnail of Sexual selection and speciation: mechanisms in Lake Victoria cichlid fish

The several hundred species of cichlid fish endemic to Lake Victoria (East-Africa) are textbook e... more The several hundred species of cichlid fish endemic to Lake Victoria (East-Africa) are textbook examples of explosive speciation. This thesis evaluates the hypothesis that sexual selection by female mate choice has contributed to this process of divergence, by investigating the mechanisms that drive the evolution and divergence of mating preferences for colour patterns. I studied two representative model systems, the

Research paper thumbnail of Using radar to obtain a quantitative description of summer bird movements in the Dutch coastal area

During 1999, several feasibility studies on the creation of an airfield at an artificial island i... more During 1999, several feasibility studies on the creation of an airfield at an artificial island in the North Sea were carried out. One of the decisive questions was whether such a location would cause an increase in the bird strike probability. In order to obtain reliable three-dimensional bird density distributions, we have carried out monthly radar observations from the piers of IJmuiden protruding 2 km into the North Sea. We used a search and tracking radar and could observe medium-sized birds at up to 7 km distance.

Research paper thumbnail of Sexual selection and speciation: mechanisms in Lake Victoria cichlid fish

M. 2003. Female sticklebacks Gasterosteus aculeatus use self-reference to optimize MHC allele num... more M. 2003. Female sticklebacks Gasterosteus aculeatus use self-reference to optimize MHC allele number during mate selection.

Research paper thumbnail of Sexual conflict over breeding substrate causes female expulsion and offspring loss in a cichlid fish

Females of the Lake Tanganyika cichlid Lamprologus callipterus exclusively breed in empty snail s... more Females of the Lake Tanganyika cichlid Lamprologus callipterus exclusively breed in empty snail shells that males collect in their territories. Male--male competition for shells is severe, leading to frequent shell stealing and territory takeover. As a consequence, males have breeding females in their shells that spawned with competitors. In this field study, we investigated both naturally occurring and experimentally induced

Research paper thumbnail of Female mating preferences and male coloration covary with water transparency in a Lake Victoria cichlid fish

Rapid speciation in Lake Victoria cichlid fish of the genus Pundamilia may be facilitated by sexu... more Rapid speciation in Lake Victoria cichlid fish of the genus Pundamilia may be facilitated by sexual selection: female mate choice exerts sexual selection on male nuptial coloration within species and maintains reproductive isolation between species. However, declining water transparency coincides with increasingly dull coloration and increasing hybridization. In the present study, we investigated the mechanism underlying this pattern in Pundamilia

Research paper thumbnail of Female preference for male color is necessary and sufficient for assortative mating in 2 cichlid sister species

A critical step for speciation in the face of gene flow is the origination of reproductive isolat... more A critical step for speciation in the face of gene flow is the origination of reproductive isolation. The evolution of assortative mating greatly facilitates this process. Assortative mating can be mediated by one or multiple cues across an array of sensory modalities. We here explore possible cues that may underlie female mate choice in a sympatric species pair of cichlid fish from Lake Victoria, Pundamilia pundamilia and Pundamilia nyererei. Previous studies identified species-specific female preferences for male coloration, but effects of other cues could not be ruled out. Therefore, we assessed female choice in a series of experiments in which we manipulated visual (color) and chemical cues. We show that the visibility of differences in nuptial hue (i.e., either blue or red) between males of the 2 species is necessary and sufficient for assortative mating by female mate choice. Such assortment mediated by a single cue may evolve relatively quickly, but could make reproductive isolation vulnerable to environmental changes. These findings confirm the important role of female mate choice for male nuptial hue in promoting the explosive speciation of African haplochromine cichlids.

Research paper thumbnail of Mechanisms of species divergence through visual adaptation and sexual selection: Perspectives from a cichlid model system

The theory of ecological speciation suggests that assortative mating evolves most easily when mat... more The theory of ecological speciation suggests that assortative mating evolves most easily when mating preferences are directly linked to ecological traits that are subject to divergent selection. Sensory adaptation can play a major role in this process, because selective mating is often mediated by sexual signals: bright colours, complex song, pheromone blends and so on. When divergent sensory adaptation affects the perception of such signals, mating patterns may change as an immediate consequence.

Research paper thumbnail of Species-Specific Relationships between Water Transparency and Male Coloration within and between Two Closely Related Lake Victoria Cichlid Species

Environmental variation in signalling conditions affects animal communication traits, with possib... more Environmental variation in signalling conditions affects animal communication traits, with possible consequences for sexual selection and reproductive isolation. Using spectrophotometry, we studied how male coloration within and between populations of two closely related Lake Victoria cichlid species (Pundamilia pundamilia and P. nyererei) covaries with water transparency. Focusing on coloration patches implicated in sexual selection, we predicted that in clear waters, with broad-spectrum light, (1) colours should become more saturated and (2) shift in hue away from the dominant ambient wavelengths, compared to more turbid waters. We found support for these predictions for the red and yellow coloration of P. nyererei but not the blue coloration of P. pundamilia. This may be explained by the species difference in depth distribution, which generates a steeper gradient in visual conditions for P. nyererei compared to P. pundamilia. Alternatively, the importance of male coloration in intraspecific sexual selection may differ between the species. We also found that anal fin spots, that is, the orange spots on male haplochromine anal fins that presumably mimic eggs, covaried with water transparency in a similar way for both species. This is in contrast to the other body regions studied and suggests that, while indeed functioning as signals, these spots may not play a role in species differentiation.

Research paper thumbnail of What do we need to know about speciation?

Trends in Ecology & Evolution, 2012

Research paper thumbnail of Poison Frog Colors Are Honest Signals of Toxicity, Particularly for Bird Predators

The American Naturalist, 2012

Online enhancement: appendix. Dryad data: http://dx.doi.org/10.5061/dryad.t979dp85.

Research paper thumbnail of Sensory Drive in Cichlid Speciation

The American Naturalist, 2006

The role of selection in speciation is a central yet poorly understood problem in evolutionary bi... more The role of selection in speciation is a central yet poorly understood problem in evolutionary biology. The rapid radiations of extremely colorful cichlid fish in African lakes have fueled the hypothesis that sexual selection can drive species divergence without geographical isolation. Here we present experimental evidence for a mechanism by which sexual selection becomes divergent: in two sibling species from Lake Victoria, female mating preferences for red and blue male nuptial coloration coincide with their contextindependent sensitivities to red and blue light, which in turn correspond to a difference in ambient light in the natural habitat of the species. These results suggest that natural selection on visual performance, favoring different visual properties in different spectral environments, may lead to divergent sexual selection on male nuptial coloration. This interplay of ecological and sexual selection along a light gradient may provide a mechanism of rapid speciation through divergent sensory drive.

Research paper thumbnail of Colour variation in cichlid fish: Developmental mechanisms, selective pressures and evolutionary consequences

Seminars in Cell & Developmental Biology, 2013

Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to co... more Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to complex social behaviour and morphological versatility, they are characterised by extensive diversity in colouration, both within and between species. Here, we review the cellular and molecular mechanisms underlying colour variation in this group and the selective pressures responsible for the observed variation. We specifically address the evidence for the hypothesis that divergence in colouration is associated with the evolution of reproductive isolation between lineages. While we conclude that cichlid colours are excellent models for understanding the role of animal communication in species divergence, we also identify taxonomic and methodological biases in the current research effort. We suggest that the integration of genomic approaches with ecological and behavioural studies, across the entire cichlid family and beyond it, will contribute to the utility of the cichlid model system for understanding the evolution of biological diversity.

Research paper thumbnail of Intraspecific sexual selection on a speciation trait, male coloration, in the Lake Victoria cichlid Pundamilia nyererei

Proceedings of the Royal Society B: Biological Sciences, 2004

The haplochromine cichlids of Lake Victoria constitute a classical example of explosive speciatio... more The haplochromine cichlids of Lake Victoria constitute a classical example of explosive speciation. Extensive intra-and interspecific variation in male nuptial coloration and female mating preferences, in the absence of postzygotic isolation between species, has inspired the hypothesis that sexual selection has been a driving force in the origin of this species flock. This hypothesis rests on the premise that the phenotypic traits that underlie behavioural reproductive isolation between sister species diverged under sexual selection within a species. We test this premise in a Lake Victoria cichlid, by using laboratory experiments and field observations. We report that a male colour trait, which has previously been shown to be important for behavioural reproductive isolation between this species and a close relative, is under directional sexual selection by female mate choice within this species. This is consistent with the hypothesis that female choice has driven the divergence in male coloration between the two species. We also find that male territoriality is vital for male reproductive success and that multiple mating by females is common.

Research paper thumbnail of Sexual dimorphism and directional sexual selection on aposematic signals in a poison frog

Proceedings of the National Academy of Sciences, 2009

It is commonly assumed that natural selection imposed by predators is the prevailing force drivin... more It is commonly assumed that natural selection imposed by predators is the prevailing force driving the evolution of aposematic traits. Here, we demonstrate that aposematic signals are shaped by sexual selection as well. We evaluated sexual selection for coloration brightness in populations of the poison frog Oophaga [Dendrobates] pumilio in Panama's Bocas del Toro archipelago. We assessed female preferences for brighter males by manipulating the perceived brightness of spectrally matched males in two-way choice experiments. We found strong female preferences for bright males in two island populations and weaker or ambiguous preferences in females from mainland populations. Spectral reflectance measurements, coupled with an O. pumilio-specific visual processing model, showed that O. pumilio coloration was significantly brighter in island than in mainland morphs. In one of the island populations (Isla Solarte), males were significantly more brightly colored than females. Taken together, these results provide evidence for directional sexual selection on aposematic coloration and document sexual dimorphism in vertebrate warning coloration. Although aposematic signals have long been upheld as exemplars of natural selection, our results show that sexual selection should not be ignored in studies of aposematic evolution.

Research paper thumbnail of Speciation through sensory drive in cichlid fish

Nature, 2008

Theoretically, divergent selection on sensory systems can cause speciation through sensory drive.... more Theoretically, divergent selection on sensory systems can cause speciation through sensory drive. However, empirical evidence is rare and incomplete. Here we demonstrate sensory drive speciation within island populations of cichlid fish. We identify the ecological and molecular basis of divergent evolution in the cichlid visual system, demonstrate associated divergence in male colouration and female preferences, and show subsequent differentiation at neutral loci, indicating reproductive isolation. Evidence is replicated in several pairs of sympatric populations and species. Variation in the slope of the environmental gradients explains variation in the progress towards speciation: speciation occurs on all but the steepest gradients. This is the most complete demonstration so far of speciation through sensory drive without geographical isolation. Our results also provide a mechanistic explanation for the collapse of cichlid fish species diversity during the anthropogenic eutrophication of Lake Victoria.

Research paper thumbnail of Heritability and heterochrony of polychromatism in a Lake Victoria Cichlid fish: stepping stones for speciation?

Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, 2006

In many haplochromine cichlid fish, male nuptial coloration is subject to female mate choice and ... more In many haplochromine cichlid fish, male nuptial coloration is subject to female mate choice and plays a central role in the evolution of reproductive isolation between incipient species. Intraspecific variation in male coloration may serve as a target for diversifying sexual selection and provide a starting point for species divergence. Here, we investigated a polychromatism in Neochromis omnicaeruleus, a haplochromine from Lake Victoria, East-Africa. In this species, male coloration ranges from skyblue to yellow-red and females are grey-blue to yellow. We found that both genetic and environmental factors influence the expression of these colours during individual development. In a natural population, we found that male colour was associated with size and sexual maturity: yellow males were smaller than blue males and tended to be sexually immature. In females, size and maturity did not differ between colour types. Laboratory crosses revealed that there is a heritable component to the observed colour variation: yellow parents produced more yellow offspring than blue parents. Together with repeated aquarium observations of yellow individuals that gradually become blue, these data suggest that yellow males change to blue as they approach sexual maturity, and that the occurrence and timing of this transition is influenced by both environmental and genetic effects. The significance of this mechanism of colour expression as a possible target for divergent selection remains to be evaluated.

Research paper thumbnail of Behavioural isolation may facilitate homoploid hybrid speciation in cichlid fish

Journal of Evolutionary Biology, 2014

Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental speci... more Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental species. In the absence of post-mating isolation due to structural changes in the genome, or temporal or spatial premating isolation, another form of reproductive isolation would be needed for homoploid hybrid speciation to occur. Here, we investigate the potential of behavioural mate choice to generate assortative mating among hybrids and parental species. We made three-first-generation hybrid crosses between different species of African cichlid fish. In three-way mate-choice experiments, we allowed hybrid and nonhybrid females to mate with either hybrid or nonhybrid males. We found that hybrids generally mated nonrandomly and that hybridization can lead to the expression of new combinations of traits and preferences that behaviourally isolate hybrids from both parental species. Specifically, we find that the phenotypic distinctiveness of hybrids predicts the symmetry and extent of their reproductive isolation. Our data suggest that behavioural mate choice among hybrids may facilitate the establishment of isolated hybrid populations, even in proximity to one or both parental species.

Research paper thumbnail of Evidence for selection on coloration in a Panamanian poison frog: a coalescent-based approach

Journal of Biogeography, 2010

Research paper thumbnail of Behavioural isolation may facilitate homoploid hybrid speciation in cichlid fish

Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental speci... more Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental species. In the absence of post-mating isolation due to structural changes in the genome, or temporal or spatial premating isolation, another form of reproductive isolation would be needed for homoploid hybrid speciation to occur. Here, we investigate the potential of behavioural mate choice to generate assortative mating among hybrids and parental species. We made three-first-generation hybrid crosses between different species of African cichlid fish. In three-way mate-choice experiments, we allowed hybrid and nonhybrid females to mate with either hybrid or non-hybrid males. We found that hybrids generally mated nonrandomly and that hybridization can lead to the expression of new combinations of traits and preferences that behaviourally isolate hybrids from both parental species. Specifically, we find that the phenotypic distinctiveness of hybrids predicts the symmetry and extent of their reproductive isolation. Our data suggest that behavioural mate choice among hybrids may facilitate the establishment of isolated hybrid populations, even in proximity to one or both parental species.

Research paper thumbnail of Species-Specific Relationships between Water Transparency and Male Coloration within and between Two Closely Related Lake Victoria Cichlid Species

Environmental variation in signalling conditions affects animal communication traits, with possib... more Environmental variation in signalling conditions affects animal communication traits, with possible consequences for sexual selection and reproductive isolation. Using spectrophotometry, we studied how male coloration within and between populations of two closely related Lake Victoria cichlid species (Pundamilia pundamilia and P. nyererei) covaries with water transparency. Focusing on coloration patches implicated in sexual selection, we predicted that in clear waters, with broad-spectrum light, (1) colours should become more saturated and (2) shift in hue away from the dominant ambient wavelengths, compared to more turbid waters. We found support for these predictions for the red and yellow coloration of P. nyererei but not the blue coloration of P. pundamilia. This may be explained by the species difference in depth distribution, which generates a steeper gradient in visual conditions for P. nyererei compared to P. pundamilia. Alternatively, the importance of male coloration in intraspecific sexual selection may differ between the species. We also found that anal fin spots, that is, the orange spots on male haplochromine anal fins that presumably mimic eggs, covaried with water transparency in a similar way for both species. This is in contrast to the other body regions studied and suggests that, while indeed functioning as signals, these spots may not play a role in species differentiation.

Research paper thumbnail of Sexual selection and speciation: mechanisms in Lake Victoria cichlid fish

The several hundred species of cichlid fish endemic to Lake Victoria (East-Africa) are textbook e... more The several hundred species of cichlid fish endemic to Lake Victoria (East-Africa) are textbook examples of explosive speciation. This thesis evaluates the hypothesis that sexual selection by female mate choice has contributed to this process of divergence, by investigating the mechanisms that drive the evolution and divergence of mating preferences for colour patterns. I studied two representative model systems, the

Research paper thumbnail of Using radar to obtain a quantitative description of summer bird movements in the Dutch coastal area

During 1999, several feasibility studies on the creation of an airfield at an artificial island i... more During 1999, several feasibility studies on the creation of an airfield at an artificial island in the North Sea were carried out. One of the decisive questions was whether such a location would cause an increase in the bird strike probability. In order to obtain reliable three-dimensional bird density distributions, we have carried out monthly radar observations from the piers of IJmuiden protruding 2 km into the North Sea. We used a search and tracking radar and could observe medium-sized birds at up to 7 km distance.

Research paper thumbnail of Sexual selection and speciation: mechanisms in Lake Victoria cichlid fish

M. 2003. Female sticklebacks Gasterosteus aculeatus use self-reference to optimize MHC allele num... more M. 2003. Female sticklebacks Gasterosteus aculeatus use self-reference to optimize MHC allele number during mate selection.

Research paper thumbnail of Sexual conflict over breeding substrate causes female expulsion and offspring loss in a cichlid fish

Females of the Lake Tanganyika cichlid Lamprologus callipterus exclusively breed in empty snail s... more Females of the Lake Tanganyika cichlid Lamprologus callipterus exclusively breed in empty snail shells that males collect in their territories. Male--male competition for shells is severe, leading to frequent shell stealing and territory takeover. As a consequence, males have breeding females in their shells that spawned with competitors. In this field study, we investigated both naturally occurring and experimentally induced

Research paper thumbnail of Female mating preferences and male coloration covary with water transparency in a Lake Victoria cichlid fish

Rapid speciation in Lake Victoria cichlid fish of the genus Pundamilia may be facilitated by sexu... more Rapid speciation in Lake Victoria cichlid fish of the genus Pundamilia may be facilitated by sexual selection: female mate choice exerts sexual selection on male nuptial coloration within species and maintains reproductive isolation between species. However, declining water transparency coincides with increasingly dull coloration and increasing hybridization. In the present study, we investigated the mechanism underlying this pattern in Pundamilia

Research paper thumbnail of Female preference for male color is necessary and sufficient for assortative mating in 2 cichlid sister species

A critical step for speciation in the face of gene flow is the origination of reproductive isolat... more A critical step for speciation in the face of gene flow is the origination of reproductive isolation. The evolution of assortative mating greatly facilitates this process. Assortative mating can be mediated by one or multiple cues across an array of sensory modalities. We here explore possible cues that may underlie female mate choice in a sympatric species pair of cichlid fish from Lake Victoria, Pundamilia pundamilia and Pundamilia nyererei. Previous studies identified species-specific female preferences for male coloration, but effects of other cues could not be ruled out. Therefore, we assessed female choice in a series of experiments in which we manipulated visual (color) and chemical cues. We show that the visibility of differences in nuptial hue (i.e., either blue or red) between males of the 2 species is necessary and sufficient for assortative mating by female mate choice. Such assortment mediated by a single cue may evolve relatively quickly, but could make reproductive isolation vulnerable to environmental changes. These findings confirm the important role of female mate choice for male nuptial hue in promoting the explosive speciation of African haplochromine cichlids.

Research paper thumbnail of Mechanisms of species divergence through visual adaptation and sexual selection: Perspectives from a cichlid model system

The theory of ecological speciation suggests that assortative mating evolves most easily when mat... more The theory of ecological speciation suggests that assortative mating evolves most easily when mating preferences are directly linked to ecological traits that are subject to divergent selection. Sensory adaptation can play a major role in this process, because selective mating is often mediated by sexual signals: bright colours, complex song, pheromone blends and so on. When divergent sensory adaptation affects the perception of such signals, mating patterns may change as an immediate consequence.

Research paper thumbnail of Species-Specific Relationships between Water Transparency and Male Coloration within and between Two Closely Related Lake Victoria Cichlid Species

Environmental variation in signalling conditions affects animal communication traits, with possib... more Environmental variation in signalling conditions affects animal communication traits, with possible consequences for sexual selection and reproductive isolation. Using spectrophotometry, we studied how male coloration within and between populations of two closely related Lake Victoria cichlid species (Pundamilia pundamilia and P. nyererei) covaries with water transparency. Focusing on coloration patches implicated in sexual selection, we predicted that in clear waters, with broad-spectrum light, (1) colours should become more saturated and (2) shift in hue away from the dominant ambient wavelengths, compared to more turbid waters. We found support for these predictions for the red and yellow coloration of P. nyererei but not the blue coloration of P. pundamilia. This may be explained by the species difference in depth distribution, which generates a steeper gradient in visual conditions for P. nyererei compared to P. pundamilia. Alternatively, the importance of male coloration in intraspecific sexual selection may differ between the species. We also found that anal fin spots, that is, the orange spots on male haplochromine anal fins that presumably mimic eggs, covaried with water transparency in a similar way for both species. This is in contrast to the other body regions studied and suggests that, while indeed functioning as signals, these spots may not play a role in species differentiation.

Research paper thumbnail of What do we need to know about speciation?

Trends in Ecology & Evolution, 2012

Research paper thumbnail of Poison Frog Colors Are Honest Signals of Toxicity, Particularly for Bird Predators

The American Naturalist, 2012

Online enhancement: appendix. Dryad data: http://dx.doi.org/10.5061/dryad.t979dp85.

Research paper thumbnail of Sensory Drive in Cichlid Speciation

The American Naturalist, 2006

The role of selection in speciation is a central yet poorly understood problem in evolutionary bi... more The role of selection in speciation is a central yet poorly understood problem in evolutionary biology. The rapid radiations of extremely colorful cichlid fish in African lakes have fueled the hypothesis that sexual selection can drive species divergence without geographical isolation. Here we present experimental evidence for a mechanism by which sexual selection becomes divergent: in two sibling species from Lake Victoria, female mating preferences for red and blue male nuptial coloration coincide with their contextindependent sensitivities to red and blue light, which in turn correspond to a difference in ambient light in the natural habitat of the species. These results suggest that natural selection on visual performance, favoring different visual properties in different spectral environments, may lead to divergent sexual selection on male nuptial coloration. This interplay of ecological and sexual selection along a light gradient may provide a mechanism of rapid speciation through divergent sensory drive.

Research paper thumbnail of Colour variation in cichlid fish: Developmental mechanisms, selective pressures and evolutionary consequences

Seminars in Cell & Developmental Biology, 2013

Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to co... more Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to complex social behaviour and morphological versatility, they are characterised by extensive diversity in colouration, both within and between species. Here, we review the cellular and molecular mechanisms underlying colour variation in this group and the selective pressures responsible for the observed variation. We specifically address the evidence for the hypothesis that divergence in colouration is associated with the evolution of reproductive isolation between lineages. While we conclude that cichlid colours are excellent models for understanding the role of animal communication in species divergence, we also identify taxonomic and methodological biases in the current research effort. We suggest that the integration of genomic approaches with ecological and behavioural studies, across the entire cichlid family and beyond it, will contribute to the utility of the cichlid model system for understanding the evolution of biological diversity.

Research paper thumbnail of Intraspecific sexual selection on a speciation trait, male coloration, in the Lake Victoria cichlid Pundamilia nyererei

Proceedings of the Royal Society B: Biological Sciences, 2004

The haplochromine cichlids of Lake Victoria constitute a classical example of explosive speciatio... more The haplochromine cichlids of Lake Victoria constitute a classical example of explosive speciation. Extensive intra-and interspecific variation in male nuptial coloration and female mating preferences, in the absence of postzygotic isolation between species, has inspired the hypothesis that sexual selection has been a driving force in the origin of this species flock. This hypothesis rests on the premise that the phenotypic traits that underlie behavioural reproductive isolation between sister species diverged under sexual selection within a species. We test this premise in a Lake Victoria cichlid, by using laboratory experiments and field observations. We report that a male colour trait, which has previously been shown to be important for behavioural reproductive isolation between this species and a close relative, is under directional sexual selection by female mate choice within this species. This is consistent with the hypothesis that female choice has driven the divergence in male coloration between the two species. We also find that male territoriality is vital for male reproductive success and that multiple mating by females is common.

Research paper thumbnail of Sexual dimorphism and directional sexual selection on aposematic signals in a poison frog

Proceedings of the National Academy of Sciences, 2009

It is commonly assumed that natural selection imposed by predators is the prevailing force drivin... more It is commonly assumed that natural selection imposed by predators is the prevailing force driving the evolution of aposematic traits. Here, we demonstrate that aposematic signals are shaped by sexual selection as well. We evaluated sexual selection for coloration brightness in populations of the poison frog Oophaga [Dendrobates] pumilio in Panama's Bocas del Toro archipelago. We assessed female preferences for brighter males by manipulating the perceived brightness of spectrally matched males in two-way choice experiments. We found strong female preferences for bright males in two island populations and weaker or ambiguous preferences in females from mainland populations. Spectral reflectance measurements, coupled with an O. pumilio-specific visual processing model, showed that O. pumilio coloration was significantly brighter in island than in mainland morphs. In one of the island populations (Isla Solarte), males were significantly more brightly colored than females. Taken together, these results provide evidence for directional sexual selection on aposematic coloration and document sexual dimorphism in vertebrate warning coloration. Although aposematic signals have long been upheld as exemplars of natural selection, our results show that sexual selection should not be ignored in studies of aposematic evolution.

Research paper thumbnail of Speciation through sensory drive in cichlid fish

Nature, 2008

Theoretically, divergent selection on sensory systems can cause speciation through sensory drive.... more Theoretically, divergent selection on sensory systems can cause speciation through sensory drive. However, empirical evidence is rare and incomplete. Here we demonstrate sensory drive speciation within island populations of cichlid fish. We identify the ecological and molecular basis of divergent evolution in the cichlid visual system, demonstrate associated divergence in male colouration and female preferences, and show subsequent differentiation at neutral loci, indicating reproductive isolation. Evidence is replicated in several pairs of sympatric populations and species. Variation in the slope of the environmental gradients explains variation in the progress towards speciation: speciation occurs on all but the steepest gradients. This is the most complete demonstration so far of speciation through sensory drive without geographical isolation. Our results also provide a mechanistic explanation for the collapse of cichlid fish species diversity during the anthropogenic eutrophication of Lake Victoria.

Research paper thumbnail of Heritability and heterochrony of polychromatism in a Lake Victoria Cichlid fish: stepping stones for speciation?

Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, 2006

In many haplochromine cichlid fish, male nuptial coloration is subject to female mate choice and ... more In many haplochromine cichlid fish, male nuptial coloration is subject to female mate choice and plays a central role in the evolution of reproductive isolation between incipient species. Intraspecific variation in male coloration may serve as a target for diversifying sexual selection and provide a starting point for species divergence. Here, we investigated a polychromatism in Neochromis omnicaeruleus, a haplochromine from Lake Victoria, East-Africa. In this species, male coloration ranges from skyblue to yellow-red and females are grey-blue to yellow. We found that both genetic and environmental factors influence the expression of these colours during individual development. In a natural population, we found that male colour was associated with size and sexual maturity: yellow males were smaller than blue males and tended to be sexually immature. In females, size and maturity did not differ between colour types. Laboratory crosses revealed that there is a heritable component to the observed colour variation: yellow parents produced more yellow offspring than blue parents. Together with repeated aquarium observations of yellow individuals that gradually become blue, these data suggest that yellow males change to blue as they approach sexual maturity, and that the occurrence and timing of this transition is influenced by both environmental and genetic effects. The significance of this mechanism of colour expression as a possible target for divergent selection remains to be evaluated.

Research paper thumbnail of Behavioural isolation may facilitate homoploid hybrid speciation in cichlid fish

Journal of Evolutionary Biology, 2014

Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental speci... more Hybrid speciation is constrained by the homogenizing effects of gene flow from the parental species. In the absence of post-mating isolation due to structural changes in the genome, or temporal or spatial premating isolation, another form of reproductive isolation would be needed for homoploid hybrid speciation to occur. Here, we investigate the potential of behavioural mate choice to generate assortative mating among hybrids and parental species. We made three-first-generation hybrid crosses between different species of African cichlid fish. In three-way mate-choice experiments, we allowed hybrid and nonhybrid females to mate with either hybrid or nonhybrid males. We found that hybrids generally mated nonrandomly and that hybridization can lead to the expression of new combinations of traits and preferences that behaviourally isolate hybrids from both parental species. Specifically, we find that the phenotypic distinctiveness of hybrids predicts the symmetry and extent of their reproductive isolation. Our data suggest that behavioural mate choice among hybrids may facilitate the establishment of isolated hybrid populations, even in proximity to one or both parental species.

Research paper thumbnail of Evidence for selection on coloration in a Panamanian poison frog: a coalescent-based approach

Journal of Biogeography, 2010