Unraveling the thread of nature's tapestry: The genetics of diversity and convergence in animal pigmentation

Pigment Cell and Melanoma Research

Volumn 25, Issue 4, 2012, Pages 411-433

  Kronforst, Marcus R ^a   Barsh, Gregory S ^b,c   Kopp, Artyom ^d   Mallet, James ^e,f   Monteiro, Antónia ^g   Mullen, Sean P ^h   Protas, Meredith ⁱ   Rosenblum, Erica B ^j   Schneider, Christopher J ^h   Hoekstra, Hopi E ^e  

^a HARVARD UNIVERSITY   (United States)

^b STANFORD UNIVERSITY SCHOOL OF MEDICINE   (United States)

^c HUDSONALPHA INSTITUTE FOR BIOTECHNOLOGY   (United States)

^d UNIVERSITY OF CALIFORNIA   (United States)

^e HARVARD UNIVERSITY   (United States)

^f UNIVERSITY COLLEGE LONDON   (United Kingdom)

^g YALE UNIVERSITY   (United States)

^h Boston University   (United States)

ⁱ UNIVERSITY OF CALIFORNIA   (United States)

^j UNIVERSITY OF CALIFORNIA   (United States)

more..

Author keywords

Adaptation; Butterfly; Convergence; Drosophila; Evolution; Reptile

Indexed keywords

MELANIN;

BUTTERFLY; CONVERGENT EVOLUTION; DROSOPHILA; FORELIMB; GENETIC VARIABILITY; LIZARD; NONHUMAN; PIGMENTATION; REVIEW;

ANIMALS; BUTTERFLIES; GENETIC VARIATION; MELANINS; NATURE; PIGMENTATION; WING;

EID: 84862661954     PISSN: 17551471     EISSN: 1755148X     Source Type: Journal    
DOI: 10.1111/j.1755-148X.2012.01014.x     Document Type: Review

References (252)

1. Alfoldi, J., Di Palma, F., Grabherr, M. et al. (2011). The genome of the green anole lizard and a comparative analysis with birds and mammals. Nature 477, 587-591.
2. Anderson, T.M., Vonholdt, B.M., Candille, S.I. et al. (2009). Molecular and evolutionary history of melanism in North American gray wolves. Science 323, 1339-1343.
3. Bagnara, J.T., and Hadley, M.E. (1973). Chromatophores and Color Change: The Comparative Physiology of Animal Pigmentation, (Englewood Cliffs, NJ: Prentice-Hall, Inc.).
4. Barrett, R.D.H., and Hoekstra, H.E. (2011). Molecular spandrels: tests of adaptation at the genetic level. Nat. Rev. Genet. 12, 767-780.
5. Barsh, G.S. (1996). The genetics of pigmentation: from fancy genes to complex traits. Trends Genet. 12, 299-305.
6. Bates, H.W. (1862). Contributions to an insect fauna of the Amazon valley. Lepidoptera: Heliconidae. Trans. Linn. Soc. Lond. 23, 495-566.
7. Baxter, S.W., Papa, R., Chamberlain, N., Humphray, S.J., Joron, M., Morrison, C., Ffrench-Constant, R.H., Mcmillan, W.O., and Jiggins, C.D. (2008). Convergent evolution in the genetic basis of Müllerian mimicry in Heliconius butterflies. Genetics 180, 1567-1577.
8. Baxter, S.W., Nadeau, N.J., Maroja, L.S. et al. (2010). Genomic hotspots for adaptation: the population genetics of Müllerian mimicry in the Heliconius melpomene clade. PLoS Genet. 6, e1000794.
9. Beldade, P., Brakefield, P.M., and Long, A.D. (2002). Contribution of Distal-less to quantitative variation in butterfly eyespots. Nature 415, 315-318.
10. Beldade, P., Rudd, S., Gruber, J.D., and Long, A.D. (2006). A wing expressed sequence tag resource for Bicyclus anynana butterflies, an evo-devo model. BMC Genomics 7, 130.
11. Beldade, P., Saenko, S.V., Pul, N., and Long, A.D. (2009). A gene-based linkage map for Bicyclus anynana butterflies allows for a comprehensive analysis of synteny with the lepidopteran reference genome. PLoS Genet. 5, e1000366.
12. Beltran, M., Jiggins, C.D., Brower, A.V.Z., Bermingham, E., and Mallet, J. (2007). Do pollen feeding, pupal-mating and larval gregariousness have a single origin in Heliconius butterflies? Inferences from multilocus DNA sequence data Biol. J. Linn. Soc. 92, 221-239.
13. Bennett, D.C., and Lamoreux, M.L. (2003). The color loci of mice - a genetic century. Pigment Cell Res. 16, 333-344.
14. Benson, W.W. (1972). Natural selection for Müllerian mimicry in Heliconius erato in Costa Rica. Science 176, 936-939.
15. Berner, D., Grandchamp, A.C., and Hendry, A.P. (2009). Variable progress toward ecological speciation in parapatry: stickleback across eight lake-stream transitions. Evolution 63, 1740-1753.
16. Bickel, D.R., Kopp, A., and Nuzhdin, S. (2011). Composite effects of polymorphisms near multiple regulatory elements create a major-effect QTL. PLoS Genet. 7, e1001275.
17. Bock, I.R. (1976). Drosophilidae of Australia I. Drosophila (Insecta: Diptera). Aust. J. Zool. 40, 1-105.
18. Braasch, I., Schartl, M., and Volff, J.N. (2007). Evolution of pigment synthesis pathways by gene and genome duplication in fish. BMC Evol. Biol. 7, 74.
19. Brisson, J.A., De Toni, D.C., Duncan, I., and Templeton, A.R. (2005). Abdominal pigmentation variation in Drosophila polymorpha: geographic variation in the trait, and underlying phylogeography. Evolution 59, 1046-1059.
20. Brisson, J.A., Wilder, J., and Hollocher, H. (2006). Phylogenetic analysis of the Cardini group of Drosophila with respect to changes in pigmentation. Evolution 60, 1228-1241.
21. Brower, A.V.Z. (1994). Phylogeny of Heliconius butterflies inferred from mitochondrial DNA sequences (Lepidoptera: Nymphalidae). Mol. Phylogenet. Evol. 3, 159-174.
22. Brown, K.S. (1981). The biology of Heliconius and related genera. Annu. Rev. Entomol. 26, 427-456.
23. Brown, K.S., Sheppard, P.M., and Turner, J.R.G. (1974). Quaternary refugia in tropical America: evidence from race formation in Heliconius butterflies. Proc. R. Soc. Lond. B 187, 369-378.
24. Brunetti, C.R., Selegue, J.E., Monteiro, A., French, V., Brakefield, P.M., and Carroll, S.B. (2001). The generation and diversification of butterfly eyespot color patterns. Curr. Biol. 11, 1578-1585.
25. Carbone, M.A., Llopart, A., Deangelis, M., Coyne, J.A., and Mackay, T.F. (2005). Quantitative trait loci affecting the difference in pigmentation between Drosophila yakuba and D. santomea. Genetics 171, 211-225.
26. Carroll, S.B., Gates, J., Keys, D.N., Paddock, S.W., Panganiban, G.E.F., Selegue, J.E., and Williams, J.A. (1994). Pattern formation and eyespot determination in butterfly wings. Science 265, 109-114.
27. Castrucci, A.M.D. (1997). Regulation of physiological color change in dorsal skin of male tree lizards, Urosaurus ornatus. Herpetologica 53, 405-410.
28. Chamberlain, N.L., Hill, R.I., Kapan, D.D., Gilbert, L.E., and Kronforst, M.R. (2009). Polymorphic butterfly reveals the missing link in ecological speciation. Science 326, 847-850.
29. Chamberlain, N.L., Hill, R.I., Baxter, S.W., Jiggins, C.D., and Kronforst, M.R. (2011). Comparative population genetics of a mimicry locus among hybridizing Heliconius butterfly species. Heredity 107, 200-204.
30. Chan, R., Stuart-Fox, D., and Jessop, T.S. (2009a). Why are females ornamented? A test of the courtship stimulation and courtship rejection hypotheses Behav. Ecol. 20, 1334-1342.
31. Chan, Y.F., Marks, M.E., Jones, F.C. et al. (2009b). Adaptive evolution of pelvic reduction in sticklebacks by recurrent deletion of a Pitx1 enhancer. Science 327, 302-305.
32. Clark, R., Brown, S.M., Collins, S.C., Jiggins, C.D., Heckel, D.G., and Vogler, A.P. (2008). Colour pattern specification in the Mocker swallowtail Papilio dardanus: the transcription factor invected is a candidate for the mimicry locus H. Proc. R. Soc. B 275, 1181-1188.
33. Clusella Trullas, S., Van Wyk, J.H., and Spotila, J.R. (2007). Thermal melanism in ectotherms. J. Therm. Biol 32, 235-245.
34. Colosimo, P.F., Hosemann, K.E., Balabhadra, S., Villarreal Jr, G., Dickson, M., Grimwood, J., Schmutz, J., Myers, R.M., Schluter, D., and Kingsley, D.M. (2005). Widespread parallel evolution in sticklebacks by repeated fixation of Ectodysplasin alleles. Science 307, 1928-1933.
35. Cooper, W.E., and Burns, N. (1987). Social significance of ventrolateral coloration in the fence lizard, Sceloporus undulatus. Anim. Behav. 35, 526-532.
36. Cott, H.B. (1940). Adaptive Coloration in Animals, (London: Methuen and Co.).
37. Counterman, B.A., Araujo-Perez, F., Hines, H.M. et al. (2010). Genomic hotspots for adaptation: the population genetics of Mullerian mimicry in Heliconius erato. PLoS Genet. 6, e1000796.
38. Cox, R.M., and Calsbeek, R. (2010). Severe costs of reproduction persist in Anolis lizards despite the evolution of a single-egg clutch. Evolution 64, 1321-1330.
39. Cresko, W.A., Amores, A., Wilson, C., Murphy, J., Currey, M., Phillips, P., Bell, M.A., Kimmel, C.B., and Postlethwait, J.H. (2004). Parallel genetic basis for repeated evolution of armor loss in Alaskan threespine stickleback populations. Proc. Natl Acad. Sci. USA 101, 6050-6055.
40. Croucher, P.J.P., Oxford, G.S., Lam, A., and Gillespie, R.G. (2011). Stabilizing selection maintains exuberant colour polymorphism in the spider Theridion californicum (Araneae, Theridiidae). Mol. Ecol. 20, 206-218.
41. Culver, D.C., and Wilkens, H. (2000). Critical review of the relevant theories of the evolution of subterranean animals. In Subterranean Ecosystems, H. Wilkens, D.C. Culver, and W.F. Humphreys, eds. (Amsterdam: Elsevier), pp. 381-398.
42. Davis, M.M., O'keefe, S.L., Primrose, D.A., and Hodgetts, R.B. (2007). A neuropeptide hormone cascade controls the precise onset of post-eclosion cuticular tanning in Drosophila melanogaster. Development 134, 4395-4404.
43. Di Palma, F., Kidd, C., Borowsky, R., and Kocher, T.D. (2007). Construction of bacterial artificial chromosome libraries for the Lake Malawi cichlid (Metriaclima zebra), and the blind cavefish (Astyanax mexicanus). Zebrafish 4, 41-47.
44. Dixon, J.R. (1967). Aspects of the biology of the lizards of the White Sands, New Mexico. Los Angeles County Museum Contribut. Sci. 129, 1-22.
45. Dombeck, I., and Jaenike, J. (2004). Ecological genetics of abdominal pigmentation in Drosophila falleni: a pleiotropic link to nematode parasitism. Evolution 58, 587-596.
46. Dowling, T.E., Martasian, D.P., and Jeffery, W.R. (2002). Evidence for multiple genetic forms with similar eyeless phenotypes in the blind cavefish, Astyanax mexicanus. Mol. Biol. Evol. 19, 446-455.
47. Edmunds, M. (1974). Defense in Animals: A Survey of Anti-predator Defenses, (London: Longman).
48. Edwards, K.A., Doescher, L.T., Kaneshiro, K.Y., and Yamamoto, D. (2007). A database of wing diversity in the Hawaiian Drosophila. PLoS ONE 2, e487.
49. Endler, J.A. (1978). A predator's view of animal color patterns. Evol. Biol. 11, 319-364.
50. Fitze, P.S., Cote, J., San-Jose, L.M., Meylan, S., Isaksson, C., Andersson, S., Rossi, J.M., and Clobert, J. (2009). Carotenoid-based colours reflect the stress response in the common lizard. PLoS ONE 4, e5111.
51. Flanagan, N.S., Tobler, A., Davison, A., Pybus, O.G., Kapan, D.D., Planas, S., Linares, M., Heckel, D., and Mcmillan, W.O. (2004). Historical demography of Müllerian mimicry in the neotropical Heliconius butterflies. Proc. Natl Acad. Sci. USA 101, 9704-9709.
52. Forsman, A. (1995). Heating rates and body temperature variation in melanistic and zigzag Vipera berus: does colour make a difference? Ann. Zool. Fenn. 32, 365-374.
53. Ghiradella, H. (1984). Structure of iridescent Lepidopteran scales - variations on several themes. Ann. Entomol. Soc. Am. 77, 637-645.
54. Ghiradella, H. (1989). Structure and development of iridescent butterfly scales - lattices and laminae. J. Morphol. 202, 69-88.
55. Gibert, P., Moreteau, B., Munjal, A., and David, J.R. (1999). Phenotypic plasticity of abdominal pigmentation in Drosophila kikkawai: multiple interactions between a major gene, sex, abdomen segment and growth temperature. Genetica 105, 165-176.
56. Gilbert, L.E. (2003). Adaptive novelty through introgression in Heliconius wing patterns: evidence for shared genetic "tool box" from synthetic hybrid zones and a theory of diversification. In Ecology and Evolution Taking Flight: Butterflies as Model Systems, C.L. Boggs, W.B. Watt, and P.R. Ehrlich, eds. (Chicago, IL: University of Chicago Press), pp. 281-318.
57. Gilbert, L.E., Forrest, H.S., Schultz, T.D., and Harvey, D.J. (1988). Correlations of ultrastructure and pigmentation suggest how genes control development of wing scales of Heliconius butterflies. J. Res. Lepid. 26, 141-160.
58. Gillespie, R.G., and Oxford, G.S. (1998). Selection on the color polymorphism in Hawaiian happy-face spiders: evidence from genetic structure and temporal fluctuations. Evolution 52, 775-783.
59. Gompel, N., and Prud'homme, B. (2009). The causes of repeated genetic evolution. Dev. Biol. 332, 36-47.
60. Gratten, J., Beraldi, D., Lowder, B.V., Mcrae, A.F., Visscher, P.M., Pemberton, J.M., and Slate, J. (2007). Compelling evidence that a single nucleotide substitution in TYRP1 is responsible for coat-colour polymorphism in a free-living population of Soay sheep. Proc. R. Soc. B 274, 619-626.
61. Gratten, J., Wilson, A.J., Mcrae, A.F., Beraldi, D., Visscher, P.M., Pemberton, J.M., and Slate, J. (2008). A localized negative genetic correlation constrains microevolution of coat color in wild sheep. Science 319, 318-320.
62. Gratten, J., Pilkington, J.G., Brown, E.A., Beraldi, D., Pemberton, J.M., and Slate, J. (2010). The genetic basis of recessive self-colour pattern in a wild sheep population. Heredity 104, 206-214.
63. Greenwood, A.K., Jones, F.C., Chan, Y.F., Brady, S.D., Absher, D.M., Grimwood, J., Schmutz, J., Myers, R.M., Kingsley, D.M., and Peichel, C.L. (2011). The genetic basis of divergent pigment patterns in juvenile threespine sticklebacks. Heredity 107, 155-166.
64. Gross, J.B., Protas, M., Conrad, M., Scheid, P.E., Vidal, O., Jeffery, W.R., Borowsky, R., and Tabin, C.J. (2008). Synteny and candidate gene prediction using an anchored linkage map of Astyanax mexicanus. Proc. Natl Acad. Sci. USA 105, 20106-20111.
65. Gross, J.B., Borowsky, R., and Tabin, C.J. (2009). A novel role for Mc1r in the parallel evolution of depigmentation in independent populations of the cavefish Astyanax mexicanus. PLoS Genet. 5, e1000326.
66. Hager, S.B. (2001). The role of nuptial coloration in female Holbrookia maculata: evidence for a dual signaling system. J. Herpetol. 35, 624-632.
67. Han, Q., Fang, J., Ding, H., Johnson, J.K., Christensen, B.M., and Li, J. (2002). Identification of Drosophila melanogaster yellow-f and yellow-f2 proteins as dopachrome-conversion enzymes. Biochem. J. 368, 333-340.
68. Hawlena, D., Boochnik, R., Abramsky, Z., and Bouskila, A. (2006). Blue tail and striped body: why do lizards change their infant costume when growing up? Behav. Ecol. 17, 889-896.
69. Hines, H.M., Counterman, B.A., Papa, R. et al. (2011). Wing patterning gene redefines the mimetic history of Heliconius butterflies. Proc. Natl Acad. Sci. USA 108, 19666-19671.
70. Hoekstra, H.E. (2006). Genetics, development and evolution of adaptive pigmentation in vertebrates. Heredity 97, 222-234.
71. Hoekstra, H.E., and Coyne, J.A. (2007). The locus of evolution: evo devo and the genetics of adaptation. Evolution 61, 995-1016.
72. Hoekstra, H.E., Krenz, J.G., and Nachman, M.W. (2005). Local adaptation in the rock pocket mouse (Chaetodipus intermedius): natural selection and phylogenetic history of populations. Heredity 94, 217-228.
73. Hoekstra, H.E., Hirschmann, R.J., Bundey, R.A., Insel, P.A., and Crossland, J.P. (2006). A single amino acid mutation contributes to adaptive beach mouse color pattern. Science 313, 101-104.
74. Hofreiter, M., and Schoeneberg, T. (2010). The genetic and evolutionary basis of colour variation in vertebrates. Cell. Mol. Life Sci. 67, 2591-2603.
75. Hohenlohe, P.A., Bassham, S., Etter, P.D., Stiffler, N., Johnson, E.A., and Cresko, W.A. (2010). Population genomics of parallel adaptation in threespine stickleback using sequenced RAD tags. PLoS Genet. 6, e1000862.
76. Hollocher, H., Hatcher, J.L., and Dyreson, E.G. (2000). Evolution of abdominal pigmentation differences across species in the Drosophila dunni subgroup. Evolution 54, 2046-2056.
77. Hubbard, J.K., Uy, J.A.C., Hauber, M.E., Hoekstra, H.E., and Safran, R.J. (2010). Vertebrate pigmentation: from underlying genes to adaptive function. Trends Genet. 26, 231-239.
78. Jacobs, M.E. (1985). Role of beta-alanine in cuticular tanning, sclerotization, and temperature regulation in Drosophila melanogaster. J. Insect Physiol. 31, 509-515.
79. Jeong, S., Rokas, A., and Carroll, S.B. (2006). Regulation of body pigmentation by the Abdominal-B Hox protein and its gain and loss in Drosophila evolution. Cell 125, 1387-1399.
80. Jeong, S., Rebeiz, M., Andolfatto, P., Werner, T., True, J., and Carroll, S.B. (2008). The evolution of gene regulation underlies a morphological difference between two Drosophila sister species. Cell 132, 783-793.
81. Jiggins, C.D., and Mcmillan, W.O. (1997). The genetic basis of an adaptive radiation: warning colour in two Heliconius species. Proc. R. Soc. Lond. B 264, 1167-1175.
82. Jiggins, C.D., Naisbit, R.E., Coe, R.L., and Mallet, J. (2001). Reproductive isolation caused by colour pattern mimicry. Nature 411, 302-305.
83. Jiggins, C.D., Mavarez, J., Beltran, M., Mcmillan, W.O., Johnston, J.S., and Bermingham, E. (2005). A genetic linkage map of the mimetic butterfly Heliconius melpomene. Genetics 171, 557-570.
84. Jones, L., and Lovich, R. (2009). Lizards of the Southwest, (Arizona: Rio Nuevo Publishers).
85. Jones, J.S., Leith, R.H., and Rawlings, P. (1977). Polymorphism in Cepaea: a problem with too many solutions? Ann. Rev. Ecol. Syst. 8, 109-143.
86. Jones, F.C., Grabherr, M.G., Chan, Y.F. et al. (2012). The genomic basis of adaptive evolution in threespine sticklebacks. Nature 484, 55-61.
87. Joron, M., Jiggins, C.D., Papanicolaou, A., and Mcmillan, W.O. (2006a). Heliconius wing patterns: an evo-devo model for understanding phenotypic diversity. Heredity 97, 157-167.
88. Joron, M., Papa, R., Beltran, M. et al. (2006b). A conserved supergene locus controls colour pattern diversity in Heliconius butterflies. PLoS Biol. 4, 1831-1840.
89. Joron, M., Frezal, L., Jones, R.T. et al. (2011). Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry. Nature 477, 203-206.
90. Kalmus, H. (1941). The resistance to desiccation of Drosophila mutants affecting body colour. Proc. R. Soc. Lond. B 130, 185-201.
91. Kapan, D.D. (2001). Three-butterfly system provides a field test of Müllerian mimicry. Nature 409, 338-340.
92. Kapan, D.D., Flanagan, N.S., Tobler, A. et al. (2006). Localization of Müllerian mimicry genes on a dense linkage map of Heliconius erato. Genetics 173, 735-757.
93. Kaufman, D.W. (1974). Adaptive coloration in Peromyscus polionotus: experimental selection by owls. J. Mammal. 55, 271-283.
94. Kawabata, T., Yasuhara, Y., Ochiai, M., Matsuura, S., and Ashida, M. (1995). Molecular cloning of insect pro-phenol oxidase - a copper-containing protein homologous to arthropod hemocyanin. Proc. Natl Acad. Sci. USA 92, 7774-7778.
95. Kelsh, R.N. (2004). Genetics and evolution of pigment patterns in fish. Pigment Cell Res. 17, 326-336.
96. Kettelwell, H.B.D. (1956). A resume of the investigations of the evolution of melanism in the Lepidoptera. Proc. R. Soc. Lond. B 145, 297-303.
97. Keys, D.N., Lewis, D.L., Selegue, J.E., Pearson, B.J., Goodrich, L.V., Johnson, R.L., Gates, J., Scott, M.P., and Carroll, S.B. (1999). Recruitment of a hedgehog regulatory circuit in butterfly eyespot evolution. Science 283, 532-534.
98. King, R.B. (1988). Polymorphic populations of the garter snake Thamnophis sirtalis near Lake Erie. Herpetologica 44, 451-458.
99. King, R.B. (2003). Mendelian inheritance of melanism in the garter snake Thamnophis sirtalis. Herpetologica 59, 484-489.
100. Kingsley, E.P., Manceau, M., Wiley, C.D., and Hoekstra, H.E. (2009). Melanism in Peromyscus Is caused by independent mutations in Agouti. PLoS ONE 4, e6435.
101. Koch, P.B. (1991). Precursors of pattern specific ommatin in red wing scales of the polyphenic butterfly Araschnia levana L.: haemolymph tryptophan and 3-hydroxykynurenine. Insect Biochem. 21, 785-794.
102. Koch, B., Keys, D.N., Rocheleau, T., Aronstein, K., Blackburn, M., Carroll, S.B., and Ffrench-Constant, R.H. (1998). Regulation of dopa decarboxylase expression during colour pattern formation in wild-type and melanic tiger swallowtail butterflies. Development 125, 2303-2313.
103. Kocher, T.D. (2004). Adaptive evolution and explosive speciation: the cichlid fish model. Nat. Rev. Genet. 5, 288-298.
104. Kocurek, G., Carr, M., Ewing, R., Havholm, K.G., Nagar, Y.C., and Singhvi, A.K. (2007). White Sands Dune Field, New Mexico: age, dune dynamics and recent accumulations. Sediment. Geol. 197, 313-331.
105. Kodandaramaiah, U. (2011). The evolutionary significance of butterfly eyespots. Behav. Ecol. 22, 1264-1271.
106. Kondo, S., Iwashita, M., and Yamaguchi, M. (2009). How animals get their skin patterns: fish pigment pattern as a live Turing wave. Int. J. Dev. Biol. 53, 851-856.
107. Kopp, A. (2009). Metamodels and phylogenetic replication: a systematic approach to the evolution of developmental pathways. Evolution 63, 2771-2789.
108. Kopp, A., and Duncan, I. (1997). Control of cell fate and polarity in the adult abdominal segments of Drosophila by optomotor-blind. Development 124, 3715-3726.
109. Kopp, A., and True, J.R. (2002). Evolution of male sexual characters in the oriental Drosophila melanogaster species group. Evol. Devel. 4, 278-291.
110. Kopp, A., Muskavitch, M.A., and Duncan, I. (1997). The roles of hedgehog and engrailed in patterning adult abdominal segments of Drosophila. Development 124, 3703-3714.
111. Kopp, A., Blackman, R.K., and Duncan, I. (1999). Wingless, decapentaplegic and EGF receptor signaling pathways interact to specify dorso-ventral pattern in the adult abdomen of Drosophila. Development 126, 3495-3507.
112. Kopp, A., Duncan, I., Godt, D., and Carroll, S.B. (2000). Genetic control and evolution of sexually dimorphic characters in Drosophila. Nature 408, 553-559.
113. Kopp, A., Graze, R.M., Xu, S., Carroll, S., and Nuzhdin, S.V. (2003). Quantitative trait loci responsible for variation in sexually dimorphic traits in Drosophila melanogaster. Genetics 163, 771-787.
114. Kraminsky, G.P., Clark, W.C., Estelle, M.A., Gietz, R.D., Sage, B.A., O'connor, J.D., and Hodgetts, R.B. (1980). Induction of translatable mRNA for dopa decarboxylase in Drosophila: an early response to ecdysterone. Proc. Natl Acad. Sci. USA 77, 4175-4179.
115. Kronforst, M.R., Kapan, D.D., and Gilbert, L.E. (2006a). Parallel genetic architecture of parallel adaptive radiations in mimetic Heliconius butterflies. Genetics 174, 535-539.
116. Kronforst, M.R., Young, L.G., Kapan, D.D., Mcneely, C., O'neill, R.J., and Gilbert, L.E. (2006b). Linkage of butterfly mate preference and wing color preference cue at the genomic location of wingless. Proc. Natl Acad. Sci. USA 103, 6575-6580.
117. Kuriyama, T., Miyaji, K., Sugimoto, M., and Hasegawa, M. (2006). Ultrastructure of the dermal chromatophores in a lizard (Scincidae: Plestiodon latiscutatus) with conspicuous body and tall coloration. Zoolog. Sci. 23, 793-799.
118. Lindsley, D.L., Grell, E.H., and Bridges, C.B. (1968). Genetic Variations of Drosophila melanogaster, (Washington, DC: Carnegie Inst.).
119. Llopart, A., Elwyn, S., Lachaise, D., and Coyne, J.A. (2002). Genetics of a difference in pigmentation between Drosophila yakuba and Drosophila santomea. Evolution 56, 2262-2277.
120. Losos, J.B. (2009). Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles, (Berkeley, CA: University of California Press).
121. Losos, J.B. (2011). Convergence, adaptation, and constraint. Evolution 65, 1827-1840.
122. Lowe, C.H., and Norris, K.S. (1956). A subspecies of the lizard Sceloporus undulatus from the White Sands of New Mexico. Herpetologica 12, 125-127.
123. Maan, M.E., Seehausen, O., and Van Alphen, J.J.M. (2010). Female mating preferences and male coloration covary with water transparency in a Lake Victoria cichlid fish. Biol. J. Linn. Soc. 99, 398-406.
124. Mallet, J. (1989). The genetics of warning color in peruvian hybrid zones of Heliconius erato and Heliconius melpomene. Proc. R. Soc. Lond. B 236, 163-185.
125. Mallet, J. (2010). Shift happens! Shifting balance and the evolution of diversity in warning colour and mimicry. Ecol. Entomol. 35, 90-104.
126. Mallet, J., and Barton, N.H. (1989). Strong natural selection in a warning color hybrid zone. Evolution 43, 421-431.
127. Mallet, J., and Gilbert, L.E. (1995). Why are there so many mimicry rings? Correlations between habitat, behavior and mimicry in Heliconius butterflies Biol. J. Linn. Soc. 55, 159-180.
128. Manceau, M., Domingues, V.S., Linnen, C.R., Rosenblum, E.B., and Hoekstra, H.E. (2010). Convergence in pigmentation at multiple levels: mutations, genes and function. Phil. Tran. R. Soc. B 365, 2439-2450.
129. Manceau, M., Domingues, V.S., Mallarino, R., and Hoekstra, H.E. (2011). The developmental role of agouti in color pattern evolution. Science 331, 1062-1065.
130. Martinez, M.N., and Cordeiro, A.R. (1970). Modifiers of color pattern genes in Drosophila polymorpha. Genetics 64, 573-584.
131. Mason, C.W. (1926). Structural colors in insects I. J. Phys. Chem. 30, 383-395.
132. Mason, C.W. (1927a). Structural colors in insects II. J. Phys. Chem. 31, 321-354.
133. Mason, C.W. (1927b). Structural colors in insects III. J. Phys. Chem. 31, 1856-1872.
134. McMillan, W.O., Monteiro, A., and Kapan, D.D. (2002). Development and evolution on the wing. Trends Ecol. Evol. 17, 125-133.
135. Merrill, R.M., Van Schooten, B., Scott, J.A., and Jiggins, C.D. (2011). Pervasive genetic associations between traits causing reproductive isolation in Heliconius butterflies. Proc. R. Soc. B 278, 511-518.
136. Michie, L.J., Mallard, F., Majerus, M.E.N., and Jiggins, F.M. (2010). Melanic through nature or nurture: genetic polymorphism and phenotypic plasticity in Harmonia axyridis. J. Evol. Biol. 23, 1699-1707.
137. Michie, L.J., Masson, A., Ware, R.L., and Jiggins, F.M. (2011). Seasonal phenotypic plasticity: wild ladybirds are darker at cold temperatures. Evol. Ecol. 25, 1259-1268.
138. Miller, C.T., Beleza, S., Pollen, A.A., Schluter, D., Kittles, R.A., Shriver, M.D., and Kingsley, D.M. (2007). cis-Regulatory changes in Kit ligand expression and parallel evolution of pigmentation in sticklebacks and humans. Cell 131, 1179-1189.
139. Mills, M.G., Nuckels, R.J., and Parichy, D.M. (2007). Deconstructing evolution of adult phenotypes: genetic analyses of kit reveal homology and evolutionary novelty during adult pigment pattern development of Danio fishes. Development 134, 1081-1090.
140. Monteiro, A. (2008). Alternative models for the evolution of eyespots and of serial homology on lepidopteran wings. BioEssays 30, 358-366.
141. Monteiro, A. (2012). Gene regulatory networks reused to build novel traits. BioEssays 34, 191-196.
142. Monteiro, A., and Podlaha, O. (2009). Wings, horns, and butterfly eyespots: how do complex traits evolve? PLoS Biol. 7, e1000037.
143. Monteiro, A., and Prudic, K.L. (2010). Multiple approaches to study color pattern evolution in butterflies. Trends Evol. Biol. 2, e2.
144. Monteiro, A., Glaser, G., Stockslagger, S., Glansdorp, N., and Ramos, D.M. (2006). Comparative insights into questions of lepidopteran wing pattern homology. BMC Dev. Biol. 6, 52.
145. Morrison, R.L., Sherbrooke, W.C., and Frostmason, S.K. (1996). Temperature-sensitive, physiologically active iridophores in the lizard Urosaurus ornatus: an ultrastructural analysis of color change. Copeia 1996, 804-812.
146. Moussian, B. (2010). Recent advances in understanding mechanisms of insect cuticle differentiation. Insect Biochem. Mol. Biol. 40, 363-375.
147. Müller, F. (1879). Ituna and Thyridia; a remarkable case of mimicry in butterflies. Trans. Entomol. Soc. Lond. 1879, xx-xxix.
148. Mundy, N.I. (2005). A window on the genetics of evolution: MC1R and plumage colouration in birds. Proceedings 272, 1633-1640.
149. Munjal, A.K., Karan, D., Gibert, P., Moreteau, B., Parkash, R., and David, J.R. (1997). Thoracic trident pigmentation in Drosophila melanogaster: latitudinal and altitudinal clines in Indian populations. Genet. Sel. Evol. 29, 601-610.
150. Nadeau, N.J., Minvielle, F., and Mundy, N.I. (2006). Association of a Glu92Lys substitution in MC1R with extended brown in Japanese quail (Coturnix japonica). Anim. Genet. 37, 287-289.
151. Naisbit, R.E., Jiggins, C.D., and Mallet, J. (2003). Mimicry: developmental genes that contribute to speciation. Evol. Dev. 5, 269-280.
152. Ng, C.-S., Hamilton, A., Frank, A., Barmina, O., and Kopp, A. (2008). Genetic basis of sex-specific color patterns in Drosophila malerkotliana. Genetics 180, 421-429.
153. Nicholson, K.E., Harmon, L.J., and Losos, J.B. (2007). Evolution of Anolis lizard dewlap diversity. PLoS ONE 2, e274.
154. Nijhout, H.F. (1986). Pattern and pattern diversity on Lepidopteran wings. Bioscience 36, 527-533.
155. Nijhout, H.F. (1991). The Development and Evolution of Butterfly Wing Patterns, (Washington, DC: Smithsonian Institution Press).
156. Nijhout, H.F., and Koch, P.B. (1991). The distribution of radiolabeled pigment precursors in the wing patterns of nymphalid butterflies. J. Res. Lepid. 30, 1-13.
157. Nosil, P., and Sandoval, C.P. (2008). Ecological niche dimensionality and the evolutionary diversification of stick insects. PLoS ONE 3, e1907.
158. Nosil, P., Crespi, B.J., and Sandoval, C.P. (2002). Host-plant adaptation drives the parallel evolution of reproductive isolation. Nature 417, 440-443.
159. Ohnishi, S., and Watanabe, T.K. (1985). Genetic analysis of color dimorphism in the Drosophila montium subgroup. Jpn. J. Genet. 60, 355-358.
160. Okada, T., and Carson, H.L. (1983). The genera Phorticella Duda and Zaprionus Coquillett (Diptera, Drosophilidae) of the Oriental region and New Guinea. Kontyu, Tokyo 51, 539-553.
161. Oliver, J.C., Robertson, K.A., and Monteiro, A. (2009). Accommodating natural and sexual selection in butterfly wing pattern evolution. Proc. R. Soc. B 276, 2369-2375.
162. Olofsson, M., Vallin, A., Jakobsson, S., and Wiklund, C. (2010). Marginal eyespots on butterfly wings deflect bird attacks under low light intensities with UV wavelengths. PLoS ONE 5, e10798.
163. Orr, H.A. (2010). The population genetics of beneficial mutations. Phil. Tran. R. Soc. B 365, 1195-1201.
164. Oxford, G.S., and Gillespie, R.G. (2001). Portraits of evolution: studies of coloration in Hawaiian spiders. Bioscience 51, 521-528.
165. Papa, R., Martin, A., and Reed, R.D. (2008). Genomic hotspots of adaptation in butterfly wing pattern evolution. Curr. Opin. Genet. Dev. 18, 559-564.
166. Parichy, D.M. (2003). Pigment patterns: fish in stripes and spots. Curr. Biol. 13, R947-R950.
167. Parichy, D.M. (2006). Evolution of danio pigment pattern development. Heredity 97, 200-210.
168. Parichy, D.M. (2007). Homology and the evolution of novelty during Danio adult pigment pattern development. J. Exp. Zool. B Mol. Dev. Evol. 308, 578-590.
169. Parkash, R., Rajpurohit, S., and Ramniwas, S. (2009a). Impact of darker, intermediate and lighter phenotypes of body melanization on desiccation resistance in Drosophila melanogaster. J Insect Sci 9, 1-10.
170. Parkash, R., Singh, S., and Ramniwas, S. (2009b). Seasonal changes in humidity level in the tropics impact body color polymorphism and desiccation resistance in Drosophila jambulina- evidence for melanism-desiccation hypothesis. J. Insect Physiol. 55, 358-368.
171. Pearse, D.E., and Pogson, G.H. (2000). Parallel evolution of the melanic form of the California legless lizard, Anniella pulchra, inferred from mitochondrial DNA sequence variation. Evolution 54, 1041-1046.
172. Pool, J.E., and Aquadro, C.F. (2007). The genetic basis of adaptive pigmentation variation in Drosophila melanogaster. Mol. Ecol. 16, 2844-2851.
173. Protas, M.E., and Patel, N.H. (2008). Evolution of coloration patterns. Annu. Rev. Cell Dev. Biol. 24, 425-446.
174. Protas, M.E., Hersey, C., Kochanek, D., Zhou, Y., Wilkens, H., Jeffery, W.R., Zon, L.I., Borowsky, R., and Tabin, C.J. (2006). Genetic analysis of cavefish reveals molecular convergence in the evolution of albinism. Nat. Genet. 38, 107-111.
175. Prud'homme, B., Gompel, N., Rokas, A., Kassner, V.A., Williams, T.M., Yeh, S.D., True, J.R., and Carroll, S.B. (2006). Repeated morphological evolution through cis-regulatory changes in a pleiotropic gene. Nature 440, 1050-1053.
176. Prudic, K.L., Jeon, C., Cao, H., and Monteiro, A. (2011). Developmental plasticity in sexual roles of butterfly species drives mutual sexual ornamentation. Science 331, 73-75.
177. Quek, S.P., Counterman, B.A., Albuquerque De Moura, P., Cardoso, M.Z., Marshall, C.R., Mcmillan, W.O., and Kronforst, M.R. (2010). Dissecting comimetic radiations in Heliconius reveals divergent histories of convergent butterflies. Proc. Natl Acad. Sci. USA 107, 7365-7370.
178. Rebeiz, M., Pool, J.E., Kassner, V.A., Aquadro, C.F., and Carroll, S.B. (2009a). Stepwise modification of a modular enhancer underlies adaptation in a Drosophila population. Science 326, 1663-1667.
179. Rebeiz, M., Ramos-Womack, M., Jeong, S., Andolfatto, P., Werner, T., True, J., Stern, D.L., and Carroll, S.B. (2009b). Evolution of the tan locus contributed to pigment loss in Drosophila santomea: a response to Matute et al. Cell 139, 1189-1196.
180. Reed, R.D., and Nagy, L. (2005). Evolutionary redeployment of a biosynthetic module: expression of eye pigment gene vermilion, cinnabar, and white in butterfly wing development. Evol. Dev. 7, 301-311.
181. Reed, R.D., and Serfas, M.S. (2004). Butterfly wing pattern evolution is associated with changes in a Notch/Distal-less temporal pattern formation process. Curr. Biol. 14, 1159-1166.
182. Reed, R.D., Papa, R., Martin, A. et al. (2011). optix drives the repeated convergent evolution of butterfly wing pattern mimicry. Science 333, 1137-1141.
183. Reeder, T.W., and Wiens, J.J. (1996). Evolution of the lizard family Phrynosomatidae as inferred from diverse types of data. Herpetol. Monogr. 10, 43-84.
184. Roberts, R.B., Ser, J.R., and Kocher, T.D. (2009). Sexual conflict resolved by invasion of a novel sex determiner in Lake Malawi cichlid fishes. Science 326, 998-1001.
185. Robertson, K.A., and Monteiro, A. (2005). Female Bicyclus anynana butterflies choose males on the basis of their UV-reflective dorsal eyespot pupils. Proc. R. Soc. B 272, 1541-1546.
186. Rosenblum, E.B. (2005). The role of phenotypic plasticity in color variation of Tularosa Basin lizards. Copeia 2005, 586-596.
187. Rosenblum, E.B. (2006). Convergent evolution and divergent selection: lizards at the White Sands ecotone. Am. Nat. 167, 1-15.
188. Rosenblum, E.B., and Harmon, L.J. (2011). "Same same but different": replicated ecological speciation at White Sands. Evolution 65, 946-960.
189. Rosenblum, E.B., Hoekstra, H.E., and Nachman, M.W. (2004). Adaptive reptile color variation and the evolution of the Mc1r gene. Evolution 58, 1794-1808.
190. Rosenblum, E.B., Hickerson, M.J., and Moritz, C. (2007). A multilocus perspective on colonization accompanied by selection and gene flow. Evolution 61, 2971-2985.
191. Rosenblum, E.B., Rompler, H., Schoneberg, T., and Hoekstra, H.E. (2010). Molecular and functional basis of phenotypic convergence in white lizards at White Sands. Proc. Natl Acad. Sci. USA 107, 2113-2117.
192. Sadoglu, P.A. (1957). Mendelian gene for albinism in natural cave fish. Experientia 1957, 394.
193. Sadoglu, P., and Mckee, A. (1969). A second gene that affects eye and body color in Mexican blind cave fish. J. Hered. 60, 10-14.
194. Saenko, S.V., Marialva, M.S., and Beldade, P. (2011). Involvement of the conserved Hox gene Antennapedia in the development and evolution of a novel trait. Evodevo 2, 9.
195. Salzburger, W., Braasch, I., and Meyer, A. (2007). Adaptive sequence evolution in a color gene involved in the formation of the characteristic egg-dummies of male haplochromine cichlid fishes. BMC Biol. 5, 51.
196. Savage, J.M., and Slowinski, J.B. (1992). The coloration of the venomous coral snakes (family Elapidae) and their mimics (families Aniliidae and Colubridae). Biol. J. Linn. Soc. 45, 235-254.
197. Schlotterer, C. (2003). Hitchhiking mapping - functional genomics from the population genetics perspective. Trends Genet. 19, 32-38.
198. Schneider, C.J. (2008). Exploiting genomic resources in studies of speciation and adaptive radiation of lizards in the genus Anolis. Integr. Comp. Biol. 48, 520-526.
199. Seehausen, O., Terai, Y., Magalhaes, I.S. et al. (2008). Speciation through sensory drive in cichlid fish. Nature 455, 620-626.
200. Shapiro, M.D., Marks, M.E., Peichel, C.L., Blackman, B.K., Nereng, K.S., Jonsson, B., Schluter, D., and Kingsley, D.M. (2004). Genetic and developmental basis of evolutionary pelvic reduction in threespine sticklebacks. Nature 428, 717-723.
201. Sheppard, P.M., Turner, J.R.G., Brown, K.S., Benson, W.W., and Singer, M.C. (1985). Genetics and the evolution of Muellerian mimicry in Heliconius butterflies. Phil. Trans. R. Soc. Lond. B 308, 433-610.
202. Sherbrooke, W.C., and Montanucci, R.R. (1988). Stone mimicry in the round-tailed horned lizard, Phrynosoma modestum (Sauria, Iguanidae). J. Arid Environ. 14, 275-284.
203. Shirai, L.T., Saenko, S.V., Keller, R.A., Jeronimo, M.A., Brakefield, P.M., Descimon, H., Wahlberg, N., and Beldade, P. (2012). Evolutionary history of the recruitment of conserved developmental genes in association to the formation and diversification of a novel trait. BMC Evol. Biol. 12, 21.
204. Sinervo, B., and Lively, C.M. (1996). The rock-paper-scissors game and the evolution of alternative male strategies. Nature 380, 240-243.
205. Sinervo, B., and Svensson, E. (2002). Correlational selection and the evolution of genomic architecture. Heredity 89, 329-338.
206. Singh, B.N., and Chatterjee, S. (1987). Greater mating success of Drosophila biarmipes males possessing an apical dark black wing patch. Ethology 75, 81-83.
207. Smith, H.M. (1943). The White Sands earless lizard. Zool. Ser. Field Mus. Nat. His. 24, 339-344.
208. Spicer, G.S. (1991). The genetic basis of a species-specific character in the Drosophila virilis species group. Genetics 128, 331-337.
209. Steiner, C.C., Weber, J.N., and Hoekstra, H.E. (2007). Adaptive variation in beach mice produced by two interacting pigmentation genes. PLoS Biol. 5, 1880-1889.
210. Steiner, C.C., Rompler, H., Boettger, L.M., Schoneberg, T., and Hoekstra, H.E. (2009). The genetic basis of phenotypic convergence in beach mice: similar pigment patterns but different genes. Mol. Biol. Evol. 26, 35-45.
211. Stern, D.L., and Orgogozo, V. (2008). The loci of evolution: how predictable is genetic evolution? Evolution 62, 2155-2177.
212. Stevens, M., Hardman, C.J., and Stubbins, C.L. (2008a). Conspicuousness, not eye mimicry, makes "eyespots" effective antipredator signals. Behav. Ecol. 19, 525-531.
213. Stevens, M., Stubbins, C.L., and Hardman, C.J. (2008b). The anti-predator function of 'eyespots' on camouflaged and conspicuous prey. Behav. Ecol. Sociobiol. 62, 1787-1793.
214. Stinchcombe, J.R., and Hoekstra, H.E. (2008). Combining population genomics and quantitative genetics: finding the genes underlying ecologically important traits. Heredity 100, 158-170.
215. Storz, J.F. (2005). Using genome scans of DNA polymorphism to infer adaptive population divergence. Mol. Ecol. 14, 671-688.
216. Strecker, U., Faundez, V.H., and Wilkens, H. (2004). Phylogeography of surface and cave Astyanax (Teleostei) from Central and North America based on cytochrome b sequence data. Mol. Phylogenet. Evol. 33, 469-481.
217. Streelman, J.T., Albertson, R.C., and Kocher, T.D. (2003). Genome mapping of the orange blotch colour pattern in cichlid fishes. Mol. Ecol. 12, 2465-2471.
218. Stuart-Fox, D., and Moussalli, A. (2009). Camouflage, communication and thermoregulation: lessons from colour changing organisms. Phil. Tran. R. Soc. B 364, 463-470.
219. Sultana, F., Kimura, M.T., and Toda, M.J. (1999). Anthophilic Drosophila of the elegans species-subgroup from Indonesia, with description of a new species (Diptera: Drosophilidae). Entomol. Sci. 2, 121-126.
220. Tan, C.C. (1946). Mosaic dominance in the inheritance of color patterns in the ladybird beetle, Harmonia axyridis. Genetics 31, 195-210.
221. Theron, E., Hawkins, K., Bermingham, E., Ricklefs, R.E., and Mundy, N.I. (2001). The molecular basis of an avian plumage polymorphism in the wild: a melanocortin-1-receptor point mutation is perfectly associated with the melanic plumage morph of the bananaquit, Coereba flaveola. Curr. Biol. 11, 550-557.
222. Tobler, A., Kapan, D., Flanagan, N.S., Gonzalez, C., Peterson, E., Jiggins, C.D., Johntson, J.S., Heckel, D.G., and Mcmillan, W.O. (2005). First-generation linkage map of the warningly colored butterfly Heliconius erato. Heredity 94, 408-417.
223. True, J.R. (2003). Insect melanism: the molecules matter. Trends Ecol. Evol. 18, 640-647.
224. True, J.R., Kopp, A., and Carroll, S.B. (2001). Prophenoloxidase as a reporter of gene expression in Drosophila. Biotechniques 30, 1004-1006, 1008-9.
225. True, J.R., Yeh, S.D., Hovemann, B.T., Kemme, T., Meinertzhagen, I.A., Edwards, T.N., Liou, S.R., Han, Q., and Li, J. (2005). Drosophila tan encodes a novel hydrolase required in pigmentation and vision. PLoS Genet. 1, e63.
226. Turner, J.R.G. (1971). The genetics of some polymorphic forms of the butterflies Heliconius melpomene (Linnaeus) and H. erato (Linneaus), 2: the hybridization of subspecies from Surinam and Trinidad. Zoologica 56, 125-157.
227. Turner, J.R.G. (1976). Adaptive radiation and convergence in subdivisions of butterfly genus Heliconius (Lepidoptera: Nymphalidae). Zool. J. Linn. Soc. 58, 297-308.
228. Turner, J.R.G., and Crane, J. (1962). The genetics of some polymorphic forms of the butterflies Heliconius melpomene Linnaeus and H. erato Linneaus, I: major genes. Zoologica 47, 141-152.
229. Turner, J.R.G., and Mallet, J.L.B. (1996). Did forest islands drive the diversity of warningly coloured butterflies? Biotic drift and the shifting balance Phil. Trans. R. Soc. Lond. B 351, 835-845.
230. Uy, J.A.C., Moyle, R.G., Filardi, C.E., and Cheviron, Z.A. (2009). Difference in plumage color used in species recognition between incipient species is linked to a single amino acid substitution in the melanocortin-1 receptor. Am. Nat. 174, 244-254.
231. Vallin, A., Jakobsson, S., Lind, J., and Wiklund, C. (2005). Prey survival by predator intimidation: an experimental study of peacock butterfly defence against blue tits. Proc. R. Soc. Lond. B 272, 1203-1207.
232. Van't Hof, A.E., Edmonds, N., Dalikova, M., Marec, F., and Saccheri, I.J. (2011). Industrial melanism in British peppered moths has a singular and recent mutational origin. Science 332, 958-960.
233. Vignieri, S.N., Larson, J.G., and Hoekstra, H.E. (2010). The selective advantage of crypsis in mice. Evolution 64, 2153-2158.
234. Vines, T.H., and Schluter, D. (2006). Strong assortative mating between allopatric sticklebacks as a by-product of adaptation to different environments. Proc. R. Soc. B 273, 911-916.
235. Walter, M.F., Zeineh, L.L., Black, B.C., Mcivor, W.E., Wright, T.R., and Biessmann, H. (1996). Catecholamine metabolism and in vitro induction of premature cuticle melanization in wild type and pigmentation mutants of Drosophila melanogaster. Arch. Insect Biochem. Physiol. 31, 219-233.
236. Weiss, S.L., Kennedy, E.A., Safran, R.J., and Mcgraw, K.J. (2011). Pterin-based ornamental coloration predicts yolk antioxidant levels in female striped plateau lizards (Sceloporus virgatus). J. Anim. Ecol. 80, 519-527.
237. Werner, T., Koshikawa, S., Williams, T.M., and Carroll, S.B. (2010). Generation of a novel wing colour pattern by the Wingless morphogen. Nature 464, 1143-1148.
238. Wiens, J.J., Kuczynski, C.A., Arif, S., and Reeder, T.W. (2010). Phylogenetic relationships of phrynosomatid lizards based on nuclear and mitochondrial data, and a revised phylogeny for Sceloporus. Mol. Phylogenet. Evol. 54, 150-161.
239. Wilkens, H. (1988). Evolution and genetics of epigean and cave Astyanax (Characidae, Pisces). Evol. Biol. 23, 271-367.
240. Wilkens, H., and Strecker, U. (2003). Convergent evolution of the cavefish Astyanax (Characidae, Teleostei): genetic evidence from reduced eye-size and pigmentation. Biol. J. Linn. Soc. 80, 545-554.
241. Williams, T.M., Selegue, J.E., Werner, T., Gompel, N., Kopp, A., and Carroll, S.B. (2008). The regulation and evolution of a genetic switch controlling sexually dimorphic traits in Drosophila. Cell 134, 610-623.
242. Wittkopp, P.J., and Beldade, P. (2009). Development and evolution of insect pigmentation: genetic mechanisms and the potential consequences of pleiotropy. Semin. Cell Dev. Biol. 20, 65-71.
243. Wittkopp, P., Vaccaro, K., and Carroll, S. (2002a). Evolution of yellow gene regulation and pigmentation in Drosophila. Curr. Biol. 12, 1547.
244. Wittkopp, P.J., True, J.R., and Carroll, S.B. (2002b). Reciprocal functions of the Drosophila Yellow and Ebony proteins in the development and evolution of pigment patterns. Development 129, 1849-1858.
245. Wittkopp, P.J., Carroll, S.B., and Kopp, A. (2003a). Evolution in black and white: genetic control of pigment patterns in Drosophila. Trends Genet. 19, 495-504.
246. Wittkopp, P.J., Williams, B.L., Selegue, J.E., and Carroll, S.B. (2003b). Drosophila pigmentation evolution: divergent genotypes underlying convergent phenotypes. Proc. Natl Acad. Sci. USA 100, 1808-1813.
247. Wittkopp, P.J., Stewart, E.E., Arnold, L.L., Neidert, A.H., Haerum, B.K., Thompson, E.M., Akhras, S., Smith-Winberry, G., and Shefner, L. (2009). Intraspecific polymorphism to interspecific divergence: genetics of pigmentation in Drosophila. Science 326, 540-544.
248. Wittkopp, P.J., Smith-Winberry, G., Arnold, L.L., Thompson, E.M., Cooley, A.M., Yuan, D.C., Song, Q., and Mcallister, B.F. (2011). Local adaptation for body color in Drosophila americana. Heredity 106, 592-602.
249. Wright, T.R. (1987). The genetics of biogenic amine metabolism, sclerotization, and melanization in Drosophila melanogaster. Adv. Genet. 24, 127-222.
250. Yamamoto, Y., Stock, D.W., and Jeffery, W.R. (2004). Hedgehog signalling controls eye degeneration in blind cavefish. Nature 431, 844-847.
251. Yamamoto, Y., Byerly, M.S., Jackman, W.R., and Jeffery, W.R. (2009). Pleiotropic functions of embryonic sonic hedgehog expression link jaw and taste bud amplification with eye loss during cavefish evolution. Dev. Biol. 330, 200-211.
252. Yassin, A., Da Lage, J.L., David, J.R., Kondo, M., Madi-Ravazzi, L., Prigent, S.R., and Toda, M.J. (2010). Polyphyly of the Zaprionus genus group (Diptera: Drosophilidae). Mol. Phylogenet. Evol. 55, 335-339.