Shawkey, M.D., M.E. Hauber, L.K. Estep and G.E. Hill. 2006. Evolutionary  transitions and structural mechanisms of avian plumage coloration in grackles and allies (Icteridae). Journal of the Royal Society Interface 3:777-783. pdf

Shawkey, M.D. and G.E. Hill. 2006. Significance of a basal melanin layer to production of non-iridescent structural plumage colour: evidence from an amelanotic Stellers' Jay (Cyanocitta stelleri).  Journal of Experimental Biology 209:1245-1250. pdf

Shawkey, M.D., S.L. Balenger, G.E. Hill, L.S. Johnson, A.J. Keyser and L. Siefferman. 2006. Mechanisms of evolutionary change in structural plumage coloration among bluebirds (Sialia spp.). Journal of the Royal Society Interface 3:527-532 (cover article). pdf

Doucet, S.M., M.D. Shawkey, G.E. Hill and R. Montgomerie. 2006.  Iridescent color in satin bowerbirds: Structure, mechanisms and individual variation. Journal of Experimental Biology 209:380-390 (cover article). pdf

Shawkey, M.D. and G.E. Hill. 2005. Carotenoids need structural colours to shine. Biology Letters 1:121-124. pdf

Shawkey, M.D., K.L. Mills, C. Dale and G.E. Hill. 2005. Microbial diversity of wild bird feathers revealed through culture-based and culture-independent techniques. Microbial Ecology 50:40-47. pdf

Shawkey, M.D., A.M. Estes, L.M. Siefferman and G.E. Hill. 2005. The anatomical basis of sexual dichromatism in non-iridescent ultraviolet-blue structural coloration of feathers. Biological Journal of the Linnean Society 85:259-271. pdf

Shawkey, M.D. and G.E. Hill. 2004. Feathers at a fine scale. Auk. 141:652-655. pdf

Doucet, S.M., M.D. Shawkey, M.K. Rathburn, H.L. Mays, Jr. and R. Montgomerie. 2004. Concordant evolution of plumage colour, feather microstructure, and a melanocortin receptor gene between mainland and island populations of a fairy-wren. Proceedings of the Royal Society of London B 271:1663-1670. pdf

Shawkey, M.D., R. Bowman and G.E. Woolfenden. 2004. Why is partial brood loss among Florida Scrub-Jays higher in suburban than wildland habitat? Canadian Journal of Zoology 82:1427-1435. pdf

Shawkey, M.D., A.M. Estes, L.M. Siefferman and G.E. Hill. 2003. Nanostructure predicts intraspecific variation in ultraviolet-blue plumage colour.  Proceedings of the Royal Society of London B 270:1455-60. Commentary on the above article appears in Nature 423:820. pdf

Shawkey, M.D., S.R. Pillai and G.E. Hill. 2003.  Chemical warfare? Effects of uropygial oil on feather-degrading bacteria. Journal of Avian Biology 34:345-349. pdf

Shawkey, M.D., M.L. Beck and G.E. Hill. 2003.  Use of a gel documentation system to measure feather growth bars.  Journal of Field Ornithology 74:125-128. pdf

Ellis, D.E., B. Clauss, T. Watanabe, R.C. Mykut, M.D. Shawkey, D.P. Mummert, D.T. Sprague, C.H. Ellis and F.B.  Trahan.  2001. Results of the second (1996) experiment to lead cranes on migration behind a motorized ground vehicle.  Pp. 122-127 in Proceedings of the Eighth North American Crane Workshop.  International Crane Foundation, Baraboo, WI.

Shawkey, M.D., V. Saranathan, H. Palsdottir, J.C. Crum, M.L. Auer and R.O. Prum. 2009. Electron tomography, 3D Fourier analysis and color prediction of a 3D bio-photonic nanostructure. Journal of the Royal Society Interface. vol. 6  Suppl 2: S213–S220 pdf

Shawkey, M.D., N.I. Morehouse and P.R. Vuksuic. 2009. A protean palette: Color materials and mixing in birds and butterflies. Journal of the Royal Society Interface vol. 6  Suppl 2:  S221-S231 pdf

Shawkey, M.D., S. Pillai and G.E. Hill. 2009. Do feather-degrading bacteria affect sexually selected plumage color? Naturwissenschaften 96:123-128. pdf

Johnson, V. M., C. Guyer, M.D. Shawkey, and S. R. Roberts. 2008. Abundance, identification, and prospective participation of bacteria on gopher tortoise shell degradation. Journal of Alabama Academy of Science 79:190 -199. pdf

Shawkey, M.D., M. Liu, E. Loos, F. Rowher, J.M. Wang, and S.R. Beissinger. 2008. Do birds differentially deposit anti-microbial proteins in clutches of eggs?  Behavioral Ecology 19:920-927. pdf

Siefferman L., M.D. Shawkey, R. Bowman and G.E. Woolfenden. 2008  Juvenile coloration of Florida scrub-jays (Aphelocoma coerulescens) is sexually dichromatic and correlated with condition. Journal of Ornithology 149:357-363. pdf

Shawkey, M.D., S.R. Pillai, G.E. Hill, L.M. Siefferman and S.R. Roberts. 2007. Bacteria as an agent for change in structural plumage color: correlational and experimental evidence. The American Naturalist S112-117. pdf

Estep, L.K., M.D. Shawkey and G.E. Hill. 2006. Carotenoid-based breast plumage  colour, body condition and clutch size in red fodies (Foudia madagascariensis).  Ostrich 77:164-169. pdf

Shawkey, M.D., J. Hussain, J.L. Hagelin, A.C. Vollmer, and G.E. Hill. 2006. Culture-independent characterization of bacteria on feathers of Crested and Least Auklets (Aethia cristatella and Aethia pusilla). Waterbirds 29:507-510. pdf

Shawkey, M.D., G.E. Hill,  K.J. McGraw, W.R. Hood and K.L. Huggins. 2006. An experimental test of the relative contribution and condition-dependence of microstructure and carotenoids in yellow plumage coloration. Proceedings of the Royal Society of London B 273:2985-2991. pdf

  1. L.D’Alba, R. Maia, M.E. Hauber and M. D. SHAWKEY. In press. Evolution of eggshell cuticle in relation to nesting ecology. Proceedings of the Royal Society of London B.

M. Xiao, Y. Li, M. Allen, D. Deheyn, X. Yue, J, Zhao, N. Gianneschi, and A.

Dhinojwala, and M.D. SHAWKEY. In press. Stimuli-Responsive Structurally Colored Films from Bioinspired Synthetic Melanin Nanoparticles. Chemistry of Materials.

B. Igic, L. D’Alba, and M.D. SHAWKEY. 2016. Manakins produce bright iridescent feather

colors without melanosomes. Journal of Experimental Biology 219: 1851-1859. pdf

J-P Iskander, C.M. Eliason, R. Maia, and M.D. SHAWKEY. 2016. A nanostructural basis for

glossy red plumage colors. Biological Journal of the Linnaean Society. DOI: 10.1111/bij.12810 pdf

R. Maia, D. R. Rubenstein, and M.D. SHAWKEY. 2016. Selection, constraint and the

evolution of coloration in African starlings. Evolution 70:1064-1069. pdf

C.M. Eliason, M.D. SHAWKEY, and J.A. Clarke. 2016. Evolutionary shifts in the melanin-

based color systems of birds. Evolution 70:445-455. pdf

B. Hsiung, D. Deheyn, M.D. SHAWKEY, and T.A. Blackledge. 2015. Blue color in

tarantulas is evolutionarily conserved despite nanostructural diversity. Science Advances 1: e1500709

B. Hsiung, T.A. Blackledge, and M.D. SHAWKEY. 2015. Spiders do have melanin after all.

Journal of Experimental Biology 218:3632-3635. pdf

B. Igic, E. Zarate, M. A. Sewell, C. Moskát, P. Cassey, J. Rutila, T. Grim, M.D. SHAWKEY, and M.

E. Hauber. 2015. A comparison of egg yolk lipid components between parasitic

Common Cuckoos and their hosts. The Auk: Ornithological Advances 132:817-825. (cover article) pdf

M.E. Hauber, Z. Aidala, B.Igic, M.D. SHAWKEY, and C. Moskát. 2015. Orange is not the

new green: Experimental shifts in nest-egg contrasts do not predict egg rejection

decisions by great reed warbler hosts of brood parasitic common cuckoos. Animal Cognition DOI 10.1007/s10071-015-0886-9. pdf

M. Xiao, Y. Li, M. Allen, D. Deheyn, X. Yue, J, Zhao, N. Gianneschi, M.D. SHAWKEY, and A.

Dhinojwala. 2015. Bio-inspired structural colors produced via self-assembly of

synthetic melanin nanoparticles. ACS Nano 9:5454-5460. (Editor’s choice)

B. Igic, V.Nunez, H.U. Voss, R. Croston, Z. Aidala, A. Lopez, A.Van Tatenhove, M. Holford, M.D.

SHAWKEY, and M.E. Hauber. 2015. Examining egg-rejection behavior of wild

birds using 3D printed eggs. PeerJ 3:e965

D. C. Fecheyr-Lippens, B. Igic, L. D'Alba, D. Hanley, A. Verdes, M. Holford, G.I.N. Waterhouse,

T. Grim, M.E. Hauber, and M.D. SHAWKEY. 2015. The cuticle modulates ultraviolet reflectance of avian eggshells. Biology Open doi:10.1242/bio.01221.

L. D’Alba and M.D. SHAWKEY. 2015. Antimicrobial defenses in avian eggs: a review. J.

Ornithology. DOI:0.1007/s10336-015-1226-1 pdf

C.M. Eliason, R. Maia, and M.D. SHAWKEY. 2015. Modular color evolution in birds

facilitated by a complex nanostructure. Evolution 69: 357-367. pdf

B. Igic, Daphne Fecheyr-Lippens, A. Chan, D. Hanley, P. Brennan, T. Grim, G.I.N. Waterhouse,

M.E. Hauber, and M.D. SHAWKEY. 2015. A nanostructural basis for gloss of avian eggshells.  Journal of the Royal Society Interface DOI: 10.1098/rsif.2014.1210 pdf

M.D. SHAWKEY, L. D’Alba, M. Xiao, M. Schutte and R. Buchholz. 2015. Ontogeny of

an iridescent nanostructure composed of hollow melanosomes. Journal of Morphology. doi: 10.1002/jmor.20347 pdf

B. Hsiung, T.A. Blackledge and M.D. SHAWKEY. 2014. Structural color and its interactions

with other color-producing elements: perspectives from spiders. Proceedings of SPIE 9187. pdf

S. Liu, M.D. SHAWKEY, D. Parkinson and M. Ahmed. 2014. Elucidation of the chemical

composition of avian melanin. Royal Society of Chemistry Advances 4:40396-99. pdf

M.E. Hauber, L.Tong, M. Bán, R. Croston, T. Grim, G.I.N. Waterhouse, M.D. SHAWKEY, and C.

Moskát. 2015. The value of artificial stimuli in behavioral ecology research:

Making the case for egg rejection studies in avian brood parasitism. Ethology 121:521-528 pdf

L. D’Alba, C. van Hemert, K.A. Spencer, B.J. Heidinger, L. Gill, N.P. Evans, P. Monaghan, C.M.

Handel, and M.D. SHAWKEY. 2014. Melanin-based color of plumage: Role of condition

and of feathers’ microstructure. Integrative and Comparative Biology doi: 10.1093/icb/icu094 pdf

M. Xiao, Ali Dhinojwala and M.D. SHAWKEY. 2014. Nanostructural basis of rainbow-like

iridescence in common bronzewing Phaps chalcoptera feathers. Optics Express 22:14625-


C. M. Eliason and M.D. SHAWKEY. 2014. Antireflection-enhanced color

by a natural graded refractive index (GRIN) structure. Optics Express 22:A642-A650.

J. Prokop, C. Schmidt, D. Gasper, R.J. Duff, A. Milsted, T. Ohkubo, H.C. Ball, M.D. SHAWKEY,

H.L. Mays, L.A. Cogburn, R.L. Londraville. 2014. Discovery of the elusive leptin in

birds: identification of several “missing links” in the evolution of leptin and its receptor. PLoS ONE 9:e92751

L. D’Alba, C.M. Eliason, H. Badawy, D.N. Jones and M.D. SHAWKEY. 2014. Antimicrobial

effects of a nanostructured eggshell in a compost-nesting bird. Journal of Experimental

Biology 217:1116-1121. pdf

Q. Li, J.A. Clarke, K. Gao, C. Zhou, Q. Meng, D. Li, L. D’Alba and M.D. SHAWKEY. 2014.

Melanosome evolution indicates a key physiological shift within feathered dinosaurs.

Nature. pdf

J. Lindgren, P. Sjovall, J.A. Gren, P. Udval, G. Dyke, R.M. Carney, B.P. Schultz, M.D. SHAWKEY,

K.R. Barnes, and M.J. Polcyn. 2014. Skin pigmentation provides evidence of

convergent melanism in extinct marine reptiles. Nature. pdf

C.M. Eliason, P.P. Bitton, M.D. SHAWKEY 2013. How hollow melanosomes affect iridescent plumage color in birds. Proc. Roy. Soc. Lond. B pdf

R. Maia, D.R. Rubenstein and M.D. SHAWKEY. 2013. Key ornamental innovations facilitate diversification in an avian radiation. PNAS pdf

R. Maia, C.M. Eliason, P.P. Bitton, S.M. Doucet, and M.D. SHAWKEY. 2013. pavo: an R package for the analysis, visualization and organization of spectral data. Methods in Ecology and Evolution doi: 10.1111/2041-210X.12069  pavo on CRAN pdf

D. Field, L. D’Alba, J. Vinther and M.D. SHAWKEY. 2013. Melanin concentration gradients in extant and fossil feathers. PLoS One 8(3):e59451 pdf

Eliason C. M. and M.D. SHAWKEY. 2012. A 2D biophotonic heterostructure produces diverse iridescent colors in dabbling ducks. Journal of the Royal Society Interface doi:

10.1098/ rsif.2012.0118. Coverage in Science pdf


Q. Li, K. Gao, Q. Meng, J.A. Clarke, M.D. SHAWKEY*, L. D’Alba, R. Pei, M. Ellison, M.A. Norell, J. A. Vinther. 2012. A new specimen of Microraptor and the evolution of iridescent plumage color. Science. pdf     Media coverage

*corresponding author


D’Alba,L., Briggs L. and M.D. SHAWKEY. 2012. Relative contributions of pigments and biophotonic nanostructures to natural color production: a case study in Budgerigars (Melopsittacus undulatus). Journal of Experimental Biology 215:1272-1277.pdf


H. K. Snyder, R. Maia, L. D’Alba, A. J. Schulz, K. Rowe, S. Rowe, and M.D. SHAWKEY. 2012.

Iridescent colour production in hairs of blind golden moles (Chrysochloridae). Biology Letters. doi: 10.1098/rsbl.2011.1168. pdf

R. Carney, J. Vinther, M.D. SHAWKEY, L. D’Alba, and J. Ackermann. 2012. New evidence on the color, ultrastructure and nature of the isolated Archaeopteryx feather. Nature Communications 3, 637. pdf

D’Alba, L.,  Van Hemert, C. Handel C.M. and M.D. SHAWKEY. 2011. A natural experiment on the condition-dependence of achromatic plumage coloration in black-capped chickadees. PLoS One 6: e25877. pdf


Maia, R., Macedo R.H. and M.D. SHAWKEY. 2011. Self-assembly of iridescent barbules through depletion attraction of melanosomes during keratinization. Journal of the Royal Society Interface: doi: 10.1098/ rsif.2011.0456.

Shawkey M.D., Maia R., D’Alba L. 2011. Proximate bases of silver color in Anhinga (Anhinga anhinga) feathers. Journal of Morphology. DOI: 10.1002/jmor.10993

Eliason C.M., Shawkey M.D. 2011. Decreased hydrophobicity of iridescent feathers: a potential cost of shiny plumage. Journal of Experimental Biology. 214:2157-63.

D’Alba L., Saranathan V. Clarke, J.A. Vinther, J.A. Prum, R.O. Shawkey, M.D. 2011. Colour producing β-keratin nanofibres in blue penguin feathers. Biology Letters. DOI: 10.1098/rsbl.2010.1163

Shawkey M.D., D’Alba L., Wozny J., Eliason C., Koop J. and Jia L. 2011. Structural color change following hydration and dehydration of iridescent mourning dove (Zenaida macroura) feathers. Zoology. DOI:10.1016/j.zool.2010.11.001

Maia R., D’Alba L., Shawkey M.D. 2011. What makes a feather shine? A nanostructural basis for glossy black colors in feathers. Proceedings of the Royal Society B. 278: 1973-1980.pdf

D’Alba, L., Oborn A., Shawkey M.D. 2010 Experimental evidence that keeping eggs dry is a mechanism for the antimicrobial effects of avian incubation. Naturwissenschaften.


Eliason, C.M., Shawkey M.D. 2010. Rapid, reversible response of iridescent feather color to ambient humidity. Opt. Express 18, 21284-21292. DOI:10.1364/OE.18.021284

Clarke J.A., Ksepka D., Salas-Gismondi R., Altamirano A.J., Shawkey M.D., D’Alba L., Vinther J., DeVries J.  Baby P. 2010. Fossil evidence for evolution of the shape and color of penguin feathers. Science. DOI: 10.1126/science.1193604

D’Alba L., Shawkey M.D., Korsten P., Komdeur J. Beissinger S.R. et al. 2010. Differential deposition of antimicrobial proteins in blue tit (Cyanistes caeruleus) clutches by laying order and male attractiveness. Behavioural Ecology and Sociobiology 64:1037-1045 pdf


Li, Q., Keqin, G., Vinther, J., Shawkey, M.D., Clarke J.A., D’Alba L., Meng, Q., Briggs D.E.G., Prum, R.O. 2010 Plumage color patterns of an extinct non-avian dinosaur. Science. DOI: 10.1126/science.1186290                                   

Shawkey, M.D., E.L. Brodie, M.K. Firestone and S.R. Beissinger. 2009. Avian incubation inhibits bacterial growth and diversification on eggs. PLoS ONE 4(2): e4522. PDF


The Shawkey Lab