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Here is a list of the most interesting "foraging-antioxidant-immunity-sex signals" papers. Here is a file with all references (carotenoids.ris). You can download the papers individually or, if you want ,at the bottom of the page there are zip files with all(most?) of the papers in pfd format.



Lozano, G. A. 1994. Carotenoids, parasites, and sexual selection. Oikos 70: 309-311 (pdf file).



Perrin, N. 1995. Signalling, mating success and paternal investment in sticklebacks (Gasterosteus aculeatus): a theoretical model. Behaviour 132: 1037-1057 (pdf file).



Bortolotti, G. R., Negro, J. J., Tella, J. L., Marchant, T. A. and Bird, D. M. 1996. Sexual dichromatism in birds independent of diet, parasites and androgens. Proceedings of the Royal Society of London, Series B 263: 1171-1176 (pdf file).

Shykoff, J. A. and Widmer, A. 1996. Parasites and carotenoid based signal intensity: How general should the relationship be? Naturwissenschaften 83: 113-121 (pdf file).

Skarstein, F. and Folstad, I. 1996. Sexual dichromatism and the immunocompetence handicap: An observational approach using Arctic charr. Oikos 76: 359-367 (pdf file).

Tella, J. L., Forero, M .G., Gajon, A., Hiraldo, F. and Donazar, J. A. 1996. Absence of blood-parasitization effects on Lesser Kestrel fitness. Auk 113: 253-256 (pdf file).



Forero, M. G., Tella, J. L. and Gajon, A. 1997. Absence of blood parasites in the Red-Necked Nightjar. Journal of Field Ornithology 68: 575-579 (pdf file).

Götmark, F. and Ahlstrom, M. 1997. Parental preference for red mouth of chicks in a songbird. Proceedings of the Royal Society of London, Series B 264: 959-962 (pdf file).

Houde, A.E. 1997. Sex, color, and mate choice in guppies. Princeton: Princeton University Press.



Kodric-Brown, A. 1998. Sexual dichromatism and temporary color changes in the reproduction of fishes. American Zoologist 38: 70-81 (pdf file).

Lozano, G. A. 1998. Parasitic stress and self-medication in wild animals. In . Mřller, A. P., Milinski, M., and Slater, P. J. B. (Eds.). Advances in the Study of Behavior Volume 27, Stress and Behavior. Chapter 6, pp. 291-317 (pdf file).

Negro, J. J., Bortolotti, G. R., Tella, J. L., Fernie, K. J. and Bird, D. M. 1998. Regulation of integumentary colour and plasma carotenoids in American Kestrels consistent with sexual selection theory. Functional Ecology 12: 307-312 (pdf file).

Svensson, E., Raberg, L., Koch, C. and Hasselquist, D. 1998. Energetic stress, immunosuppression and the costs of an antibody response. Functional Ecology 12: 912-919 (pdf file).

Wedekind, C., Meyer, P., Frischknecht, M., Niggli, U. A., Pfander, H. 1998. Different carotenoids and potential information content of red coloration of male three-spined stickleback. Journal of Chemical Ecology 24: 787- 801 (pdf file).



Candolin, U. 1999. Male-male competition facilitates female choice in sticklebacks. Proceedings of the Royal Society of London, Series B 266: 785-789 (pdf file).

Candolin, U. 1999. The relationship between signal quality and physical condition: is sexual signalling honest in the three-spined stickleback? Animal Behaviour 58: 1261-1267 (pdf file).

Gonzalez, G., Sorci, G., Mřller, A. P., Ninni, P., Hussy, C., and De Lope, F. 1999. Immunocompetence and condition-dependent sexual advertisement in male house sparrows (Passer domesticus). Journal of Animal Ecology 68: 1225-1234 (pdf file).

Grahn, M. 1999. MHC genotype and ornamentation. In: Animal signals: signalling and signal design in animal communication (Espmark, Y., Amundsen, T. & Rosenqvist, G., eds.). The Royal Norwegian Society of Sciences and Letters. Tapir Publishers, Trondheim, Norway (book chapter - pdf file).

Grether, G. F., Hudon, J., and Millie, D. F. 1999. Carotenoid limitation of sexual coloration along an environmental gradient in guppies. Proceedings of the Royal Society of London, Series B 266: 1317-1322 (pdf file).

Hill, G. E. 1999. Is there an immunological cost to carotenoid-based ornamental coloration? American Naturalist 154: 589-595 (pdf file).

Saino, N., Stradi, R., Ninni, P., Pini, E. and Mřller, A. P. 1999. Carotenoid plasma concentration, immune profile, and plumage ornamentation of male barn swallows (Hirundo rustica). American Naturalist 154: 441-448 (pdf file).

von Schantz, T., Bensch, S., Grahn, M., Hasselquist, D. and Wittzell, H. 1999. Good genes, oxidative stress and condition-dependent sexual signals. Proceedings of the Royal Society of London, Series B 266: 1-12 (pdf file).




Andrersson, S. 2000. Efficacy and content in avian colour signals. In Animal Signals: signallling and signal design in animal communication (Y. Espmark, T. Amudsen and G. Rosenqvist, eds), The Royal Norwegian Society of Sciences and Letters, pp. 47-60. Trondheim, Norway: Tapir Academic Press.

Badyaev, A. and Hill, G. E. 2000. Evolution of sexual dichromatism: contribution of carotenoid- versus melanin-based colorarion. Biol J. Linn Soc. 69: 153-172 (pfd file).

Bortolotti, G. R., Tella, J. L., Forero, M. G., Dawson, R. D. and Negro, J. J. 2000. Genetics, local environment and health as factors influencing plasma carotenoids in wild American kestrels (Falco sparverius). Proceedings of the Royal Society of London, Series B 267: 1433-1438 (pdf file).

Brawner, W. R., Hill, G. E., and Sundermann, C. A. 2000. Effects of coccidial and mycoplasmal infections on carotenoid-based plumage pigmentation in male House Finches. Auk 117: 952-963 (pdf file).

Candolin, U. 2000. Changes in expression and honesty of sexual signalling over the reproductive lifetime of sticklebacks. Proceedings of the Royal Society of London, Series B 267: 2425-2430 (pdf file).

Candolin, U. 2000. Male-male competition ensures honest signaling of male parental ability in the three-spined stickleback (Gasterosteus aculeatus). Behavioral Ecology and Sociobiology 49: 57-61 (pdf file).

Grether, G. F. 2000. Carotenoid limitation and mate preference evolution: A test of the indicator hypothesis in guppies (Poecilia reticulata). Evolution 54: 1712-1724 (pdf file).

Hőrak, P., Vellau, H., Ots, I. and Mřller, A. P. 2000. Growth conditions affect carotenoid-based plumage coloration of great tit nestlings. Naturwissenschaften 87: 460-464 (pdf file).

Mřller, A. P., Biard, C., Blount, J. D., Houston, D. C., Ninni, P., Saino, N., Surai, P. F. 2000. Carotenoid-dependent signals: Indicators of foraging efficiency, immunocompetence or detoxification ability? Avian and Poultry Biology Reviews 11: 137-159 (pdf file).

Negro, J. J. and Garrido-Fernandez, J. 2000. Astaxanthin is the major carotenoid in tissues of white storks (Ciconia ciconia) feeding on introduced crayfish (Procambarus clarkii). Comparative Biochemistry and Physiology B 126: 347-352 (pdf file).

Negro, J. J., Tella, J. L., Blanco, G., Forero, M. G. and Garrido-Fernandez, J. 2000. Diet explains interpopulation variation of plasma carotenoids and skin pigmentation in nestling white storks. Physiological and Biochemical Zoology 73: 97-101 (pdf file).

Poiani, A. Goldsmith, A. R. and Evans, M. R. 2000. Ectoparasites of house sparrows (Passer domesticus): an experimental test of the immunocompetence handicap hypothesis and a new model. Behavioral Ecology and Sociobiology 47: 230-242 (pdf file).

Saino, N., Ninni, P., Incagli, M., Calza, S., Sacchi, R. and Mřller, A. P. 2000. Begging and parental care in relation to offspring need and condition in the barn swallow (Hirundo rustica). American Naturalist 156: 637-649 (pdf file).



Candolin, U. and Voigt, H. R. 2001. No effect of a parasite on reproduction in stickleback males: a laboratory artefact? Parasitology 122: 457-464 (pdf file).

Faivre, B., Préault, M., Théry, M., Secondi, J., Patris, B. and Cézilly, F. 2001. Breeding strategy and morphological characters in an urban population of blackbirds, Turdus merula. Animal Behaviour 61: 969-974 (pdf file).

Grether, G. F., Hudon, J. and Endler, J. A. 2001. Carotenoid scarcity, synthetic pteridine pigments and the evolution of sexual coloration in guppies (Poecilia reticulata). Proceedings of the Royal Society of London, Series B 268: 1245-1253 (pdf file).

Hill, G. E. 2001. A red bird in a brown bag: the function and evolution of colourful plumage in the house finch. Oxford University Press (Book).

Hőrak, P., Ots, I., Vell , H., Spottiswoode, C. and Mřller, A. P. 2001. Carotenoid-based plumage coloration reflects hemoparasite infection and local survival in breeding great tits. Oecologia 126: 166-173 (pdf file).

Lozano, G. A. 2001. Carotenoids, immunity, and sexual selection: comparing apples and oranges? American Naturalist 158: 200-203 (pdf file).

Negro, J. J., Tella, J. L., Hiraldo, F., Bortolotti, G. R., Prieto, P. 2001. Sex-and age-related variation in plasma carotenoids despite a constant diet in the Red-legged Partridge Alectoris rufa. Ardea 89: 275-280 (pdf file).

Pryke, S.R., Lawes, M. J., Andersson, S. 2001. Agonistic carotenoid signalling in male red-collared widowbirds: aggression related to the colour signal of both the territory owner and model intruder. Animal Behaviour 62: 695-704 (pdf file).



Arlettaz, R., Christe, P., Surai, P. F. and Mřller, A. P. 2002. Deliberate rusty staining of plumage in the bearded vulture: does function precede art? Animal Behaviour 64: F1-F3 (pdf file).

Bennett, P. and Owens, I. 2002. Evolutionary Ecology of Birds: Life Histories, Mating systems, and Extinction: Oxford University Press.

Blount, J. D., Surai, P. F., Nager, R. G., Houston, D. C., Mřller, A. P., Trewby, M. L. and Kennedy, M. W. 2002. Carotenoids and egg quality in the lesser black-backed gull Larus fuscus: a supplemental feeding study of maternal effects. Proceedings of the Royal Society of London, Series B 269: 29-36 (pdf file).

Bouslama, Z., Lambrechts, M. M., Ziane, N., Djenidi, R. D. and Chabi, Y. 2002. The effect of nest ectoparasites on parental provisioning in a north-African population of the Blue Tit Parus caeruleus. Ibis 144: E73-E78 (pdf file).

Hőrak, P., Surai, P.F. and Mřller, A.P. 2002. Fat-soluble antioxidants in the eggs of great tits Parus major in relation to breeding habitat and laying sequence. Avian Sci., 2: 123-130 (pdf file).

Martín-Vivaldi, M. and Cabrero, J. 2002. Selección sexual. In Evolución: La base de la biología (M. Soler, ed), pp. 235-259. Espańa: Proyecto Sur (pdf file).

Moore, J. 2002. Parasites and the Behavior of Animals. Oxford University Press (book)

Wiebe, K. L. and Bortolotti, G. R. 2002. Variation in carotenoid-based color in Northern flickers in a hybrid zone. Wilson Bulletin 114: 393-400 (pdf file).

Zuk, M. and Stoehr, A. M. 2002. Immune defense and host life history. American Naturalist 160: S9-S22 (pdf file).



Blount, J. D., Metclafe, N. B., Birkhead, T. R., and Surai, P. F. 2003. Carotenoid modulation of immune function and sexual attractiveness in zebra finches. Science 300: 125-127 (pdf file).

Faivre, B., Préault, M., Salvadori, F., Théry, M., Gaillard, M. and Cézilly, F. 2003. Bill colour and immunocompetence in the European blackbird. Animal Behaviour 65: 1125-1131 (pdf file).

Godin, J. G. J. and McDonough, H. E. 2003. Predator preference for brightly colored males in the guppy: a viability cost for a sexually selected trait. Behavioural Ecology 14: 194-200 (pdf file).

Heindl, M. and Winkler, H. 2003. Female canaries (Serinus canaria) associate more with males that contrast strongly against the background. Ethology 109: 259-271(pdf file).

Hőrak, P. and Saks, L. 2003. Animal allure and health linked by plant pigments. BioEssays 25: 1-2 (pdf file).

Hudon, J., Grether, GF., and Millie, DF. 2003. Marginal differentiation between the sexual and general carotenoid pigmentation of guppies (Poecilia reticulata) and a possible visual explanation. Physiological and Biochemical Zoology 76: 776-790 (pdf file).

Jawor, J. M. and Breitwisch, R. 2003. Melanin ornaments, honesty, and sexual selection. Auk 120: 249-265 (pdf file).

Kurtz, J. 2003. Sex, parasites and resistance - an evolutionary approach. Zoology 106: 327-339 (pdf file).

Mahler, B, Araujo, L. S. and Turabo, P. L. 2003. Dietary and sexual correlated of carotenoid pigment expression in dove plumage Condor 105: 258-267 (pdf file).

Massaro, M., Davis, LS and Darby, JT. 2003. Carotenoid-derived ornaments reflect parental quality in male and female yellow-eyed penguins (Megadyptes antipodes). Behavioral Ecology and Sociobiology 55: 169-175 (pdf file).

McGraw, KJ, Ardia, DR. 2003. Carotenoids, immunocompetence, and the information content of sexual colors: An experimental test. American Naturalist 162: 704-712 (pfd file).

Moreno, J. and Osorno, J. L. 2003. Avian egg colour and sexual selection: does eggshell pigmentation reflect female condition and genetic quality? Ecology Letters 6 803- 806 (pdf file).

Navara, KJ, Hill, GE. 2003. Dietary carotenoid pigments and immune function in a songbird with extensive carotenoid-based plumage coloration. Behavioral Ecology 14: 909-916 (pdf file).

Ohlsson, T., Smith, H. G., Raberg, L. and Hasselquist, D. 2003. Effects of nutrition on sexual ornaments and humoral immune responsiveness in adult male pheasants. Ethology, Ecology and Evolution 15: 31-42 (pdf file).

Pryke, S. R. and Andersson, S. 2003. Carotenoid-based epaulettes reveal male competitive ability: experiments with resident and floater red-shouldered widowbirds. Animal Behaviour 66: 217-224 (pdf file).

Pryke, S. R. and Andersson, S. 2003. Carotenoid-based status signalling in red-shouldered widowbirds (Euplectes axillaris): epaulet size and redness affect captive and territorial competition. Behavioural Ecology and Sociobiology 53: 393-401 (pdf file).

Saino, N., Ambrosini, R., Martinelli, R., Ninni, P. and Mřller, A.P. 2003. Gape coloration reliably reflects immunocompetence of barn swallow (Hirundo rustica) nestlings. Behavioral Ecology 14: 16-22 (pdf file).

Saks, L. McGraw, K. and Hőrak, P. 2003. How feather colour reflects its carotenoid content. Functional Ecology 17: 555-561 (pdf file).

Saks, L., Ots, I. and Hőrak, P. 2003. Carotenoid-based plumage coloration of male greenfinches reflects health and immunocompetence. Oecologia 134 : 301-307 (pdf file).

Torres, R., Velando, A. 2003. A dynamic trait affects continuous pair assessment in the blue-footed booby, Sula nebouxii. Behavioral Ecology and Sociobiology 55: 65-72 (pdf file).

Tschirren, R., Fitze, P. S. and Richner, H. 2003. Proximate mechanisms of variation in the carotenoid-based plumage coloration of nestling great tits (Parus major L.) Journal of Evolutionary Biology 16: 91-100 (pdf file).



Alonso-Alvarez C, Bertrand S, Devevey G, Gaillard M, Prost J, Faivre B, Sorci G. 2004. An experimental test of the dose-dependent effect of carotenoids and immune activation on sexual signals and antioxidant activity. American Naturalist 164 (5): 651-659 (pdf file).

Badyaev AV. 2004. Developmental perspective on the evolution of sexual ornaments. Evolutionary Ecology Research 6: 975-991 (pdf file).

Blount JD. 2004. Carotenoids and life-history evolution in animals. Archives of Biochemistry and Biophysics 430: 10-15 (pdf file).

Bright, A., Waas, JR., King, CM., Cuming, PD. 2004. Bill colour and correlates of male quality in blackbirds: an analysis using canonical ordination. Behavioural Processes 65: 123-132 (pdf file).

Fenoglio S, Cucco M, Fracchia L, Martinotti MG, Malacarne G. 2004. Shield colours of the Moorhen are differently related to bacterial presence and health parameters. Ethology, ecology & Evolution 16: 171-180 (pdf file).

Gaillard M, Juillet C, Cezilly F, Perrot-Minnot MJ. 2004. Carotenoids of two freshwater amphipod species (Gammarus pulex and . roeseli) and their common acanthocephalan parasite Polymorphus minutes. Comparative Biochemistry and Physiology B- Biochemistry & Molecular Biology 139 (1): 129-136 (pdf file).

Grande, J. M., Negro, J. J. and Torres, M. J. 2004. The evolution of bird plumage colouration: a role for feather-degrading bacteria? Ardeola 51: 375-383 (pdf file).

Gregoire, A., Preault, M., Cezilly,F., Wood,M. J., Pradel, R., Faivre B. 2004. Stabilizing natural selection on the early expression of a secondary sexual trait in a passerine bird. Journal of Evolutionary Biology 17 (5): 1152-1156 (pdf file).

Grether, G. F., Kasahara, S., Kolluru, G.R., Cooper, E. L. 2004. Sex-specific effects of carotenoid intake on the immunological response to allografts in guppies (Poecilia reticulata). Proceedings of the Royal Society of London Series B. 271: 45-49 (pdf file).

Grether GF, Kolluru GR, Nersissian K. 2004. Individual colour patches as multicomponent signals. Biological Reviews 79 (3): 583-610 (pdf file).

Hill, G. E., Farmer, K. L. and Beck, M. L. 2004. The effects of mycoplasmosis on carotenoid plumage coloration in male house finches. Journal of Experimental Biology 207: 2095-2099 (pdf file).

Hőrak, P., Saks, L., Karu, U., Ots, I., Surai, P. F. and McGraw, K. J. 2004. How coccidian parasites affect health and appearance of greenfinches. Jourmal of Animal Ecology 73: 935-947 (pdf file)

Hőrak, P., Surai, PF., Ots, I., Moller, AP. 2004. Fat soluble antioxidants in brood-rearing great tits Parus major: relations to health and appearance. Journal of Avian Biology 35: 63-70 (pdf file)

Karino K, Haijima Y. 2004. Algal-diet enhances sexual ornament, growth and reproduction in the guppy . BEHAVIOUR 141: 585-601 (pdf file).

Kurtz, J., Kalbe, M., Aeschlimann, PB., Haberli, MA., Wegner, KM., Reusch, TBH., Milinski, M. 2004. Major histocompatibility complex diversity influences parasite resistance and innate immunity in sticklebacks. Proceedings of the Royal Society of London Series B 271: 197-204 (pdf file).

Lozano, G. A. and Lank, D. B. 2004. Immunocompetence and testosterone-induced condition traits in male ruffs (Philomachus pugnax). Animal Biology 54: 315-329 (pdf file).

McGraw KJ. 2004. Colorful songbirds metabolize carotenoids at the integument. Journal of Avian Biology 35: 471-476 (pdf file).

McGraw, KJ.,  Ardia, DR. 2004. Immunoregulatory activity of different dietary carotenoids in male zebra finches. Chemoecology 14: 25-29 (pfd file)

McGraw KJ, Nogare MC. 2004. Carotenoid pigments and the selectivity of psittacofulvin-based coloration systems in parrots. Comparative Biochemistry and Physiology B (3): 229-233 (pdf file).

McGraw, K. J., Safran, R, J., Evans, M. R., Wakamatsu, K. 2004. European barn swallows use melanin to color their feathers brown. Behavioural Ecology 15: 889-891 (pdf file).

McGraw KJ, Schuetz JG. 2004. The evolution of carotenoid coloration in estrildid finches: a biochemical analysis. Comparative Biochemistry and Physiology B 139: 45-51 (pdf file).

Moreno, J. 2004. Avian reproduction in a Mediterranean context: contribution of ornithological research in Spain. Ardeola 51: 51-70 (pdf file).

Ninni, P., de Lope, F., Saino, N., Haussy, C., Moller, AP. 2004. Antioxidants and condition-dependence of arrival date in a migratory passerine. Oikos 105: 55-64 (pdf file).

Peters A, Denk AG, Delhey K, Kempenaers B. 2004. Carotenoid-based bill colour as an indicator of immunocompetence and sperm performance in male mallards. Journal of Evolutionary Biology 17 (5): 1111-1120 (pdf file).

Peters, A., Delhey, K., Denk, A. G. and Kempenaers, B. 2004. Trade-offs between immune investment and sexual signalling in male mallards. American Naturalist 164: 51-59 (pdf file).

Tella, JL, Figuerola, J., Negro, J. J., Blanco, G., Rodriguez-Estrella, R., Forero, MG., Blazquez, MC., Green, AJ., and Hiraldo, F. 2004. Ecological, morphological and phylogenetic correlates of interspecific variation in plasma carotenoid concentration in birds. Journal of Evolutionary Biology 17: 156-164 (pdf file).


Zip file 1995-2004




Alvarez, F. and Sanchez, C. and Angulo, S. 2005. The frontal shield of the moorhen: sex differences and relationship with body condition. Ethology, Ecology & Evolution 17(2): 135-148 (pdf file).

Cassey, P., Ewen, J. G., Boulton, R. L., Blackburn, T. M., Moller, A. P., Biard, C., Olson, V. and Karadas, F. 2005. Egg carotenoids in passerine birds introduced to New Zealand: relations to ecological factors, integument coloration and phylogeny. Functional Ecology 19 (4): 719-726 (pdf file).

Criscuolo, F., Gonzalez-Barroso, M. D., Bouillaud, F., Ricquier, D., Miroux, B., and Sorci, G. 2005. Mitochondrial uncoupling proteins: new perspectives for evolutionary ecologists. American Naturalist 166 (6): 686-699 (pdf file).

Figuerola, J.  Torres, J. Garrido, J., Green, A., Negro, J. J. 2005. Do carotenoids and spleen size vary with helminth load in greylag geese?  Canadian Journal of Zoology 83: 389-395 (pdf file).

Grether, G. F. , Kolluru, G. R., Rodd, F. H. , de la Cerda, J. and Shimazaki, K. 2005. Carotenoid availability affects the development of a colour-based mate preference and the sensory bias to which it is genetically linked. Proceedings of the Royal Society of London B. 272: 2181-2188 (pdf file).

Hill, G. E. and Farmer, K. L. 2005. Carotenoid-based plumage coloration predicts resistance to a novel parasite in the house finch. Naturwissenschaften 92: 30-34 (pdf file).

Karino, K., Utagawa, T. and Shinjo, S. 2005. Heritability of the algal-foraging ability: an indirect benefit of female mate preference for males' carotenoid-based coloration in the guppy, Poecilia reticulata. Behavioral Ecology and Sociobiology 59 (1): 1-5 (pdf file).

Magurran, A.E. 2005. Evolutionary Ecology: The Trinidadian Guppy. Oxford: Oxford University Press.

McGraw, K. J. 2005. The antioxidant function of many animal pigments: are there consistent health benefits of sexually selected colorants? Animal Behaviour 69:757-764 (pdf file).

McGraw, K. J. and D. R. Ardia. 2005. Sex differences in carotenoid status and immune performance in zebra finches. Evol. Ecol. Res. 7:251-262 (pdf file).

Olson, V. A. and Owens, I. P. F. 2005. Interspecific variation in the use of carotenoid-based coloration in birds: diet, life history and phylogeny. Journal of Evolutionary Biology 18 (6): 1534-1546 (pdf file)

Preault. M., Chastel. O., Cezilly, F. and Faivre, B. 2005. Male bill colour and age are associated with parental abilities and breeding performance in blackbirds. Behavioral Ecology and Sociobiology 58 (5): 497-505 (pdf file)

Rutowski, R. L., Macedonia, J. M., Morehouse, N. and Taylor-Taft, L. 2005. Pterin pigments amplify iridescent ultraviolet signal in males of the orange sulphur butterfly, Colias eurytheme. Proceedings of the Royal Society of London B. 272 (1578): 2329-2335 (pdf file).

Searcy, W. A. and Nowicki, S. 2005. The evolution of Animal communication: reliability and deception in signaling systems. Monographs in Behavior and Ecology, Edited by J. R. Krebs and T. Clutton-Brock. Princeton University Press (book chapter).

Torres, R. and Velando, A. 2005. Male preference for female foot colour in the socially monogamous blue-footed booby, Sula nebouxii. Animal Behaviour 69: 59-65 (pdf file).

Van der Veen, I. T. 2005. Costly carotenoids: a trade-off between predation and infection risk? Journal of Evolutionary Biology 18: 992-999 (pdf file).

Van Oort, H. and Dawson, R. D. 2005. Carotenoid ornamentation of adult male Common Redpolls predicts probability of dying in a salmonellosis outbreak. Functional Ecology 19 (5): 822-827 (pdf file).

 Velando, A., Torres, R. and Espinosa, I. 2005. Male coloration and chick condition in blue-footed booby: a cross-fostering experiment. Behavioral Ecology and Sociobiology 58: 175-180 (pdf file).




Arriero, E. and Fargallo, J. A. 2006. Habitat structure is associated with the expression of carotenoid-based coloration in nestling blue tits Parus caeruleus Naturwissenschaften 93: 173-180 (pdf file).

Basolo, A.L.2006. Genetic linkage and color polymorphism in the southern platyfish (Xiphophorus maculatus): A model system for studies of color pattern evolution. Zebrafish 3 (1): 65-83 (pdf file).

Bertrand, S., Alonso-Alvarez, C., Devevey, G., Faivre, B., Prost, J., and Sorci, G. 2006. Carotenoids modulate the trade-off between egg production and resistance to oxidative stress in zebra finches. Oecologia 147: 576-584 (pdf file).

Bertrand S, Faivre B, Sorci G. 2006. Do carotenoid-based sexual traits signal the availability of non-pigmentary antioxidants? Journal of Experimental Biology  209: 4414-4419 (pdf file).

Blount JD, Matheson SM. 2006. Effects of carotenoid supply on escape flight responses in zebra finches, Taeniopygia guttata. Animal Behaviour 72: 595-601(pdf file).

Costantini, D., Dell'Omo, G. 2006. Effects of T-cell-mediated immune response on avian oxidative stress. Comparative Biochemistry and Physiology, Part A 145: 137-142(pdf file).

Cucco M, Guasco B, Malacarne G, Ottonelli R. 2006. Effects of β-carotene supplementation on chick growth, immune status and behaviour in the grey partridge, Perdix perdix. Behavioural Processes 73: 325-332 (pdf file).

Dawson, R. D. and Bortolotti, G. R. 2006. Carotenoid-dependent coloration of male American kestrels predicts ability to reduce parasitic infections. Naturwissenschaften 93: 597-602 (pdf file).

Dowling DK, Mulder RA. 2006. Plumage and its association with reproductive success in red-capped robins. Annales Zoologici Fennici  43: 311-321(pdf file).

Hadfield JD, Owens IPF. 2006. Strong environmental determination of a carotenoid-based plumage trait is not mediated by carotenoid availability. Journal of Evolutionary Biology 19: 1104-1114 (pdf file).

Heath JA, Frederick PC. 2006. White Ibis integument color during the breeding season. Journal of Field Ornithology  77: 141-150 (pdf file).

Hőrak, P., Saks, L. Karu, U. and Ots, I. 2006. Host resistance and parasite virulence in greenfinch coccidiosis.  Journal of Evolutionary Biology 19: 277-288 (pdf).

Horak P, Zilmer M, Saks L, Ots I, Karu U, Zilmer K. 2006. Antioxidant protection, carotenoids and the costs of immune challenge in greenfinches. Journal of Experimental Biology 209: 4329-4338 (pdf file).

Karadas F, Surai P, Grammenidis E, Sparks NHC, Acamovic T. 2006. Supplementation of the maternal diet with tomato powder and marigold extract: effects on the antioxidant system of the developing quail. British Poultry Science 47: 200-208 (pdf file).

Kilner, R. M. (2006). Function and evolution of color in young birds. Avian Coloration Volume II: function and evolution. G. E. Hill and K. J. McGraw. Cambridge, MA, Harvard University Press. II: 201-232.

Kolluru GR, Grether GF, South SH, Dunlop E, Cardinali A, Liu L, Carapiet A. 2006.  The effects of carotenoid and food availability on resistance to a naturally occurring parasite (Gyrodactylus turnbulli) in guppies (Poecilia reticulata) (Carapiet, Andreh). Biological Journal of the Linnean Society 89: 301-309 (pdf file).

Kolluru G.R, Ruiz NC, Del Cid N, Dunlop E, Grether GF. 2006. The effects of carotenoid and food intake on caudal fin regeneration in male guppies. Journal of Fish Biology 68: 1002-1012 (pdf file).

Kurtz J, Wegner KM, Kalbe M, Reusch TBH, Schaschl H, Hasselquist D, Milinski M. 2006. MHC genes and oxidative stress in sticklebacks: an immuno-ecological approach. Proceedings of the Royal Society B-Biological Sciences 273: 1407-1414 (pdf file).

Maan ME, van der Spoel M, Jimenez PQ, van Alphen JJM, Seehausen O. 2006.  Fitness correlates of male coloration in a Lake Victoria cichlid fish. Behavioral Ecology  17: 691-699 (pdf file).

McGraw KJ. 2006. Dietary carotenoids mediate a trade-off between egg quantity and quality in Japanese quail. Ethology Ecology & Evolution 18: 247-256 (pdf file).

McGraw, K. J. 2006. Mechanisms of melanin based coloration. In: Hill G. E. and McGraw K. J. (eds). Avian Coloration Vol I: Mechanisms and measurements. Harvard University Press.

McGraw, K. J. 2006. Mechanisms of carotenoid based coloration. In: Hill G. E. and McGraw K. J. (eds). Avian Coloration Vol I: Mechanisms and measurements. Harvard University Press.

McGraw KJ, Crino OL, Medina-Jerez W, Nolan PM. 2006. Effect of dietary carotenoid supplementation on food intake and immune function in a songbird with no carotenoid coloration. Ethology 112: 1209-1216 (pdf file).

McGraw KJ, Klasing KC. 2006. Carotenoids, immunity, and integumentary coloration in red junglefowl (Gallus gallus). Auk 123: 1161-1171 (pdf file).

Negro, J.J. , Sarasola, J.H. , Fariñas, F. , Zorrilla, I. 2006. Function and occurrence of facial flushing in birds. Comparative Biochemistry and Physiology - Part A 143: 78-84 (pdf file).

Olson, V.A. 2006. Estimating nutrient intake in comparative studies of animals: An example using dietary carotenoid content in birds. Oikos 112: 620-628 (pdf file).

Owens, I. P. F.  2006. Ecological explanations for interspecific variability in coloration. In: Hill G. E. and McGraw K. J. (eds). Avian Coloration Vol II: Function and evolution. Harvard University Press.

Santos S., Elward, B., Lumeij, J. T. 2006.  Sexual dichromatism in the blue-fronted amazon parrot (Amazona aestiva) revealed by multiple-angle spectrometry. Journal of Avian Medicine and Surgery   20: 8-14 (pdf file).

Tummeleht L, Magi M, Kilgas P, Mand R, Horak P. 2006. Antioxidant protection and plasma carotenoids of incubating great tits (Parus major L.) in relation to health state and breeding conditions. Comparative Biochemistry and Physiology, Part C 144:  166-172 (pdf file).

Vaclav R. 2006. Within-male melanin-based plumage and bill elaboration in male house sparrows. Zoological Science   23: 1073-1078 (pdf file).

Velando A, Beamonte-Barrientos R, Torres R. 2006. Pigment-based skin colour in the blue-footed booby: an honest signal of current condition used by females to adjust reproductive investment. Oecologia 149: 535-542 (pdf file).



Aguilera, E. and Amat, J. A. 2007. Carotenoids, immune response and the expression of sexual ornaments in male greenfinches (Carduelis chloris). Naturwissenschaften 94: 895-902 (pdf file).

Andersson S, Prager M, Johansson EIA. 2007. Carotenoid content and reflectance of yellow and red nuptial plumages in widowbirds (Euplectes spp.). Functional Ecology 21: 272-281 (pdf file).

Boughman JW. 2007. Condition-dependent expression of red colour differs between stickleback species Journal of Evolutionary Biology 20: 1577-1590 (pdf file).

Clotfelter ED, Ardia DR, McGraw KJ. 2007. Red fish, blue fish: trade-offs between pigmentation and immunity in Betta splendens. Behavioural Ecology 18: 1139-1145 (pdf file).

Costantini, D., Coluzza, C., Fanfani, A. and Dell'Omo, G. 2007. Effects of carotenoid supplementation on colour expression, oxidative stress and body mass of rehabilitated captive adult kestrels (Falco tinnunculus). Journal of Comparative Physiology, Part B 177(7):723-731 (pdf file).

Dijkstra PD, Hekman R, Schulz RW, Groothuis TGG. 2007. Social stimulation, nuptial colouration, androgens and immunocompetence in a sexual dimorphic cichlid fish. Behavioral Ecology and Sociobiology 61: 599-609 (pdf file).

Eraud C, Devevey G, Gaillard M, Prost J, Sorci G. and Faivre B. 2007. Environmental stress affects the expression of a carotenoid-based sexual trait in male zebra finches. Journal of Experimental Biology 210(20): 3571-3578 (pdf file).

Griggio, M., Serra, L., Licheri, D., Monti, A. and Pilastro, A. 2007. Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal. Behavioural Ecology and Sociobiology 61: 423-433.(pdf file).

Hartley, I. R. 2007. Sexual dimorphism. Pages 121-141. in B. G. M. Jamieson, editor. Reproductive biology and phylogeny of birds, Part B: Sexual Selection, Behavior, Conservation, Embryology and Genetics. Science Publishers Inc., Enfield.

Horak P, Saks L, Zilmer M, Karu U, Zilmer K. 2007. Do dietary antioxidants alleviate the cost of immune activation? An experiment with greenfinches. American Naturalist 170: 625-635 (pdf file).

Isaksson C, Andersson S. 2007. Carotenoid diet and nestling provisioning in urban and rural great tits Parus major. Journal of Avian Biology 38: 564-572 (pdf file).

Isaksson C, McLaughlin P, Monaghan P, Andersson S, 2007. Carotenoid pigmentation does not reflect total non-enzymatic antioxidant activity in plasma of adult and nestling great tits, Parus major. Functional Ecology 21: 1123-1129 (pdf file).

Isaksson C, Von Post M, Andersson S. 2007. Sexual, seasonal, and environmental variation in plasma carotenoids in great tits, Parus major. Biological Journal of the Linnean Society  92: 521-527 (pdf file).

Jouventin, P., K. J. McGraw, M. Morel & A. Célerier (2007) Dietary carotenoid supplementation affects orange beak but not foot coloration in Gentoo Penguins Pygoscelis papua. Waterbirds, 30, 573-578 (pdf file).

Karino K, Shinjo S. 2007. Relationship between algal-foraging ability and expression of sexually selected traits in male guppies. Zoological Science 24: 571-576 (pdf file).

Karino K, Shinjo S, Sato A. 2007. Algal-searching ability in laboratory experiments reflects orange spot coloration of the male guppy in the wild. Behaviour 144: 101-113 (pdf file).

Karu U, Saks L, Horak P. 2007. Carotenoid coloration in greenfinches is individually consistent irrespective of foraging ability. Physiological and Biochemical Zoology  80 (6): 663-670 (pdf file).

Kurtz J, Kalbe M, Langefors S, Mayer I, Milinski M, Hasselquist D. 2007. An experimental test of the immunocompetence handicap hypothesis in a teleost fish: 11-ketotestosterone suppresses innate immunity in three-spined sticklebacks. American Naturalist 170: 509-519 (pdf file).

Martinez-Padilla J, Mougeot F, Perez-Rodriguez L, Bortolotti, GR. 2007. Nematode parasites reduce carotenoid-based signalling in male red grouse. Biology Letters 3: 161-164 (pdf file).

McGraw KJ. 2007. Dietary mineral content influences the expression of melanin-based ornamental coloration. Behavioural Ecology 18: 137-142 (pdf file).

McGraw KJ, Medina-Jerez W, Adams H. 2007. Carotenoid-based plumage coloration and aggression during molt in male house finches. Behaviour 144: 165-178 (pdf file).

Mougeot, F., Perez-Rodriguez, L., Martinez-Padilla, J., Leckie, F., Redpath, S. M. 2007.  Parasites, testosterone and honest carotenoid-based signalling of health. Functional Ecology 21: 886-898 (pdf file).

Norris, D. R., Marra, P. P., Kyser, T. K., Ratcliffe, L. M. and Montgomerie, R. 2007. Continent-wide variation in feather colour of a migratory songbird in relation to body condition and moulting locality. Biology Letters 3: 16-19 (pdf file).

Perez-Rodriguez, L., Alonso-Alvarez, C., Vinuela, J. 2007. Repeated sampling but not sampling hour affects plasma carotenoid levels. Physiological and Biochemical Zoology  80: 250-254 (pdf file).

Pike, T. W., Blount, J. D., Bjerkeng, B., Lindstrom, J., Metcalfe, N. B. 2007. Carotenoids, oxidative stress and female mating preference for longer lived males. Proceedings of the Royal Society B  274: 1591-1596 (pdf file).

Pike,T. W.,  Blount, J. D., Lindström, J. and Metcalfe, N. B. 2007. Availability of non-carotenoid antioxidants affects the expression of a carotenoid-based sexual ornament. Biology Letters 3(4):  353-356 (pdf file).

Pike, T. W, Blount, J. D., Lindstrom, J., Metcalfe, N. B. 2007. Dietary carotenoid availability influences a male's ability to provide parental care. Behavioural Ecology 18: 1100-1105 (pdf file).

Sandre SL, Tammaru T, Esperk T, Julkunen-Tiitto R, Mappes J. 2007. Carotenoid-based colour polyphenism in a moth species: search for fitness correlates. Entomologia Experimentalis Et Applicata  124: 269-277 (pdf file).

Serra L, Griggio M, Licheri D, Pilastro A. 2007. Moult speed constrains the expression of a carotenoid-based sexual ornament. Journal of Evolutionary Biology 20: 2028-2034 (pdf file).

Smith, H. G., Rĺberg, L., Ohlsson, T., Granbom, M. and Hasselquist, D. 2007. Carotenoid and protein supplementation have differential effects on pheasant ornamentation and immunity. Journal of Evolutionary Biology 20: 310–319.(pdf file).



Baeta, R., B. Faivre, S. Motreuil, M. Gaillard & J. Moreau . 2008. Carotenoid trade-off between parasitic resistance and sexual display: An experimental study in the blackbird (Turdus merula). Proc. R. Soc. B Biol. Sci., 275, 427 (pdf file).

Blount, J.D. and McGraw, K. 2008. Signal functions of carotenoids. In Carotenoids. Volume 4: Natural Functions (G. Britton, S. Liaaen-Jensen and H. Pfander, eds), pp. 213-236. Berlin: Springer.

Buehler DM, Piersma T. 2008. Travelling on a budget: predictions and ecological evidence for bottlenecks in the annual cycle of long-distance migrants. Philosophical Transactions of the Royal Society B-Biological Sciences  363(490): 247-266 (pdf file).

Costantini, D. 2008. Oxidative stress in ecology and evolution: lessons from avian studies. Ecology Letters 11: 1238-1251 (pdf file).

De Neve, L., J. A. Fargallo, Vergara, P., Lemus, J. A. and Luaces, I.. 2008. Effects of maternal carotenoid availability in relation to sex, parasite infection and health status of nestling kestrels (Falco tinnunculus). Journal of Experimental Biology 211(9): 1414-1425 .(pdf file).

Doutrelant C, Gregoire A, Grnac N, Gomez D., Lambrechts MM, Perret, P. 2008. Female coloration indicates female reproductive capacity in blue tits. Journal of Evolutionary Biology 21(1): 226-233 (pdf file).

Gautier, P., M. Barroca, S. Bertrand, C. Eraud, M. Gaillard, M. Hamman, S. Motreuil, G. Sorci & B. Faivre. 2008. The presence of females modulates the expression of a carotenoid-based sexual signal. Behav. Ecol. Sociobiol., 62, 1159 (pdf file).

Isaksson C, Andersson S. 2008. Oxidative stress does not influence carotenoid mobilization and plumage pigmentation. Proceedings of the Royal Society B 275(1632): 309-314 (pdf file).

Juola, F. A., K. McGraw & D. C. Dearborn. 2008. Carotenoids and throat pouch coloration in the great frigatebird (Fregata minor). Comparative Biochemistry and Physiology, Part B 149, 370 (pdf file).

Karu, U., Saks, L., Horak, P. 2008. Carotenoid-based plumage coloration is not affected by vitamin E supplementation in male greenfinches. Ecological Research 23(6): 931-935 (pdf file).

Laaksonen, T., Negro, J. J., Lyytinen, S., Valkama, J., Ots, I., annd Korpimaki, J. 2008. Effects of experimental brood size manipulation and gender on carotenoid levels of Eurasian kestrels Falco tinnunculus. PLoS ONE 3(6) (pdf file).

Maan, M. E., A. M. C. Van Rooijen, J. J. M. Van Alphen & O. Seehausen. 2008. Parasite-mediated sexual selection and species divergence in Lake Victoria cichlid fish. Biol. J. Linn. Soc., 94, 53-60 (pdf file).

Perez, C., Lores, M, Velando, A. 2008. Availability of nonpigmentary antioxidant affects red coloration in gulls. Behavioral Ecology 19(5): 967-973 (pdf file).

Perez-Rodriguez, L., Mougeot, F.Alonso-Alvarez, C., Blas, J., Vińuela, J., Bortolotti, G. R. 2008. Cell-mediated immune activation rapidly decreases plasma carotenoids but does not affect oxidative stress in red-legged partridges (Alectoris rufa). Journal of Experimental Biology 211(13): 2155-2161 (pdf file).

Perez-Rodriguez, L. and Vińuela, J., 2008. Carotenoid-based bill and eye ring coloration as honest signals of condition: an experimental test in the red-legged partridge (Alectoris rufa). Naturwissenschaften 95(9): 821-830 (pdf file).

Pizzolon M, Rasotto MB, Mazzoldi C. 2008. Male lagoon gobies, Knipowitschia panizzae, prefer more ornamented to larger females. Behavioral Ecology and Sociobiology   62: 521-528 (pdf file).

Rowe, M. and K. McGraw (2008). Carotenoids in the seminal fluid of wild birds: interspecific variation in fairy-wrens. Condor 110(4): 694-700 (pdf file).

Zuk, M. 2008. Riddled with Life: Friendly Worms, Ladybug Sex, and the Parasites That Make Us Who We Are: Houghton Mifflin Harcourt.




Alonso-Alvarez, C., Pérez-Rodríguez, L., Garcia, J. T. & Vińuela, J. (2009) Testosterone-mediated trade-offs in the old age: A new approach to the immunocompetence handicap and carotenoid-based sexual signalling. Proceedings of the Royal Society B 276:2093-2101 (pdf file).

Biard, C., Hardy, C., Motreuil, S. & Moreau, J. (2009) Dynamics of PHA-induced immune response and plasma carotenoids in birds: Should we have a closer look? Journal of Experimental Biology 212:1336-1343 (pdf file).

Biard, C., Gil, D., Karadaş, F., Saino, N., Spottiswoode, C. N., Surai, P. F. & Mřller, A. P. 2009. Maternal effects mediated by antioxidants and the evolution of carotenoid-based signals in birds. American Naturalist 174: 696-708 (pdf file).

Bonato, M., Evans, M. R., Hasselquist, D. & Cherry, M. I. (2009) Male coloration reveals different components of immunocompetence in ostriches, Struthio camelus. Animal Behaviour 77:1033-1039 (pdf file).

David, P. and Heeb, P. 2009. Parasites and sexual selection. In Ecology and evolution of parasitism (F. Thomas, J.-F. Guégan and F. Renaud, eds), p. 31.48. Oxford: Oxford University Press.

Galván, I., Díaz, L. & Sanz, J. J. 2009. Relationships between territory quality and carotenoid-based plumage colour, cell-mediated immune response, and body mass in Great Tit Parus major nestlings. Acta Ornithologica 44: 139-150 (pdf file).

Geens, A., Dauwe, T. & Eens, M. 2009. Does anthropogenic metal pollution affect carotenoid colouration, antioxidative capacity and physiological condition of great tits (Parus major)? Comparative Biochemistry and Physiology - C Toxicology and Pharmacology 150: 155-163 (pdf file).

Gomez, D., Richardson, C., Lengagne, T., Plenet, S., Joly, P., Léna, J. P. & Théry, M. (2009) The role of nocturnal vision in mate choice: Females prefer conspicuous males in the European tree frog (Hyla arborea). Proceedings of the Royal Society B 276:2351-2358 (pdf file).

Griggio, M., Serra, L., Licheri, D., Campomori, C. & Pilastro, A. (2009) Moult speed affects structural feather ornaments in the blue tit. Journal of Evolutionary Biology 22:782-792 (pdf file).

Griggio, M., Morosinotto, C., Pilastro, A. 2009. Nestlings' carotenoid feather ornament affects parental allocation strategy and reduces maternal survival. Journal of Evolutionary Biology 22 (10): 2077-2085 (pdf file).

Hill, G. E., Hood, W. R. & Huggins, K. 2009. A multifactorial test of the effects of carotenoid access, food intake and parasite load on the production of ornamental feathers and bill coloration in American goldfinches. Journal of Experimental Biology 212:1225-1233 (pdf file).

Kappes, P.J., Stutchbury, B.J.M. and Woolfenden, B.E. 2009. The relationship between carotenoid-based coloration and pairing, within- and extra-pair mating success in the American redstart. Condor, 111: 684-693 (pdf).

Lindström, J., Pike, T.W., Blount, J.D., Metcalfe, N.B. 2009. Optimization of resource allocation can explain the temporal dynamics and honesty of sexual signals . American Naturalist 174(4): 515-525 (pdf file).

Monaghan, P., Metcalfe, N. B. & Torres, R. (2009) Oxidative stress as a mediator of life history trade-offs: Mechanisms, measurements and interpretation. Ecology Letters 12:75-92 (pdf file).

Morales, J., Velando, A. & Torres, R. (2009) Fecundity compromises attractiveness when pigments are scarce. Behavioral Ecology 20:117-123 (pdf file).

Mougeot, F., Pérez-Rodríguez, L., Sumozas, N. & Terraube, J. 2009. Parasites, condition, immune responsiveness and carotenoid-based ornamentation in male red-legged partridge Alectoris rufa. Journal of Avian Biology 40:67-74 (pdf file).

Pérez-Rodríguez, L. 2009. Carotenoids in evolutionary ecology: Re-evaluating the antioxidant role. BioEssays 31 (10): 1116-1126 (pdf file).

Prager, M., Johansson, E. I. A. & Andersson, S. 2009. Differential ability of carotenoid C4-oxygenation in yellow and red bishop species (Euplectes spp.). Comparative Biochemistry and Physiology - B Biochemistry and Molecular Biology 154: 373-380 (pdf file).

Sadd, B. M. & Schmid-Hempel, P. (2009) Principles of ecological immunology. Evolutionary Applications 2:113-121 (pdf file).

Sánchez, M. I., Hortas, F., Figuerola, J. & Green, A. J. (2009) Sandpipers select red brine shrimps rich in both carotenoids and parasites. Ethology 115:196-200 (pdf file).

Sorci, G. & Faivre, B. 2009. Inflammation and oxidative stress in vertebrate host-parasite systems. Philosophical Transactions of the Royal Society, Series B 364:71-83 (pdf file).

Stirnemann, I., Johnston, G., Rich, B., Robertson, J. & Kleindorfer, S. 2009. Phytohaemagglutinin (PHA) response and bill-hue wavelength increase with carotenoid supplementation in Diamond Firetails (Stagonopleura guttata). Emu 109: 344-351 (pdf file).

Svensson, P. A., Pélabon, C., Blount, J. D., Forsgren, E., Bjerkeng, B. & Amundsen, T. 2009. Temporal variability in a multicomponent trait: Nuptial coloration of female two-spotted gobies. Behavioral Ecology 20:346-353 (pdf file).

Tanvez, A., Amy, M., Chastel, O. & Leboucher, G. 2009. Maternal effects and β-carotene assimilation in Canary chicks. Physiology and Behavior 96:389-393 (pdf file).

Toomey, M. B. & McGraw, K. J. 2009. Seasonal, sexual, and quality related variation in retinal carotenoid accumulation in the house finch (Carpodacus mexicanus). Functional Ecology 23:321-329 (pdf file).

Van Dongen, W. F. D. & Mulder, R. A. 2009. Multiple ornamentation, female breeding synchrony, and extra-pair mating success of golden whistlers (Pachycephala pectoralis). Journal of Ornithology 150: 607-620 (pdf file).


zip 2005-2009




Ardia, D. R., Broughton, D. R. & Gleicher, M. J. 2010. Short-term exposure to testosterone propionate leads to rapid bill color and dominance changes in zebra finches. Hormones and Behavior 58: 526-532 (pdf file).

Biard, C., Saulnier, N., Gaillard, M. & Moreau, J. 2010. Carotenoid-based bill colour is an integrative signal of multiple parasite infection in blackbird. Naturwissenschaften 97(11): 987-995 (pdf file).

Costantini, D., Rowe, M., Butler, M.W. & McGraw, K.J. 2010. From molecules to living systems: historical and contemporary issues in oxidative stress and antioxidant ecology. Funct. Ecol. 24: 950-959 (pdf file).

Cote, J., Arnoux, E., Sorci, G., Gaillard, M. & Faivre, B. 2010. Age-dependent allocation of carotenoids to coloration versus antioxidant defences. Journal of Experimental Biology 213: 271-277 (pdf file).

Dattelbaum, J.D., Sieg, D., Manieri, C.M., Thomson, G. & Hill, M. 2010. Plasticity of acquired secondary metabolites in Clathria prolifera (Demospongia: Poecilosclerida): putative photoprotective role of carotenoids in a temperate intertidal sponge. The Open Marine Biology Journal 4: 87-95 ( pdf file).

del Cerro, S., Merino, S., Martínez-de la Puente, J., Lobato, E., Ruiz-de-Castańeda, R., Rivero-de Aguilar, J., Martínez, J., Morales, J., Tomás, G. & Moreno, J. 2010. Carotenoid-based plumage colouration is associated with blood parasite richness and stress protein levels in blue tits (Cyanistes caeruleus). Oecologia 162: 825-835 (pdf file).

Dugas, M. B. 2010. Nestling birds put their best flange forward. Journal of Avian Biology 41: 336-341 (pdf file).

Dunn, P. O., Garvin, J. C., Whittingham, L. A., Freeman-Gallant, C. R. & Hasselquist, D. 2010. Carotenoid and melanin-based ornaments signal similar aspects of male quality in two populations of the common yellowthroat. Functional Ecology 24: 149-158 (pdf file).

Edler, A. U. & Friedl, T. W. P. 2010. Individual quality and carotenoid-based plumage ornaments in male red bishops (Euplectes orix): plumage is not all that counts. Biological Journal of the Linnean Society 99: 384-397 (pdf file).

Edler, A.U. & Friedl, T.W.P. 2010. Plumage colouration, age, testosterone and dominance in male red bishops (Euplectes orix): a laboratory experiment. Ethology 116: 806-820 (pdf file).

Garner, S. R., Neff, B. D. & Bernards, M. A. 2010. Dietary carotenoid levels affect carotenoid and retinoid allocation in female Chinook salmon Oncorhynchus tshawytscha. Journal of Fish Biology 76: 1474-1790 (pdf file).

Gladbach, A., Gladbach, D.J., Kempenaers, B. & Quillfeldt, P. 2010. Female-specific colouration, carotenoids and reproductive investment in a dichromatic species, the upland goose Chloephaga picta leucoptera. Behav. Ecol. Sociobiol. 64: 1779-1789 (pdf file).

Gladbach, A., Gladbach, D. J. & Quillfeldt, P. 2010. Variations in leucocyte profiles and plasma biochemistry are related to different aspects of parental investment in male and female Upland geese Chloephaga picta leucoptera. Comparative Biochemistry and Physiology, Part A 156: 269-277 (pdf file).

Grether, G. F. (2010) The evolution of mate preferences, sensory biases, and indicator traits. In:  (Brockmann, H. J., Roper, T. J., Naguib, M., Wynne-Edwards, K., Mitani, J. C. & Simmons, L. W., eds.).Advances in the Study of Behavior, Vol. 41 Chapter 3, pp. 35-76 (pdf file).

Griggio, M., Hoi, H. & Pilastro, A. 2010. Plumage maintenance affects ultraviolet colour and female preference in the budgerigar. Behavioural Processes 84: 739-744 (pdf file).

Griggio, M., Zanollo, V. & Hoi, H. 2010. UV plumage color is an honest signal of quality in male budgerigars. Ecological Research 25: 1410-1703 (pdf file).

Hőrak, P., Sild, E., Soomets, U., Sepp, T. & Kilk, K. 2010. Oxidative stress and information content of black and yellow plumage coloration: an experiment with greenfinches. Journal of Experimental Biology 213: 2225-2233 (pdf file).

Huggins, K. A., Navara, K. J., Mendonça, M. T. & Hill, G. E. 2010. Detrimental effects of carotenoid pigments: the dark side of bright coloration. Naturwissenschaften 97: 637-644 (pdf file).

Kekäläinen, J., Valkama, H., Huuskonen, H. & Taskinen, J. 2010. Multiple sexual ornamentation signals male quality and predicts female preference in minnows. Ethology 116: 895-903 (pdf file).

Lindstedt, C., Morehouse, N., Pakkanen, H., Casas, J., Christides, J. P., Kemppainen, K., Lindstrom, L. & Mappes, J. 2010. Characterizing the pigment composition of a variable warning signal of Parasemia plantaginis larvae. Functional Ecology 24: 759-766 (pdf file).

Martínez-Padilla, J., Mougeot, F., Webster, L. M. I., Pérez-Rodríguez, L. & Piertney, S. B. 2010. Testing the interactive effects of testosterone and parasites on carotenoid-based ornamentation in a wild bird. Journal of Evolutionary Biology 23: 902-913 (pdf file).

Mougeot, F., Martínez-Padilla, J., Blount, J. D., Pérez-Rodríguez, L., Webster, L. M. I. & Piertney, S. B. 2010. Oxidative stress and the effect of parasites on a carotenoid-based ornament. Journal of Experimental Biology 213: 400-407 (pdf file).

Mougeot, F., Martínez-Padilla, J., Bortolotti, G. R., Webster, L. M. I. & Piertney, S. B. 2010. Physiological stress links parasites to carotenoid-based colour signals. Journal of Evolutionary Biology 23: 643-650 (pdf file).

Navarro, C., Pérez-Contreras, T., Avilés, J. M., McGraw, K. J. & Soler, J. J. 2010. Beak colour reflects circulating carotenoid and vitamin A levels in spotless starlings (Sturnus unicolor). Behavioral Ecology and Sociobiology 64: 1057-1067 (pdf file).

Pérez, C., Lores, M. & Velando, A. 2010. Oil pollution increases plasma antioxidants but reduces coloration in a seabird. Oecologia 163: 875-884 (pdf file).

Pérez-Rodríguez, L., Mougeot, F. & Alonso-Alvarez, C. 2010. Carotenoid-based coloration predicts resistance to oxidative damage during immune challenge. Journal of Experimental Biology 213: 1685-1690 (pdf file).

Prager, M. & Andersson, S. 2010. Convergent evolution of red carotenoid coloration in widowbirds and bishops (Euplectes spp.). Evolution 64: 3609-3619 (pdf file).

Rémy, A., Grégoire, A., P., P. & C., D. 2010. Mediating male–male interactions: the role of the UV blue crest coloration in blue tits. Behav. Ecol. Sociobiol. 64: 1839-1847 (pdf file).

Sternalski, A., Mougeot, F., Eraud, C., Gangloff, B., Villers, A. & Bretagnolle, V. 2010. Carotenoids in nestling Montagu's harriers: variations according to age, sex, body condition and evidence for diet-related limitations. Journal of Comparative Physiology, Part B 180: 33-43 (pdf file).

Toomey, M. B., Butler, M. W. & McGraw, K. J. 2010. Immune-system activation depletes retinal carotenoids in house finches (Carpodacus mexicanus). Journal of Experimental Biology 213: 1709-1716  (pdf file).

Torres, R. & Velando, A. (2010) Color in a long-lived tropical seabird. Sexual selection in a life-history context. In: Behavioral Ecology of Tropical Animals, Vol. 42 (Macedo, R., ed.). pp. 155-188 Advances in the Study of Behavior (pdf file).

Veuille, M. 2010. Darwin and sexual selection: One hundred years of misunderstanding. Comptes Rendus - Biologies 333: 145-156 (pdf file).

Vinkler, M. & Albrecht, T. 2010. Carotenoid maintenance handicap and the physiology of carotenoid-based signalisation of health. Naturwissenschaften 97: 19-28 (pdf file).



Casagrande, S., Dell'Omo, G., Costantini, D., Tagliavini, J. & Groothuis, T. 2011. Variation of a carotenoid-based trait in relation to oxidative stress and endocrine status during the breeding season in the Eurasian kestrel: a multi-factorial study. Comparative Biochemistry and Physiology - A 160: 16-26 (pdf file).

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García-Navas, V., Ferrer, E.S., and Sanz, J.J. 2012. Plumage yellowness predicts foraging ability in the blue tit Cyanistes caeruleus. Biological Journal of the Linnean Society 106(2): 418-429 (pdf file).

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Kelly, R.J., Murphy, T.G., Tarvin, K.A., and Burness, G. 2012. Carotenoid-based ornaments of female and male American goldfinches (Spinus tristis) show sex-specific correlations with immune function and metabolic rate. Physiological and Biochemical Zoology 85(4): 348-363 (pdf file).

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Legagneux, P., Clark, R.G., Guillemain, M., Eraud, C., Théry, M., and Bretagnolle, V. 2012. Large-scale geographic variation in iridescent structural ornaments of a long-distance migratory bird. Journal of Avian Biology 43(4): 355-361 (pdf file).

Maia, R., Brasileiro, L., Lacava, R.V., and Macedo, R.H. 2012. Social environment affects acquisition and color of structural nuptial plumage in a sexually dimorphic tropical passerine. PLoS One 7(10): e47501 (pdf file).

Mitrus, C., Mitrus, J., and Sikora, M. 2012. Badge size and arrival time predict mating success of red-breasted flycatcher Ficedula parva males. Zoological Science 29(12): 795-799 (pdf file).

Orledge, J.M., Blount, J.D., Hoodless, A.N., and Royle, N.J. 2012. Antioxidant supplementation during early development reduces parasite load but does not affect sexual ornament expression in adult ring-necked pheasants. Functional Ecology 26(3): 688-700 (pdf file).

Rosenthal, M.F., Murphy, T.G., Darling, N., and Tarvin, K.A. 2012. Ornamental bill color rapidly signals changing condition. Journal of Avian Biology 43(6): 553-564 (pdf file).

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Simons, M.J.P., Briga, M., Koetsier, E., Folkertsma, R., Wubs, M.D., Dijkstra, C., and Verhulst, S. 2012a. Bill redness is positively associated with reproduction and survival in male and female zebra finches. PLoS One 7(7): e40721 (pdf file).

Simons, M.J.P., Cohen, A.A., and Verhulst, S. 2012b. What does carotenoid-dependent coloration tell? Plasma carotenoid level signals immunocompetence and oxidative stress state in birds - a meta-analysis. PLoS One 7(8): e43088 (pdf file).

Stephen, I.D., Scott, I.M.L., Coetzee, V., Pound, N., Perrett, D.I., and Penton-Voak, I. 2012. Cross-cultural effects of color, but not morphological masculinity, on perceived attractiveness of men's faces. Evolution and Human Behavior 33(4): 260-267 (pdf file).

Sternalski, A., Mougeot, F., and Bretagnolle, V. 2012a. Phenotypic variation in nestlings of a bird of prey under contrasting breeding and diet conditions. Biological Journal of the Linnean Society 107(4): 799-812 (pdf file).

Sternalski, A., Mougeot, F., Parker, N.R., Pérez-Rodríguez, L., and Bretagnolle, V. 2012b. Carotenoid-based coloration, condition, and immune responsiveness in the nestlings of a sexually dimorphic bird of prey. Physiological and Biochemical Zoology 85(4): 364-375 (pdf file).

Theis, A., Salzburger, W., and Egger, B. 2012. The function of anal fin egg-spots in the cichlid fish Astatotilapia burtoni. PLoS One 7(1): e29878 (pdf file).

Whitehead, R.D. 2012. Attractive skin coloration: harnessing sexual selection to improve diet and health. Evolutionary Psychology 10(5): 842-854 (pdf file).

Zanollo, V., Griggio, M., Robertson, J., and Kleindorfer, S. 2012. The number and coloration of white flank spots predict the strength of a cutaneous immune response in female Diamond Firetails, Stagonopleura guttata. Journal of Ornithology 153(4): 1233-1244 (pdf file).


zip file 2010-2012




Alfonso, Y.U., Morris, H.J., Gutiérrez, A., Rodríguez-Schettino, L., Denis, D., and Steffen, J.E. 2013. Dewlap color variation based on pterin and carotenoid pigments in three subspecies of anolis jubar of the Cuban southern coast. Copeia 2013(2): 201-205 (pdf file).

Blanco, G., Hornero-Méndez, D., Lambertucci, S.A., Bautista, L.M., Wiemeyer, G., Sanchez-Zapata, J.A., Garrido-Fernández, J., Hiraldo, F., and Donázar, J.A. 2013. Need and seek for dietary micronutrients: endogenous regulation, external signalling and food sources of carotenoids in New World vultures. PLoS One 8(6): e65562 (pdf file).

Brown, A.C., McGraw, K.J., and Clotfelter, E.D. 2013. Dietary carotenoids increase yellow nonpigment coloration of female convict cichlids (Amantitlania nigrofasciata). Physiological and Biochemical Zoology 86(3): 312-322 (pdf file).

Cook, E.G., Murphy, T.G., and Johnson, M.A. 2013. Colorful displays signal male quality in a tropical anole lizard. Naturwissenschaften 100: 993-996 (pdf file).

Del Val E, Negro JJ, Senar JC (2013) Seasonal variaiton of circulating plasma carotenoids in Mediterranean Great Tit Parus major. Bird Study 60(4): 459-465 (pdf file).

Doutrelant, C., Grégoire, A., Gomez, D., Staszewski, V., Arnoux, E., Tveraa, T., Faivre, B., and Boulinier, T. 2013. Colouration in Atlantic puffins and blacklegged kittiwakes: monochromatism and links to body condition in both sexes. Journal of Avian Biology 44(5): 451-460 (pdf file).

Garcia-de Blas, E., Rafael, M., Vinuela, J., Perez-Rodriguez, L., and Alonso-Alvarez, C. 2013. Free and Esterified Carotenoids in Ornaments of an Avian Species: The Relationship to Color Expression and Sources of Variability. Physiological and Biochemical Zoology 86(5): 483-498 (pdf file).

Giraudeau, M., Sweazea, K., Butler, M.W., and McGraw, K.J. 2013. Effects of carotenoid and vitamin E supplementation on oxidative stress and plumage coloration in house finches (Haemorhous mexicanus). Comparative Biochemistry and Physiology - Part A 166(3): 406-413 (pdf file).

Gotanda, K., Delaire, L., Raeymaekers, J., Pérez-Jvostov, F., Dargent, F., Bentzen, P., Scott, M., Fussmann, G., and Hendry, A. 2012. Adding parasites to the guppy-predation story: insights from field surveys. Oecologia 172(1): 155-166 (pdf file) (pdf file).

Hamilton, D.G., Whiting, M.J., and Pryke, S.R. 2013. Fiery frills: carotenoid-based coloration predicts contest success in frillneck lizards. Behavioral Ecology 24(5): 1138-1149 (pdf file).

Hudon, J., Derbyshire, D., Leckie, S., and Flinn, T. 2013. Diet-induced plumage erythrism in Baltimore orioles as a result of the spread of introduced shrubs. The Wilson Journal of Ornithology 125(1): 88-96 (pdf file).

Johnson, J.D., and Hill, G.E. 2013. Is carotenoid ornamentation linked to the inner mitochondria membrane potential? A hypothesis for the maintenance of signal honesty. Biochimie 95(2): 436-444 (pdf file).

Lahaye, S.E.P., Eens, M., Darras, V.M., and Pinxten, R. 2013. Hot or not: the effects of exogenous testosterone on female attractiveness to male conspecifics in the budgerigar. PLoS ONE 8(8): e74005 (pdf file).

Leniowski, K., and Węgrzyn, E. 2013. The carotenoid-based red cap of the Middle Spotted Woodpecker Dendrocopos medius reflects individual quality and territory size. Ibis 155(4): 804-813 (pdf file).

Lozano, G. A., Lank, D. B., & Addison, B. (2013). Immune and oxidative stress trade-offs in four classes of ruffs (Philomachus pugnax) with different alternative reproductive strategies. Canadian Journal of Zoology, 91(4), 212-218 (pdf file).

Mehlis, M., & Bakker, T. C. M. (2013). Male reproductive traits of full-sibs of different age classes in three-spined sticklebacks (Gasterosteus). Springer Plus, 2, 175 (pdf file).

Midamegbe, A., Grégoire, A., Staszewski, V., Perret, P., Lambrechts, M.M., Boulinier, T., and Doutrelant, C. 2013. Female blue tits with brighter yellow chests transfer more carotenoids to their eggs after an immune challenge. Oecologia 173(2): 387-397 (pdf file).

Pérez-Rodríguez, L., Martínez-Padilla, J., and Mougeot, F. 2013. Carotenoid-Based Ornaments as Signals of Health Status in Birds: Evidences from Two Galliform Species, the Red-Legged Partridge (Alectoris rufa) and the Red Grouse (Lagopus lagopus scoticus). In Carotenoids: Food Sources, Production and Health Benefits. Edited by M. Yamaguchi. Nova Science Publishers, Hauppauge, New York. pp. 173-198 (pdf file).

Pickett, S.R.A., Weber, S.B., McGraw, K.J., Norris, K.J., and Evans, M.R. 2013. Environmental and parental influences on offspring health and growth in great tits Parus major. PLoS One 8(7): e69695 (pdf file).

Polo-Cavia, N., López, P., and Martín, J. 2013. Head coloration reflects health state in the red-eared slider Trachemys scripta elegans. Behavioral Ecology and Sociobiology 67(1): 153-162 (pdf file).

Rahman, M.M., Kelley, J.L., and Evans, J.P. 2013. Condition-dependent expression of pre- and postcopulatory sexual traits in guppies. Ecology and Evolution 3(7): 2197-2213 (pdf file).

Synek, P., Albrecht, T., Vinkler, M., Schnitzer, J., Votýpka, J., and Munclinger, P. 2013. Haemosporidian parasites of a European passerine wintering in South Asia: diversity, mixed infections and effect on host condition. Parasitology Research 112(4): 1667-1677 (pdf file).

Vroonen, J., Vervusrt, B., and Van Damme, R. 2013. Melanin-based colouration as a potential indicator of male quality in the lizard Zootoca vivipara (Squamata: Lacertidae). Amphibia-Reptilia 34(3): 539-549 (pdf file).

Weiss, S.L., Mulligan, E.E., Wilson, D.S., and Kabelik, D. 2013. Effect of stress on female-specific ornamentation. Journal of Experimental Biology 216(14): 2641-2647 (pdf file).

Yang, C., Wang, J., Fang, Y., and Sun, Y.-H. 2013. Is sexual ornamentation an honest signal of male quality in the Chinese grouse (Tetrastes sewerzowi)? PLoS One 8(12): e82972 (pdf file).




Brown, A.C., Leonard, H.M., McGraw, K.J., and Clotfelter, E.D. 2014. Maternal effects of carotenoid supplementation in an ornamented cichlid fish. Functional Ecology 28(3): 612-620 (pdf file).

Costantini, D. 2014. The costs of make-up in sexual selection and social signalling. In Oxidative Stress and Hormesis in Evolutionary Ecology and Physiology: a marriage between mechanistic and evolutionary approaches. Edited by D. Costantini. Springer-Verlag, Berlin. pp. 171-204.

Fairhurst, G.D., Dawson, R.D., Oort, H., and Bortolotti, G.R. 2014. Synchronizing feather-based measures of corticosterone and carotenoid-dependent signals: what relationships do we expect? Oecologia 174(3): 689-698 (pdf file).

Garcia-de Blas, E., Mateo, R., and Alonso-Alvarez, C. 2014. Accumulation of dietary carotenoids, retinoids and tocopherol in the internal tissues of a bird: a hypothesis for the cost of producing colored ornaments. Oecologia 177(1): 259-271 (pdf file).

Garcia-de Blas, E., Mateo, R., Guzman Bernardo, F.J., Rodriguez Martin-Domeadios, R.C., and Alonso-Alvarez, C. 2014. Astaxanthin and papilioerythrinone in the skin of birds: a chromatic convergence of two metabolic routes with different precursors? Naturwissenschaften 101(5): 407-416 (pdf file).

Giraudeau, M., and McGraw, K.J. 2014. Physiological correlates of urbanization in a desert songbird. Integrative and Comparative Biology 54(4): 622-632 (pdf file).

Jones, T.B., Drake, A., and Green, D.J. 2014. Individual quality explains association between plumage colouration, arrival dates and mate acquisition in yellow warblers (Setophaga petechia). BMC Ecology 14 (pdf file).

Kopena, R., Lopez, P., and Marin, J. 2014. What are carotenoids signaling? Immunostimulatory effects of dietary vitamin E, but not of carotenoids, in Iberian green lizards. Naturwissenschaften 101(12): 1107-1114 (pdf file).

Lahaye, S.E.P., Eens, M., Darras, V.M., and Pinxten, R. 2014. Bare-part color in female budgerigars changes from brown to structural blue following testosterone treatment but Is not strongly masculinized. PLoS One 9(1): e86849 (pdf file).

Leităo, A.V., Monteiro, A.H., and Mota, P.G. 2014. Ultraviolet reflectance influences female preference for colourful males in the European serin. Behavioral Ecology and Sociobiology 68(1): 63-72 (pdf file).

Lucas, A., Morales, J., and Velando, A. 2014. Differential effects of specific carotenoids on oxidative damage and immune response of gull chicks. Journal of Experimental Biology 217: 1253-1262 (pdf file).

Männiste, M., and Hőrak, P. 2014. Emerging infectious diseaese selects for darker plumage in greenfinches. Frontiers in Ecology and Evolution 2: Article 4 (1-6) (pdf file).

Martinez-Padilla, J., Perez-Rodriguez, L., Mougeot, F., Ludwig, S., and Redpath, S.M. 2014. Intra-sexual competition alters the relationship between testosterone and ornament expression in a wild territorial bird. Hormones and Behavior 65(5): 435-444 (pdf file).

Mikulas, J.E., and Brooks, D.M. 2014. A case of hermaphorditism in a wood duck (Aix sponsa). Bulletin of the Texas Ornithological Society 43(1): 63-65 (pdf file).

Sefc, K.M., Brown, A.C., and Clotfelter, E.D. 2014. Carotenoid-based coloration in cichlid fishes. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology 173: 42-51 (pdf file).

Simons, M.J.P., Maia, R., Leenknegt, B., and Verhulst, S. 2014. Carotenoid-dependent signals and the evolution of plasma carotenoid levels in birds. American Naturalist 184(6): 741-751 (pdf file).

Steffen, J.E., and Guyer, C.C. 2014. Display behaviour and dewlap colour as predictors of contest success in brown anoles. Biological Journal of the Linnean Society 111(3): 646-655 (pdf file).

Walker, L.K., Thorogood, R., Karadas, F., Raubenheimer, D., Kilner, R.M., and Ewen, J.G. 2014. Foraging for carotenoids: do colorful male hihi target carotenoid-rich foods in the wild? Behavioral Ecology 25(5): 1048-1057 (pdf file).



Anderson, C., Wong, S.C., Fuller, A., Zigelsky, K. and Earley, R.L. 2015. Carotenoid-based coloration is associated with predation risk, competition, and breeding status in female convict cichlids (Amatitlania siquia) under field conditions. Env. Biol. Fishes, 98: 1005-1013.

Beck, M. L., W. A. Hopkins, and D. M. Hawley. 2015. Relationships among plumage coloration, blood selenium concentrations, and immune responses of adult and nestling tree swallows. The Journal of Experimental Biology 218:3415-3424.

Casagrande, S., Pinxten, R., Zaid, E., and Eens, M. 2015. Birds receiving extra carotenoids keep singing during the sickness phase induced by inflammation. Behavioral Ecology and Sociobiology 69(6): 1029-1037.

Davis, A.K. and Altizer, S. 2015. New Perspectives on Monarch Migration, Evolution, and Population Biology: An Overview. In Monarchs in a Changing World: Biology and Conservation of an Iconic Butterfly (K.S. Oberhauser, K.R. Nail and S. Altizer, eds), p. 352. Ithaca: Cornell University Press.

Driessens, T., Huyghe, K., Vanhooydonck, B. and Van Damme, R. 2015. Messages conveyed by assorted facets of the dewlap, in both sexes of Anolis sagrei. Behav. Ecol. Sociobiol., 69: 1251-1264.

Heinen-Kay, J.L., Morris, K.E., Ryan, N.A., Byerley, S.L., Venezia, R.E., Peterson, M.N. and Langerhans, R.B. 2015. A trade-off between natural and sexual selection underlies diversification of a sexual signal. Behav. Ecol., 26(2): 533-542.

Jacob, S., Parthuisot, N., Vallat, A., Ramon-Portugal, F., Helfenstein, F., and Heeb, P. 2015. Microbiome affects egg carotenoid investment, nestling development and adult oxidative costs of reproduction in Great tits. Functional Ecology 29: 1048-1058..

Johnson, A. M. and R. C. Fuller (2015). "The meaning of melanin, carotenoid, and pterin pigments in the bluefin killifish, Lucania goodei." Behavioral Ecology 26: 158-167.

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Leclaire, S., Bourret, V., Blanchard, P., de Franceschi, C., Merkling, T., Hatch, S.A. and Danchin, E. 2015. Carotenoids increase immunity and sex specifically affect color and redox homeostasis in a monochromatic seabird. Behav. Ecol. Sociobiol., 69: 1097-1111.

Leitao, A.V., Ferreira, A.C., Funghi, C., Trigo, S. and Mota, P.G. 2015. Evidence for multiple functions in a sexually selected ornament. Anim. Behav., 110: 155-161.

Lopez-Rull, I., Hornero-Mendez, D., Frias, O. and Blanco, G. 2015. Age-related relationships between innate immunity and plasma carotenoids in an obligate avian scavenger. PLoS One, 10: e0141759.

Mehlis, M., Rick, I.P. and Bakker, T.C.M. 2015. Dynamic resource allocation between pre- and postcopulatory episodes of sexual selection determines competitive fertilization success. Proceedings of the Royal Society of London B: Biological Sciences, 282.

Plasman, M., Reynoso, V.H., Nicolas, L. and Torres, R. 2015. Multiple colour traits signal performance and immune response in the Dickerson’s collared lizard Crotaphytus dickersonae. Behav. Ecol. Sociobiol., 69: 765-775.

Reudink, M. W., A. E. McKellar, K. L. D. Marini, S. L. McArthur, P. P. Marra, and L. M. Ratcliffe. 2015. Inter-annual variation in American redstart (Setophaga ruticilla) plumage colour is associated with rainfall and temperature during moult: an 11-year study. Oecologia 178:161-173.

Simons, M.J.P., Groothuis, T.G.G., and Verhulst, S. 2015. An appraisal of how the vitamin A-redox hypothesis can maintain honesty of carotenoid-dependent signals. Ecology and Evolution 5(1): 224-228.

Steffen, J.E., Learn, K.M., Drumheller, J.S., Boback, S.M. and McGraw, K.J. 2015. Carotenoid composition of colorful body stripes and patches in the painted turtle (Chrysemys picta) and red-eared slider (Trachemys scripta). Chelonian Conservation and Biology, 14: 56-63.

Theis, A., T. Bosia, T. Roth, W. Salzburger, and B. Egger. 2015. Egg-spot pattern and body size asymmetries influence male aggression in haplochromine cichlid fishes. Behavioral Ecology 26(6):1512-1519.

Tian, H., Wu, P., Wang, W. and Ru, S. 2015. Disruptions in aromatase expression in the brain, reproductive behavior, and secondary sexual characteristics in male guppies (Poecilia reticulata) induced by tributyltin. Aquat. Toxicol., 162: 117-125.

Trigo, S. and Mota, P.G. 2015. What is the value of a yellow patch? Assessing the signalling role of yellow colouration in the European serin. Behav. Ecol. Sociobiol., 69: 481-490.

Wilgers, D.J., and Hebets, E.A. 2015. Functional Approach to Condition. In Animal Signaling and Function. Edited by D. J. Irschick, M. Briffa, and J. Podos. John Wiley & Sons, Inc., Hoboken, NJ, USA. pp. 229-252.




 Brown, A.C., Cahn, M.D., Choi, S., and Clotfelter, E.D. 2016. Dietary carotenoids and bacterial infection in wild and domestic convict cichlids (Amatitlania spp.). Environmental Biology of Fishes 99(4): 439-449.

Dias, R.I., Manica, L.T., Gressler, D., Bell, J.A., and Fecchio, A. 2016. Plumage coloration, body condition and immunological status in yellow-billed cardinals (Paroaria capitata). Ethology, Ecology and Evolution 28(4): 462-476.

Garcia-de Blas, E., Mateo, R., and Alonso-Alvarez, C. 2016. Specific carotenoid pigments in the diet and a bit of oxidative stress in the recipe for producing red carotenoid-based signals. PeerJ 4: e2237.

Horak, P., and M. Manniste. 2016. Viability selection affects black but not yellow plumage colour in greenfinches. Oecologia 180 (1):23-32

Lee, S.-i., Kim, M., Choe, J.C., and Jablonski, P.G. 2016. Evolution of plumage coloration in the crow family (Corvidae) with a focus on the color-producing microstructures in the feathers: a comparison of eight species. Animal Cells and Systems 20(2): 95-102.

Lifshitz, N., and St Clair, C.C. 2016. Coloured ornamental traits could be effective and non-invasive indicators of pollution exposure for wildlife. Conservation Physiology 4(1): cow028.

 Lim, H.H., and Pike, T.W. 2016. Dietary carotenoid availability affects avian color discrimination. Behavioral Ecology 27(6): 1579-1584.

López Rull, I., Nicolás, L., Neri-Vera, N., Argáez, V., Martínez, M., and Torres, R. 2016. Assortative mating by multiple skin color traits in a seabird with cryptic sexual dichromatism. Journal of Ornithology XX(oo): 1-14.

Perez-Rodriguez, L., E. Garcia de Blas, J. Martinez-Padilla, F. Mougeot, and R. Mateo. 2016. Carotenoid profile and vitamins in the combs of the red grouse (Lagopus lagopus scoticus): implications for the honesty of a sexual signal. Journal of Ornithology 157(1): 145-157

Rowe, M., Pierson, K.L., and McGraw, K.J. 2016. Exploratory behavior is associated with plasma carotenoid accumulation in two congeneric species of waterfowl. Behavioural Processes 115: 181-190.

Tomasek, O., Gabrielova, B., Kacer, P., Marsik, P., Svobodova, J., Syslova, K., Vinkler, M. and Albrecht, T. 2016. Opposing effects of oxidative challenge and carotenoids on antioxidant status and condition-dependent sexual signalling. Scientific Reports, 6: Article number 23546.



 Abrahăo, M. and Alves, M. A. S. 2017. Sexual selection in birds: Influence of ambient light and conspicuity on the reproductive success of males. Oecologia Australis 21(4): 361-373.

Broughton, D. R., Schneider, B. C., McGraw, K. J. and Ardia, D. R. 2017. Carotenoids buffer the acute phase response on fever, sickness behavior and rapid bill color change in zebra finches. Journal of Experimental Biology 220(16): 2957-2964.

Burns, K. J., McGraw, K. J., Shultz, A. J., Stoddard, M. C. and Thomas, D. B. 2017: Advanced Methods for Studying Pigments and Coloration Using Avian Specimens In: The Extended Specimen:Emerging Frontiers in Collections-Based Ornithological Research. (Webster, M. A., ed). CRC Press, Boca Raton.

Czyż, M. J. and Gołdyn, B. 2017. Relative scaling and colouration of cercopodes in Branchipus schaefferi (Crustacea: Branchiopoda). Biologia 72(3): 319-324.

Devigili, A., Belluomo, V., Locatello, L., Rasotto, M. B. and Pilastro, A. 2017. Postcopulatory cost of immune system activation in Poecilia reticulata. Ethology Ecology & Evolution 29(3): 266-279.

Fan, M., Hall, M. L., Kingma, S. A., Mandeltort, L. M., Hidalgo Aranzamendi, N., Delhey, K. and Peters, A. 2017. No fitness benefits of early molt in a fairy-wren: relaxed sexual selection under genetic monogamy? Behavioral Ecology 28(4): 1055-1067.

Foo, Y. Z., Rhodes, G. and Simmons, L. W. 2017. The carotenoid beta-carotene enhances facial color, attractiveness and perceived health, but not actual health, in humans. Behavioral Ecology 28(2): 570-578.

Foo, Y. Z., Simmons, L. W. and Rhodes, G. 2017. Predictors of facial attractiveness and health in humans. Scientific Reports 7: 39731.

Freeman-Gallant, C. R. and Taff, C. C. 2017. Age-specific patterns of infection with haemosporidians and trypanosomes in a warbler: implications for sexual selection. Oecologia 184(4): 813-823.

García-Heras, M.-S., Arroyo, B., Simmons, R. E., Camarero, P. R., Mateo, R., García, J. T. and Mougeot, F. 2017. Pollutants and diet influence carotenoid levels and integument coloration in nestlings of an endangered raptor. Science of The Total Environment 603–604: 299-307.

Henschen, A. E., Whittingham, L. A. and Dunn, P. O. 2017. The relationship between blood parasites and ornamentation depends on the level of analysis in the common yellowthroat. Journal of Avian Biology 48(9): 1263-1272.

Koch, R. E., Josefson, C. C. and Hill, G. E. 2017. Mitochondrial function, ornamentation, and immunocompetence. Biological Reviews 92(3): 1459-1474.

Lee, A.-I., Baek, M. J., Choe, J. C. and Jablonski, P. G. 2017. Characterisation of the major carotenoids in the plasma of the Eurasian magpie (Pica pica) nestlings. Korean Journal of Ornithology 24(1): 1-9.

Lehnert, S. J., Devlin, R. H., Pitcher, T. E., Semeniuk, C. A. D. and Heath, D. D. 2017. Redder isn’t always better: cost of carotenoids in Chinook salmon eggs. Behavioral Ecology 28(2): 549-555.

Theis, A., Roth, O., Cortesi, F., Ronco, F., Salzburger, W. and Egger, B. 2017. Variation of anal fin egg-spots along an environmental gradient in a haplochromine cichlid fish. Evolution 71(3): 766-777.

Václav, R. and Kolníková, Z. 2017. Effects of food and thermal regimes on body condition indices and skin colouration in corn snakes. Biologia 72: 84.

Wilkins, L. G. E., Marques da Cunha, L., Menin, L., Ortiz, D., Vocat-Mottier, V., Hobil, M., Nusbaumer, D. and Wedekind, C. 2017. Maternal allocation of carotenoids increases tolerance to bacterial infection in brown trout. Oecologia 185(3): 351-363.


zip file 2013-2017



Amundsen, T. 2018. Sex roles and sexual selection: lessons from a dynamic model system. Current Zoology 64(3): 363-392.

Bruch, E. E., and M. E. J. Newman. 2018. Aspirational pursuit of mates in online dating markets. Science Advances 4:eaap9815.

Busso, J. P. and Blanckenhorn, W. U. 2018. Disruptive sexual selection on male body size in the polyphenic black scavenger fly Sepsis thoracica. Behavioral Ecology 29(3): 769-777.

Cantarero, A., Pilastro, A. and Griggio, M. 2018. Nestling sex ratio is associated with both male and female attractiveness in rock sparrows. Journal of Avian Biology 49(8): e01666.

Freeman-Gallant, C. R. and Taff, C. C. 2018. Age and infection history are revealed by different ornaments in a warbler. Oecologia 188(4): 1025–1036.

Hudson, S. B. and Wilcoxen, T. E. 2018. Condition dependence of structural plumage coverage in Indigo Buntings Passerina cyanea. Avian Biology Research 11(1): 54-61.

Janas, K., Podmokła, E., Lutyk, D., Dubiec, A., Gustafsson, L., Cichoń, M. and Drobniak, S. 2018. Influence of haemosporidian infection status on structural and carotenoid-based colouration in the blue tit (Cyanistes caeruleus). Journal of Avian Biology 49(10): e01840.

Karvonen, A., Wagner, C. E., Selz, O. M. and Seehausen, O. 2018. Divergent parasite infections in sympatric cichlid species in Lake Victoria. Journal of Evolutionary Biology 31(9): 1313-1329.

Knight, A., Ewen, J. G., Brekke, P. and Santure, A. W. 2018: The evolutionary biology, ecology and epidemiology of Coccidia of passerine birds. In: Advances in Parasitology. Academic Press.

Koch, R. E. and Hill, G. E. 2018. Do carotenoid‐based ornaments entail resource tradeoffs? An evaluation of theory and data. Funtional Ecology 32(8): 1908-1920.

Montoya, B., Flores, C. and Torres, R. 2018. Repeatability of a dynamic sexual trait: Skin color variation in the Brown Booby (Sula leucogaster). Auk 135(3): 622-636.

Pardal, S., Alves, J., A., Mota, P., G. and Ramos, J., A. 2018. Dressed to impress: breeding plumage as a reliable signal of innate immunity. Journal of Avian Biology 49(7): e01579.

Parolini, M., Iacobuzio, R., Possenti, C. D., Bassano, B., Pennati, R. and Saino, N. 2018. Carotenoid-based skin coloration signals antioxidant defenses in the brown trout (Salmo trutta). Hydrobiologia 815(1): 267-280.

Pike, T. W. 2018. Discrimination of signal carotenoid content using multidimensional chromatic information. Behavioral Ecology 29(1): 87-92.

Scott, R. J. and Black, C. A. 2018. An experimental examination of dietary impacts on nuptial signal expression in threespine stickleback: does the nuptial signal indicate carotenoid and lipid stores? Experimental Biology of Fishes 101(5): 687-698.

Sumasgutner, P., Adrion, M. and Gamauf, A. 2018. Carotenoid coloration and health status of urban Eurasian kestrels (Falco tinnunculus). PLoS One 13(2): e0191956.

Svobodová, J., Bauerová, P., Eliáš, J., Velová, H., Vinkler, M. and Albrecht, T. 2018. Sperm variation in Great Tit males (Parus major) is linked to a haematological health-related trait, but not ornamentation. Journal of Ornithology 159(3): 815-822.

Takahashi, D. 2018. Female mate choice based on male nuptial coloration in Pale Chub, Zacco platypus. Zoological Science 35(1): 23-27.

Weaver, R. J., Santos, E. S. A., Tucker, A. M., Wilson, A. E. and Hill, G. E. 2018. Carotenoid metabolism strengthens the link between feather coloration and individual quality. Nature Communications 9(1): 73.

Wojan, E. M., Bertram, S. M., Clendenen, D. A., Castillo, C., Neldner, H. M. and Kolluru, G. R. 2018. Sexual selection on the multicomponent display of black morph male Girardinus metallicus (Pisces: Poeciliidae). Behavioural Processes 153: 1-8.

Wright, A., E., Fumagalli, M., Cooney Christopher, R., Bloch Natasha, I., Vieira Filipe, G., Buechel Severine, D., Kolm, N. and Mank Judith, E. 2018. Male-biased gene expression resolves sexual conflict through the evolution of sex-specific genetic architecture. Evolution Letters 2(2): 52-61.



 Agbali M, Smith C (2019) A role for receiver bias in rose bitterling mate choice? International Journal of Applied Science 1:386-395.

Cantarero A, Mateo R., Camarero P, Alonso D, Fernandez-Eslava B., Alonso-Alvarez, C (2019) A mitochondria-targeted antioxidant affects the carotenoid-based plumage of red crossbills. bioRxiv:839670.

Engel KC, Hwang W, Steiger S (2019) A pheromone that coordinates parental care is evolutionary conserved among burying beetles (Silphidae: Nicrophorus). Chemoecology 29:1-9.

Erritouni YR, Reinke BA, Calsbeek R (2019) A novel body coloration phenotype in Anolis sagrei: Implications for physiology, fitness, and predation. PLoS One 13:e0209261.

Franklin AM, Donatelli CM, Culligan CR, Tytell ED (2019) Meral-spot reflectance signals weapon performance in the mantis shrimp Neogonodactylus oerstedii (Stomatopoda). Biological Bulletin 236:43-54.

Leclaire S, Bourret V, Pineaux M, Blanchard P, Danchin E, Hatch SA (2019) Red coloration varies with dietary carotenoid access and nutritional condition in kittiwakes. J. Exp. Biol. 222:jeb210237.

Marson KM, Taylor DS, Earley RL (2019) Cryptic male phenotypes in the mangrove rivulus fish, Kryptolebias marmoratus. Biological Bulletin 236:13-28.

Mieno A, Karino K (2019) Male mate preference for female coloration in a Cyprinid fish, Puntius titteya. Zool. Sci. 36:504-510.

Mos B, Dworjanyn SA (2019) Ready to harvest? Spine colour predicts gonad index and gonad colour rating of a commercially important sea urchin. Aquaculture 505:510-516.

Ogita M, Karino K (2019) Effect of body coloration on male–male competition in a cyprinid fish Puntius titteya. Zool. Sci. 36:141-146.

Segura LN, Mahler B (2019) Male Red-crested Cardinal plumage coloration is associated with parental abilities and breeding performance. Scientific Reports 9:10958.

Wojan EM, Carreiro NC, Clendenen DA, Neldner HM, Castillo C, Bertram SM, Kolluru GR (2019) The effects of commonly used anaesthetics on colour measurements across body regions in the poeciliid fish, Girardinus metallicus. J. Fish Biol. 95:1320-1330.



 Cantarero A, Mateo R, Camarero P, Alonso D, Fernandez-Eslava B, Alonso-Alvarez C (2020) Testing the shared-pathway hypothesis in the carotenoid-based coloration of red crossbills. Evolution 74:2348-2364.

Costantini D, Dell'Omo DG (2020) The Kestrel. Cambridge University Press

Earley RL, Anderson CT, Moscicki MK, Norton BB, Brown AC, Clotfelter ED (2020) Carotenoid availability and tradeoffs in female convict cichlids, a reverse sexually-dichromatic fish. Env. Biol. Fish. 103:1541–1552.

Fogelholm J, Henriksen R, Höglund A, Huq N, Johnsson M, Lenz R, Jensen P, Wright D (2020) CREBBP and WDR 24 Identified as candidate genes for quantitative variation in red-brown plumage colouration in the chicken. Scientific Reports 10:1161.

García-Campa J, Müller W, González-Braojos S, García-Juárez E, Morales J (2020) Dietary carotenoid supplementation facilitates egg laying in a wild passerine. Ecol. Evol. 10: 4968-4978.

Hund AK, Hubbard JK, Albrecht T, Vortman Y, Munclinger P, Krausová S, Tomášek O, Safran RJ (2020) Divergent sexual signals reflect costs of local parasites. Evolution 74:2404-2418.

LaPlante LH, Delaney S (2020) Male mate choice for a female ornament in a monogamous cichlid fish, Mikrogeophagus ramirezi. J. Fish Biol. 96:663-668.

Longo AV, Rodríguez-Gómez CA, Zegarra JP, Monzón O, Claudio-Hernández HJ, Joglar RL, Zamudio KR, Burrowes PA, López-Torres AL (2020) Tick parasitism as a cost of sexual selection and male parental care in a Neotropical frog. Ecosphere 11:e03010.

Nguyen, C. D. H., G. Amoroso, T. Ventura, J. J. Minich, and A. Elizur. 2020. Atlantic Salmon (Salmo salar L., 1758) gut microbiota profile correlates with flesh pigmentation: cause or effect? Marine Biotechnology 22:786-804.

Prasetya AM, Peters A, Delhey K (2020) Carotenoid-based plumage colour saturation increases with temperature in Australian passerines. Journal of Biogeography 47:2671-2683.

Sawant PB, Chakravarty S, Dasgupta S, Chadha NK, Sawant BT (2020) The quintessence of colour enhancement in ornamental fishes: an empirical pathway towards rainbow revolution. Current Science 119:1093-1100.

Stephenson J, Stevens M, Troscianko J, Jokela J (2020) The size, symmetry, and color saturation of a male guppy's ornaments forecast his resistance to parasites. Am. Nat. 196:597-608.

Zorrozua N, Alonso-Álvarez C, Diaz B, Sanpera C, Jover L, Arizaga J (2020) Carbon δ13C isotopic marker values correlate with carotenoid-based bill colouration in adult yellow-legged gulss Larus michahellis. Ardeola 67:325-339, 15.



 Paukner A, Slonecker EM, Wooddell LJ (2021) Effects of dominance and female presence on secondary sexual characteristics in male tufted capuchin monkeys (Sapajus apella). Ecol. Evol. n/a:xxx-xxx.

Peneaux C, Hansbro PM, Griffin AS (2021) The potential utility of carotenoid‐based coloration as biomonitor of environmental change. Ibis 163:20-37.

Powers MJ, Hill GE (2021) A review and assessment of the Shared-Pathway Hypothesis for the maintenance of signal honesty in red ketocarotenoid-based coloration. Integr. Comp. Biol. XX:xxx-xxx.

Renthlei Z, Hmar L, Trivedi KA (2021) High temperature attenuates testicular responses in tree sparrow (Passer montanus). Gen. Comp. Endocrinol. 301:113654.


File with all references here (carotenoids.ris)


 1994-2004          2005-2009          2010-2012          2013-2017     2018->

As the fact that carotenoids have immunostimulant properties became more widely known among ecologist, many other interesting "antioxidants and immunity in ecology" papers began to appear (e.g., transfer of antioxidants to the egg, antioxidant-dependent signalling by nestlings to parents, carotenoid- vs. melanin-dependent sexual signals, etc).

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