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University of Groningen The role of visual adaptation in cichlid fish speciation Wright, Daniel Shane

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University of Groningen

The role of visual adaptation in cichlid fish speciation

Wright, Daniel Shane

IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below.

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Publication date: 2019

Link to publication in University of Groningen/UMCG research database

Citation for published version (APA):

Wright, D. S. (2019). The role of visual adaptation in cichlid fish speciation. University of Groningen.

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The role of visual adaptation

in cichlid fish speciation

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This research in this thesis was carried out in the Evolutionary Genetics, Behaviour, and Development (EGDB) group at the Groningen Institute for Evolutionary Life Sciences (GELIFES), University of Groningen, The Netherlands.

This research was supported by the Swiss National Science Foundation and the Netherlands Foundation for Scientific Research.

Cover design & layout by: Daniel Shane Wright

Printed by: ProefschriftMaken || www.proefschriftmaken.nl ISBN: 978-94-034-1539-0

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The role of visual adaptation in

cichlid fish speciation

PhD thesis

to obtain the degree of PhD at the University of Groningen

on the authority of the Rector Magnificus Prof. E. Sterken

and in accordance with the decision by the College of Deans. This thesis will be defended in public on

Friday 29 March, 2019 at 12.45 hours

by

Daniel Shane Wright

born on 20 November, 1985 in Virginia, USA

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Supervisors

Prof. M.E. Maan Prof. A.G.G. Groothuis

Assessment Committee

Prof. F.J. Weissing Prof. R.C. Fuller Prof. M.E. Ritchie

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T

ABLE OF

C

ONTENTS

CHAPTER 1 ...9

GENERAL INTRODUCTION ...9

Speciation via sexual selection ... 10

Ecological speciation ... 11

Sensory drive ... 11

Lake Victoria cichlids... 12

Pundamilia ... 13

Thesis overview ... 15

CHAPTER 2 ... 17

DEVELOPMENTAL EFFECTS OF VISUAL ENVIRONMENT ON SPECIES-ASSORTATIVE MATING PREFERENCES IN LAKE VICTORIA CICHLID FISH ... 17

Abstract ... 18 Introduction ... 19 Methods ... 20 Behaviour Scoring ... 24 Statistical Analysis ... 24 Results ... 25 Discussion ... 28 Acknowledgements ... 32 Supplementary information ... 33 CHAPTER 3 ... 41

DEVELOPMENTAL EFFECTS OF ENVIRONMENTAL LIGHT ON MALE NUPTIAL COLORATION IN LAKE VICTORIA CICHLID FISH ... 41

Abstract ... 42 Introduction ... 43 Methods ... 45 Statistical analysis ... 47 Results ... 49 Discussion ... 54 Acknowledgements ... 57 Supplementary information ... 58 CHAPTER 4 ... 67

VISUAL PIGMENT EXPRESSION COVARIES WITH VISUAL ENVIRONMENT IN LAKE VICTORIA CICHLID FISH ... 67 Abstract ... 68 Introduction ... 69 Methods ... 71 Statistical analysis ... 75 Results ... 76 Discussion ... 82 Acknowledgements ... 85 Supplementary information ... 86

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CHAPTER 5 ... 93

TESTING SENSORY DRIVE SPECIATION IN CICHLID FISH: LINKING LIGHT CONDITIONS TO OPSIN EXPRESSION, OPSIN GENOTYPE, AND FEMALE MATE PREFERENCE ... 93

Abstract ... 94 Introduction ... 95 Methods ... 97 Statistical analyses ... 99 Results ... 100 Discussion ... 107 Acknowledgements ... 111 Supplementary information ... 112 CHAPTER 6 ... 119

ENVIRONMENTAL LIGHT INFLUENCES FORAGING PERFORMANCE IN LAKE VICTORIA CICHLIDS... 119 Abstract ... 120 Introduction ... 121 Methods ... 123 Statistical Analysis ... 125 Results ... 126 Discussion ... 129 Acknowledgements ... 132 Supplementary information ... 134 CHAPTER 7 ... 139 SYNTHESIS ... 139

Developmental effects of environmental light on female preference ... 141

Developmental effects of the environmental light on male coloration ... 143

Visual pigment expression covaries with light environment in wild fish ... 143

Linking opsin expression, opsin genotype, and mate preference ... 144

Environmental light influences foraging performance ... 146

What is the role of visual adaptation in speciation? ... 148

REFERENCES ... 151

AUTHOR AFFILIATIONS ... 169

SUMMARY ... 171

NEDERLANDSE SAMENVATTING ... 177

ACKNOWLEDGEMENTS ... 183

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“…if further, the eye ever varies and the variations be inherited, as is likewise certainly the case; and if such variations should be useful to any animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, should not be considered as subversive of the theory.”

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