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Colour Variation in Cichlid Fish Seminars in Cell & Developmental Biology 24 (2013) 516–528 Contents lists available at SciVerse ScienceDirect Seminars in Cell & Developmental Biology j ournal homepage: www.elsevier.com/locate/semcdb Review Colour variation in cichlid fish: Developmental mechanisms, selective pressures and evolutionary consequences a,∗ b,1 Martine E. Maan , Kristina M. Sefc a University of Groningen, Behavioural Biology, PO Box 11103, 9700 CC Groningen, The Netherlands b Institute of Zoology, University of Graz, Universitätsplatz 2, A-8010 Graz, Austria a r a t b i c s t l e i n f o r a c t Article history: Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to complex social Available online 9 May 2013 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 Keywords: variation in this group and the selective pressures responsible for the observed variation. We specifically Cichlidae address the evidence for the hypothesis that divergence in colouration is associated with the evolution Natural selection of reproductive isolation between lineages. While we conclude that cichlid colours are excellent models Pigmentation for understanding the role of animal communication in species divergence, we also identify taxonomic Polymorphism and methodological biases in the current research effort. We suggest that the integration of genomic Sexual selection Speciation 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. © 2013 The Authors. Published by Elsevier Ltd. Open access under CC BY license. Contents 1. Introduction . 517 2. Colour pattern variation in the cichlid fish family. 517 2.1. Variation within populations. 517 2.2. Geographic variation . 517 3. Cellular and molecular basis of cichlid colouration and colour pattern differentiation . 517 3.1. Pigments and structural colours . 517 3.2. The genetics of cichlid colouration . 518 3.2.1. Genetic architecture of colour patterns . 518 3.2.2. Modularity and integration. 518 3.3. Molecular processes promoting phenotypic diversification . 518 3.3.1. Phenotype evolution associated with gene expression regulation . 518 3.3.2. Gene duplications . 520 3.3.3. Alternative splicing . 520 4. Developmental and environmental plasticity . 520 5. Evolution . 521 5.1. Drift and hybridisation . 521 5.2. Natural selection . 521 5.2.1. Linking melanin patterns with ecology and social behaviour . 521 5.2.2. Predation risk . 521 5.2.3. Foraging . 521 ∗ Corresponding author. Tel.: +31 0 50 3632196. E-mail addresses: [email protected] (M.E. Maan), [email protected] (K.M. Sefc). 1 Tel.: +43 0 316 380 5601. 1084-9521 © 2013 The Authors. Published by Elsevier Ltd. Open access under CC BY license. http://dx.doi.org/10.1016/j.semcdb.2013.05.003 M.E. Maan, K.M. Sefc / Seminars in Cell & Developmental Biology 24 (2013) 516–528 517 6. Intersexual selection: colouration-mediated mate choice . 522 7. Sexual selection, colour divergence and speciation . 523 7.1. Intraspecific sexual selection and interspecific isolation . 523 7.2. Ecological correlates of sexually selected colour variation . 523 8. Intrasexual competition: territorial interactions . 524 8.1. Colouration-mediated aggression and speciation . 524 9. Concluding remarks . 525 Acknowledgements . 525 References . 525 1. Introduction patterning to conspicuous differences in both colour and pattern (Fig. 1). This variation can interfere with species delimitations, Cichlid fishes are well-known among aquarists and biologists particularly when sexual isolation by mate choice and postzy- for their enormous colour diversity. The family comprises between gotic.
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