Dr.
Kevin McGraw
Associate
professor
Arizona
State University
School
of Life Sciences
Tempe,
AZ 85287-4501
Click here to learn about
opportunities as a graduate student in my lab
Contents
Contact
Information
Electronic mail
address: Kevin.McGraw@asu.edu
Office phone: (480)
965-5518
Lab phone: (480)
965-2593
Departmental fax:
(480) 965-6899
Education
|
Institution |
Degree |
Dates |
Department |
Advisor |
|
Post-doc Ph.D. M.S. B.S. |
1/04-8/04 8/99-12/03 9/97-8/99 8/93-5/97 |
My research
programme centers on the control and function of animal colors, particularly in
birds. The Darwinian theory of
sexual selection coined over 125 years ago remains the cornerstone of
biological explanations for the evolution of exaggerated ornamental traits. Brilliant colors, for example, 
evolve particularly in male animals in response to intense competitions
for mates. Only the most colorful
males can out-compete their rivals and serve as suitable, attractive mates for
females.
While there is now
much empirical evidence to support this basic tenet of sexual-selection theory
for many ornament types (e.g. large body-size, colorful plumage, elongate
tails), biologists have become particularly interested in why females pay particular attention to these male traits
when making their mating decisions.
What information are males communicating to females with their
color? What advantages do females
gain by mating with the most colorful male? It is with a mechanistic approach to studying the particular
factors that mold the development of colorful plumage in birds that we have
begun to understand the costs to males in developing bright colors and the
benefits accrued by females who acquire the sexiest mate.
My advisor during
my M.Sc. degree, Dr. Geoffrey Hill, helped revolutionize many of the ways in
which biologists perceive ornamental traits as mating signals. Following the theoretical framework
offered by Zahavi, Williams, Fisher, and of course Darwin, he pursued the means
by which males communicate their worth to females in a colorful species of
songbird from 
North America, the house finch (Carpodacus mexicanus). Males
derive their colorful plumage, which ranges from bright red to drab yellow,
from carotenoid pigments, the same pigments that are responsible for the yellow
color of corn, the orange color of carrots, and the redness of autumn
leaves. Only photosynthetic
organisms like plants and algae can manufacture these molecules, so birds and
other vertebrates can only display carotenoids in feathers by first acquiring
them in their diet. In a series of
elegant field and laboratory experiments, Geoff demonstrated that the more
brightly colored males are preferred as mates by females, but also obtain the
largest quantities of carotenoids in their diet, are in the best nutritional
condition, help provision females with food as she prepares for breeding, and
have the highest survival rates.
Thus, males use their color to demonstrate to females their foraging
abilities, general health, and willingness to provide parental assistance. Only with this foundation of research
in place have we begun to explore the ways in which carotenoid-derived plumage
is controlled and functions in other songbirds as well as the information
contained within other types of color signals in animals.
In fact, these two
areas of research have been the focal points of my dissertation work at Cornell
University. To dive deeper into
the world of carotenoid-signaling in birds is to understand these pigments at
the biochemical and physiological levels.
Animals certainly acquire these molecules in the diet, but also use a
complex physiological processing 
system to transport carotenoids through the body to the site of
deposition at the growing feather.
Thus, we must consider these physiological steps as potentially costly
components of pigment utilization and thus a direct means by which male birds
can communicate their nutritional, physiological, and genetic quality to
females. Preliminary work has in
fact shown that certain male songbirds (e.g. American goldfinches) develop
differently colored plumage when fed the same diet (McGraw and Hill 2001),
suggesting a role for physiological regulation of color development. In collaboration with Dr. Bob Parker in
the Nutritional Sciences division at Cornell, we are using high-performance
liquid-chromatography (HPLC) as a biochemical tool for identifying the
carotenoid pigments acquired by these animals in their diet and the degree to
which they circulate them through the body bound to lipoprotein particles and
deliver them to peripheral tissues for pigmentation. Birds are also capable of metabolically altering the
structure of carotenoids, often times rendering them more desirable as feather
colorants. It appears that certain
metabolic pathways may be favored over others en route to the development of
the most brightly colored feathers (McGraw et al. 2002). There is no doubt that this new,
mechanistic approach to evaluating the production of color signals will offer
many unique insights into the costliness and signal content of carotenoid-based
colors in birds.
Carotenoid pigments
are by no means the only source of color in animals, however. Melanin pigments are responsible for
most of the black, gray, brown, and earth-toned colors in nature (e.g. in
insect cuticles, animal fur, lizard scales, and 
bird feathers), and are found in two primary forms, eumelanin (blacks
and grays) and phaeomelanin (rust, olive). Many birds display sexually dimorphic patches of black color
in their feathers, and this has been the focus of a great body of literature
over the past 25 years. These
eumelanic colors appear to serve unique functions as signals of social status
in populations of songbirds, often times in non-breeding contexts (e.g. in a
winter foraging flock).
Again, to understand why these ornaments reliably reveal the aggressive
intentions or dominance ability of individuals is to delve into the mechanisms
underlying the production and maintenance of these signals. I have accumulated evidence that
nutritional input has little impact on the development of eumelanic plumage
colors in certain species (e.g. American goldfinch, house sparrow; see McGraw
and Hill 2000, McGraw et al. 2002).
Instead, it seems that physiological regulation occurs via
steroid-hormone circulation (e.g. testosterone) and its corresponding link to
aggressive behavior in animals. We
are currently considering the role that social factors themselves play on the
androgenic responsiveness of individuals and the associated development of
melanin-based plumage.
Lastly, an
altogether different form of color has captured the attention of behavioral
ecologists over the past few years.
Rather than having a pigmentary origin, structurally colored feathers
have unique microstructural arrangements of feather tissues and layers that
often reflect iridescent, blue, or ultraviolet wavelengths of light. Human-vision is limited for short
light-wavelengths, and as such many of the structural plumage colors in 
songbirds have been historically ignored as signals of
mate quality. Recent advances in
spectrophotometric technology, however, have allowed more objective and
quantitative means of assessing structural-color. In fact, many songbirds
exhibit sexually dichromatic patterns of structural color that function as
targets of female mate-choice.
Much less is known of the proximate factors that shape the production of
these carefully arranged feather tissues, and this has been the subject of a
series of studies in collaboration with fellow graduate students in and around
our department here at Cornell (McGraw et al., 2002).
In all, my
biological research is deeply rooted in integrative approaches, as the only
means by which we can understand how and why animals communicate with the
signals they do and can provide unique sets of information with the diversity
of ornaments that exist. Continued
progress at the biochemical, physiological, molecular, and genetic levels will
bring us that much closer to identifying the true fitness benefits that females
reap from securing a colorful mate, in currencies that not even Darwin could
have imagined.
Edited books
Hill, G. E. and K. J. McGraw. 2006b.
Bird Coloration. Volume II. Function and Evolution. Harvard University
Press, Cambridge, MA. Catalog
advertisement, Table of
Contents, Purchase
now, Read
an excerpt, Wingspan (review), Times
Literary Supplement (review), Science
(browsing), Trends in
Ecology and Evolution (review), Nature
(review), Condor
(review), Quarterly
Review of Biology (review), ISBE
newsletter (review)
Hill, G. E. and K. J. McGraw. 2006a. Bird Coloration. Volume I. Mechanisms and
Measurements. Harvard University Press, Cambridge, MA. Catalog
advertisement, Table of
Contents, Purchase
now, ASU
Insight feature (on last page), Condor
(review), Auk
(review)
Journal
Publications or Book Chapters
114. Safran, R. J., K. J. McGraw, M. R. Wilkins, J. Hubbard, and J. Marling. 2010.
Positive antioxidant balance over the breeding season predicts reproductive performance
in a wild bird. PLoS One (in press).
113. Nolan, P. M., F. S. Dobson, T. J. Karels, K.
J. McGraw, and P. Jouventin. 2010. Mutual
mate choice for colorful traits in king penguins. Ethology (in press).
112. Toomey, M. B., M. W. Butler, M. G. Meadows,
L. A. Taylor, H. Fokidis, and K. J. McGraw.
2010. A novel method for quantifying the glossiness of animals. Behav. Ecol.
Sociobiol. (in press).
111. Navarro Puig, C., T. Perez-Contreras, J.
Aviles, J. Soler, and K. J. McGraw.
2010. Beak colour reflects circulating carotenoid and vitamin A levels in
spotless starlings (Sturnus unicolor). Behav. Ecol. Sociobiol.
(in press).
110. McGraw, K. J. 2010. Avian antioxidants and oxidative stress: insights from food,
physiology, and feathers. In: Oxidative stress (Mandelker, L. and Vajdovich,
P., eds), Springer, NY.
109.
Hipfner, J. M., K. A. Hobson, J. Dale, and K. J. McGraw. 2010. Stable isotopes link diet to avian yolk
carotenoid allocation: a comparative study of five auk species
(Charadriiformes: Alcidae). Physiol. Biochem. Zool. (in press).
108. Lin, S. M., K. Nieves-Puigdoller, A. C.
Brown, K. J. McGraw, and E. D.
Clotfelter. 2010. Testing the carotenoid trade-off hypothesis in the
polychromatic Midas cichlid, Amphilophus citrinellus. Physiol.
Biochem. Zool. (in press). Uncorrected page
proofs
107. McGraw, K. J. and M. B. Toomey. 2010. Carotenoid accumulation in the tissues of
zebra finches: predictors of integumentary pigmentation and implications for
carotenoid allocation strategies. Physiol. Biochem. Zool. 83:97-109. PDF and appendix
106. Cohen, A. A., R. A. Mauck, N. T. Wheelwright,
C. E. Huntington, and K. J. McGraw.
2009. Complexity in relationships between antioxidants and individual
life-history parameters in a songbird and a seabird. Oikos 118:1854-1861. PDF
105. McGraw, K. J. and J. D. Blount. 2009. Control and function of carotenoid coloration
in birds: a review of case studies. In Carotenoids: Physical, Chemical,
and Biological Functions and Properties (J. T. Landrum, ed.). CRC Press, Boca
Raton, FL (in press; published 21 December 2009).
104. Butler, M. W. and K. J. McGraw. 2009. Indoor housing during development affects
moult, carotenoid circulation, and beak colouration of mallard ducks (Anas
platyrhynchos). Avian Biol. Res. 2:203-211. PDF
103. Dierenfeld, E.S., K. J. McGraw, K. Fritsche, J.T. Briggler, and J. Ettling. 2009.
Nutrient composition of whole crayfish (Orconectes and Procambarus species) consumed by hellbender (Cryptobranchus
alleganiensis). Herp. Rev. 40:324-330. PDF
102. Cohen, A. A., W. D. Robinson, and K. J.
McGraw. 2009. Serum antioxidant levels in
wild birds vary in relation to diet, season, life history strategy, and
species. Oecologia 161:673-683. PDF, Supplementary
Material 1, Supplementary
Material 2
101. Bascunan, A. L., E. A. Tourville, M. B.
Toomey, and K. J. McGraw. 2009. Food
color preferences of molting house finches (Carpodacus mexicanus)
in relation to sex and plumage coloration. Ethology 115:1066-1073. PDF
100. Steffen, J. and K. J. McGraw. 2009. How dewlap color reflects its carotenoid and
pterin content in brown anoles. Comp. Biochem. Physiol. B 154:334-340. PDF
99. McGraw, K. J. 2009. Identifying anatomical sites of carotenoid metabolism in birds.
Naturwissenschaften 96:987-988. PDF, Original article
critiqued, Authors’
response to my critique
98. McGraw, K. J., M. Massaro, T. J. Rivers, and T. Mattern. 2009. Annual, sexual,
size-, and condition-related variation in the colour and fluorescent pigment
content of yellow crests in Snares penguins (Eudyptes robustus).
Emu 109:93-99. PDF,
cover image
97. McGraw, K. J. 2009. Canine coloration: dark domesticated dogs and the bright side
for wild wolves. Pigment Cell Melanoma Res. 22:255-256. PDF
96. Meadows, M. G., M. W. Butler, N. I. Morehouse,
L. A. Taylor, M. B. Toomey, K. J. McGraw,
and R. L. Rutowski. 2009. Iridescence: views from many angles. J. R. Soc.
Interface 6:S107-S113. PDF
95. Cohen, A. A. and K. J. McGraw. 2009. No simple measures for antioxidant status in
birds: complexity in inter- and intraspecific correlations among circulating
antioxidant types. Funct. Ecol. 23:310-320. PDF
94. Toomey, M. B. and K. J. McGraw. 2009. Seasonal, sexual, and quality related
variation in retinal carotenoid accumulation in the house finch (Carpodacus
mexicanus). Funct. Ecol. 23:321-329. PDF
93. McGraw, K. J. and A. L. A. Middleton. 2009. American Goldfinch (Carduelis tristis).
The Birds of North America Online (A. Poole, Ed.). Ithaca, NY: Cornell
Laboratory of Ornithology.
92. Rowe, M. and K. J. McGraw. 2009. Carotenoids in the seminal fluid of wild
birds: interspecific variation in fairy-wrens. Condor 110:694-700. PDF
91. McGraw, K.
J. 2009. Visual signaling in animals.
In: Encyclopedia of Neuroscience (L. R. Squire, ed.), vol. 10. Academic Press
Oxford, pp. 307-311. PDF
90. McGraw, K. J., E. A. Tourville, and M. W. Butler. 2008. A quantitative comparison of
the commonly used methods for extracting carotenoids from avian plasma. Behav.
Ecol. Sociobiol. 62:1991-2002. PDF
89. Cohen, A. A., K. J. McGraw, P. Wiersma, J. B. Williams, W. D. Robinson, T. R.
Robinson, J. D. Brawn, and R. E. Ricklefs. 2008. Interspecific associations
between circulating antioxidant levels and life-history variation in birds. Am.
Nat. 172:178-193. PDF
88. Piault, R., J. Gasparini, P. Bize, M. Paulet, K.
J. McGraw, and A. Roulin. 2008.
Experimental support for the make-up hypothesis in nestling tawny owls (Strix
aluco). Behav. Ecol. 19:703-709. PDF
87. Safran, R. J., J. Adelman, K. J. McGraw, and M. Hau. 2008. Sexual signal elaboration
affects physiological state in a social vertebrate. Current Biology
18:R461-R462. PDF,
over
300 newspaper and online articles written on this paper, plus Good Morning America,
CNN,
and several local radio interviews (e.g. San Francisco, Houston, Rockford, IL,
New York). TREE
“Research Focus” article highlighting this paper
86.
Deviche, P., K. J. McGraw, and
J. Underwood. 2008. Age-, sex- and season-specific
accumulation of plasma carotenoid pigments in white-winged crossbills. J. Avian
Biol. 39:283-292. PDF
85. Blount, J. D. and K. J. McGraw. 2008. Signal functions of carotenoid colouration.
In: Carotenoids. Volume 4: Natural Functions (G. Britton, S. Liaaen-Jensen, and
H. Pfander, eds.). Birkhauser Verlag, basel, pp. 213-236. Uncorrected
galley proofs
84. McGraw, K. J. 2008. An update on the honesty of melanin-based color signals in birds.
Pigment Cell Melanoma Res. 21:133-138. PDF
83. Schaefer,
H. M., K. J. McGraw,
and C. Catoni. 2008. Birds use fruit colour as an honest signal of dietary
antioxidant rewards. Funct. Ecol. 22:303-310. PDF
82. Juola, F. A., K. J. McGraw, and D. C. Dearborn. 2008. Carotenoids and throat
pouch coloration in the great frigatebird (Fregata minor). Comp.
Biochem. Physiol. B. 149:370-377. PDF
81. Safran, R. J., Pilz, K. M., McGraw, K. J., Correa, S. M., and Schwabl, H. 2008. Are yolk
androgens and carotenoids in barn swallow eggs related to parental quality?
Behav. Ecol. Sociobiol. 62:427-438. PDF
80. Jouventin, P., K. J. McGraw, M. Morel, and A. Celerier. 2008. Dietary carotenoid supplementation affects orange
beak but not foot coloration in gentoo penguins. Waterbirds 30:573-578. PDF
79. Clotfelter,
E., Ardia, D., and McGraw, K. J.
2007. Red fish, blue fish: trade-offs between pigmentation and immunity in Betta
splendens. Behav. Ecol.
18:1139-1145. PDF
78. Toomey, M. B. and K. J. McGraw. 2007. Modified saponification and HPLC methods for
analyzing carotenoids from the retina of Japanese quail (Coturnix japonica):
implications for its use as a nonprimate model species. Invest. Ophthalmol.
Vis. Sci. 48:3976-3982. PDF
77. Taylor, L. A. and K. J. McGraw. 2007. Animal coloration: sexy spider scales. Curr.
Biol. 17:R592-R593. PDF
76. McGraw, K. J., M. B. Toomey, P. M. Nolan, N. I. Morehouse, M. Massaro, and P.
Jouventin. 2007. A description of unique fluorescent yellow pigments in penguin
feathers. Pigment Cell Res. 20:301-304. PDF, Supplementary
Material
75. Bezzerides, A. L., K. J. McGraw, R. S. Parker, and J. Husseini. 2007. Elytra color
as a signal of chemical defense in the Asian ladybird beetle (Harmonia
axyridis). Behav. Ecol. Sociobiol. 61:1401-1408. PDF
74. McGraw, K. J. and D. R. Ardia. 2007. Do carotenoids buffer testosterone-mediated
immunosuppression?: an experimental test in a colorful songbird. Biol. Letters
3:375-378. PDF, Electronic
Supplementary Material, Biology
Letters Highlighted Paper, ASU Headline,
ASU News,
School of Life
Sciences News, Science
Daily, Innovations
Report, Biology
News Net, PhysOrg.com,
Eureka
Alert, DentalPlans.com,
United
Press International, ImediNews, Earth Times,
Monsters
and Critics, New Kerala,
Post
Chronicle, News-Medical.net,
Associated
Content, Medical
News Today, MediLexicon,
Nutrition
Horizon, ASU
Research Magazine, Softpedia
73. Hofmann, C. M., K. J. McGraw, T. W. Cronin, and K. E. Omland. 2007. Melanin
coloration in New World orioles I: carotenoid masking and pigment dichromatism
in the orchard oriole complex. J. Avian Biol. 38:163-171. PDF
72. McGraw, K. J., W. Medina-Jerez, and H. Adams. 2007. Carotenoid-based plumage coloration and aggression
during molt in male house finches. Behaviour 144:165-178. PDF
71. Steffen, J. and K. J. McGraw. 2007. Patterns of carotenoid and pterin
pigmentation in the colorful dewlaps of two anole species. Comp. Biochem.
Physiol. B. 146:42-46. PDF
70. McGraw, K. J. 2007. Dietary mineral content influences melanin-based ornamental
coloration. Behav. Ecol. 18:137-142. PDF
69. McGraw, K. J. 2006. The mechanics of
uncommon colors in birds: pterins, porphyrins, and psittacofulvins. In: Bird
Coloration. I. Mechanisms and Measurements (G. E. Hill and K. J. McGraw, eds.).
Harvard University Press, Cambridge,
MA, pp. 354-398.
68. McGraw, K. J. 2006. The mechanics of carotenoid coloration in birds. In: Bird
Coloration. I. Mechanisms and Measurements (G. E. Hill and K. J. McGraw, eds.).
Harvard University Press, Cambridge, MA, pp. 177-242.
67. McGraw, K. J. 2006. The mechanics
of melanin coloration in birds. In: Bird Coloration. I. Mechanisms and
Measurements (G. E. Hill and K. J. McGraw, eds.). Harvard University Press,
Cambridge, MA, pp. 243-294.
66. Shawkey, M. D., G. E. Hill, K. J. McGraw, and W. R. Hood. 2006. An experimental test of the
contributions and condition-dependence of microstructure and carotenoids in
yellow plumage colouration. Proc. R. Soc. Lond. B. 273:2985-2991. PDF
65. McGraw, K. J., O. L. Crino, W. Medina-Jerez, and P. M. Nolan. 2006. Effect of
dietary carotenoid supplementation on food intake and immune function in a
songbird with no carotenoid coloration. Ethology 112: 1209-1216. PDF
64. McGraw, K. J. and K. C. Klasing. 2006. Carotenoids,
immunity, and integumentary coloration in red junglefowl (Gallus gallus).
Auk 123:1161-1171. PDF
63. McGraw, K. J. 2006. Carotenoids mediate a trade-off between egg quantity and quality
in Japanese quail. Ethol. Ecol. Evol. 18:247-256. PDF
62. McGraw, K. J. 2006. Sex-steroid dependence of carotenoid-based coloration in female
zebra finches. Physiol. Behav. 88:347-352. PDF
61. McGraw, K. J., P. M. Nolan, and O. L. Crino. 2006. Carotenoid accumulation
strategies for becoming a colorful house finch: analyses of plasma and liver
pigments in wild molting birds. Funct. Ecol. 20:678-688. PDF, Cover Image
60. McGraw, K. J., S. M. Correa, and E.
Adkins-Regan. 2006. Testosterone upregulates lipoprotein status to control
sexual attractiveness in a colorful songbird. Behav. Ecol. Sociobiol.
60:117-122. PDF
59. McGraw, K. J. and L. Hardy. 2006. Astaxanthin is responsible for the pink plumage
flush of Franklins and Ring-billed gulls. J. Field Ornithol. 77:29-33. PDF
58. McGraw, K. J. and R. S. Parker.
2006. A novel lipoprotein-mediated mechanism controlling sexual attractiveness
in a colorful songbird. Physiol. Behav. 87:103-108. PDF
57. Deviche, P., K. McGraw, and E. Greiner. 2005. Interspecific differences in
hematozoan infection in Sonoran Desert Aimophila sparrows. J.
Wild. Dis. 41:532-541. PDF
56. McGraw, K. J., E. Adkins-Regan, and R. S. Parker. 2005. Maternally derived
carotenoid pigments affect offspring survival, sex ratio, and sexual
attractiveness in a colorful songbird. Naturwiss. 92:375-380. PDF
55. McGraw, K.
J., R. J. Safran, and K.
Wakamatsu. 2005. How feather colour reflects its melanin content. Funct. Ecol.
19:816-821. PDF
54. Safran, R. J., C. R. Neuman, K. J. McGraw, and I. J. Lovette. 2005. Dynamic paternity
allocation as a function of male plumage color in barn swallows. Science
309:2210-2212. PDF, Cover image, Cornell
University, Arizona State University, ASU Insight, ASU
homepage, BBC, Cornell
Daily Sun, Cornell Lab of Ornithology, American Association for the Advancement of Science, ABC News, ABC News
Online, AOL News, Belleville News Democrat, Biology News Net, Cherry Bomb, CLAS Alumni Magazine, EurekAlert, East Valley
Tribune (AZ), East Valley Tribune homepage,
Melbourne Herald Sun, Innovations Report, Independent Online, Live
Science, MSNBC, National Geographic, National Wildlife
Federation Magazine, New Haven Herald,
New Kerala, New
Scientist, New York Times, Newswise, Noorderlicht,
NPR, OptusNet, Pet
Talk Radio, Planet Ark, Red Nova, Reuters, Salem Statesman Journal, Science Daily, School
of Life Sciences (ASU), Science News, Telegraph, The Age,
The Oregonian, Times Online, Unison,
Virtual
Life Sciences Library, Web India, World News Australia, Yahoo News, AAAS radio
clip, CLAS
News (ASU), 10,000 Birds
53. McGraw, K. J. 2005. Interspecific variation in dietary carotenoid assimilation in
birds: links to phylogeny and color ornamentation. Comp. Biochem. Physiol. B
142:245-250. PDF
52. McGraw, K. J. and M. C. Nogare. 2005. Distribution of unique red feather pigments in
parrots. Biology Letters 1:38-43. PDF,
Arizona State University homepage 1, Arizona State University homepage 2, Arizona State University press release 1, Arizona State University press release 2, Arizona State University video interview, ASU Insight Newspaper, The Arizona Republic, Innovations Report, The Biochemist, Windy City Parrot, Wissenschaft, EurekaAlert
(AAAS), ChemLin, ParrotSuperCenter, Yahoo Korea, Noorderlicht,
ScienceDaily, LiveScience, ParrotScience.com,
The Oregonian, Farbimpulse, Futura
Sciences, El Correo Digital, Vigyan Prasar, Precious
Fids, Girl Scientist, Science News, Aviary
& Cage Bird Society of South Florida, ASU Insight (Summer Highlights), ASU Alumni Magazine (Spring 2005), eNewsSource, BBC
Wildlife, BirdTalk Magazine, Grade Winner, Planned Parrothood, Chemical and Engineering News, Birdwatchers Digest
51. McGraw, K. J. 2005. The antioxidant function of many animal pigments: are there
consistent health benefits of sexually selected colorants? Anim. Behav.
69:757-764. PDF, Top 10 article in
Animal Behaviour
50. 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
49. McGraw, K. J., G. E. Hill, and R. S.
Parker. 2005. The physiological costs of being colourful: nutritional control
of carotenoid utilization in the American goldfinch (Carduelis tristis).
Anim. Behav. 69:653-660. PDF
48. McGraw, K. J., J. Hudon, G. E.
Hill, and R. S. Parker. 2005. A simple and inexpensive chemical test for
behavioral ecologists to determine the presence of carotenoid pigments in
animal tissues. Behav. Ecol. Sociobiol. 57:391-397. PDF
47. McGraw, K. J. 2004. Not all red, orange, and yellow animal colors are
carotenoid-based: the need to couple biochemical and behavioral studies of
color signals. Proc. Ind. Natl. Sci. Acad. B 70:593-598. PDF
46. McGraw, K. J., K. Wakamatsu, A. B. Clark, and K. Yasukawa. 2004. Red-winged blackbirds
Agelaius phoeniceus use carotenoid and melanin pigments to color
their epaulets. J. Avian Biol. 35:543-550. PDF
45. McGraw, K. J. 2004. Colorful songbirds
metabolize carotenoids at the integument. J. Avian Biol. 35:471-476. PDF
44. McGraw, K. J. and A. J. Gregory. 2004. Carotenoid pigments in male American
goldfinches: what is the optimal biochemical strategy for becoming colourful?
Biol. J. Linn. Soc. 83:273-280. PDF
43. McGraw, K. J. and J. G. Schuetz. 2004. The evolution of carotenoid coloration in
estrildid finches: a biochemical analysis. Comp. Biochem. Physiol. B 139:45-51.
PDF
42. McGraw, K. J. 2004. Winter plumage coloration in male
American goldfinches: do reduced ornaments serve signaling functions in the
non-breeding season? Ethology 110:707-715. PDF
41. McGraw, K. J., G. E. Hill, K. J. Navara, and R. S. Parker. 2004. Differential
accumulation and pigmenting ability of dietary carotenoids in colorful finches.
Physiol. Biochem. Zool. 77:484-491. PDF
40. McGraw, K. J. and K. Wakamatsu. 2004. Melanin basis of ornamental feather colors in
male zebra finches. Condor 106:686-690. PDF
39. Horak, P., L. Saks, U. Karu, I. Ots, P. F.
Surai, and K. J. McGraw. 2004. How
coccidian parasites affect health and appearance of greenfinches. J. Anim.
Ecol. 73:935-947. PDF
38. McGraw, K. J., R. J. Safran, M. R. Evans, and K. Wakamatsu. 2004. European barn
swallows use melanin pigments to color their feathers brown. Behav. Ecol.
15:889-891. PDF, Confirmatory reply by Stradi
37. McGraw, K. J. and G. E. Hill. 2004. Plumage color as a dynamic trait: carotenoid
pigmentation of male house finches (Carpodacus mexicanus) fades
during the breeding season. Can. J. Zool. 82:734-738. PDF
36. McGraw, K. J. and M. C. Nogare. 2004. Carotenoid pigments and the selectivity of
psittacofulvin-based coloration systems in parrots. Comp. Biochem. Physiol. B
138:229-233. PDF
35. Safran, R. J. and K. J. McGraw. 2004. Plumage
coloration, not length or symmetry of tail-streamers, is a sexually selected
trait in North American barn swallows. Behav. Ecol. 15:455-461. PDF, Nature
Australia
34. McGraw, K. J., K. Wakamatsu, S. Ito, P. M. Nolan, P. Jouventin, F. S.
Dobson, R. E. Austic, R. J. Safran, L. M. Siefferman, G. E. Hill, and R. S.
Parker. 2004. You cant judge a pigment by its color: carotenoid and melanin
content of yellow and brown feathers in swallows, bluebirds, penguins, and
domestic chickens. Condor 106:390-395. PDF
33. McGraw, K. J. 2004. Multiple UV reflectance peaks in the iridescent neck feathers of
pigeons. Naturwiss. 91:125-129. PDF
32. Mays Jr., H. L., K. J. McGraw,
G. Ritchison, S. Cooper, V. Rush, and R. S. Parker. 2004. Sexual dichromatism
in the Yellow-breasted Chat (Icteria virens): spectrophotometric
analysis and biochemical basis. J. Avian Biol. 35:125-134. PDF
31. McGraw, K. J. and G. E. Hill. 2004. Mate attentiveness, seasonal timing of breeding
and long-term pair bonding in the house finch (Carpodacus mexicanus).
Behaviour 141:1-13. PDF
30. McGraw, K. J. and D. R. Ardia. 2004. Immunoregulatory activity of different dietary
carotenoids in male zebra finches. Chemoecol. 14:25-29. PDF
29. Hill, G. E. and K. J. McGraw. 2004. Correlated changes in male plumage
coloration and female mate choice in cardueline finches. Anim. Behav. 67:27-35.
PDF
28. McGraw, K. J. and D. R. Ardia. 2003. Carotenoids, immunocompetence, and the
information content of sexual colors: an experimental test. Am. Nat.
162:704-712. PDF
27. Saks, L., K. J. McGraw, and P. Horak. 2003. How feather colour reflects
its carotenoid content. Funct. Ecol. 17:555-561. PDF
26. McGraw, K. J., M. D. Beebee, G. E. Hill, and R. S. Parker. 2003. Lutein-based
plumage coloration in songbirds is a consequence of selective pigment
incorporation into feathers. Comp. Biochem. Physiol. B. 135:689-696. PDF
25. McGraw, K. J., G. E. Hill, and R. S. Parker. 2003. Carotenoid pigments in a mutant
cardinal: implications for the genetic and enzymatic control mechanisms of
carotenoid metabolism in birds. Condor 105:587-592. PDF
24. McGraw, K. J., A. J. Gregory, R. S. Parker, and E. Adkins-Regan. 2003. Diet, plasma
carotenoids, and sexual coloration in the zebra finch (Taeniopygia guttata).
Auk 120:400-410. PDF
23. McGraw, K. J. 2003. Melanins, metals, and mate quality. Oikos 102:402-406. PDF
22. Hill, G. E and K. J. McGraw. 2003. Melanin, nutrition, and the lions mane.
Science (Letter) 299:660. PDF
21. McGraw, K. J., J. Dale, and E. A. Mackillop. 2003. Social environment during molt
and the expression of melanin-based plumage pigmentation in male house
sparrows (Passer domesticus).
Behav. Ecol. Sociobiol. 53:116-122. PDF
20. McGraw, K. J., E. A. Mackillop, J. Dale, and M. E. Hauber. 2002. Different colors
reveal different information: how nutritional stress affects the expression of
melanin- and structurally based ornamental coloration. J. Exp. Biol.
205:3747-3755. PDF, JEB 2002 Highlight
19. McGraw, K. J., E. Adkins-Regan, and R. S. Parker. 2002. Anhydrolutein in the zebra
finch: a new, metabolically derived carotenoid in birds. Comp. Biochem.
Physiol. B. 132:811-818. PDF
18. McGraw, K. J. 2002. Environmental predictors of geographic variation in human mating
preferences. Ethology 108:303-317. PDF,
Reuters
press release, New Scientist
article, Straits
Times, Ananova,
iAfrica, Daily Dispatch,
NY Post
editorial, Rochester
(N.Y.) Democrat & Chronicle, Yahoo News (in Spanish), Expressen (Swedish
newspaper), Het
Parool (Dutch daily newspaper), Ithaca
(N.Y.) Journal, Atlanta
Journal-Constitution, Cornell
Univ. press release, Australian
Broadcasting Company, Hindustan
Times (India), Megastar
(England), La Opinion
(Los Angeles), Washington
Post, Hola
Hoy (NYC Spanish newspaper), Chicago
Sun-Times, Dagbladet
(Norwegian newspaper), Indianapolis Star,
Terra Lycos,
Telegraph News (U.K.), Cornell
Chronicle, National
Review, Some
girl named Allison, San
Francisco Chronicle, Indianapolis Star again,
Chicago
Tribune, Indianapolis
Star yet again, Buffalo News,
New
Orleans Channel, Financial Times magazine
(London), EurekAlert
(AAAS), Cosmiverse,
InteliHealth,
El Norte,
El Universal, KLIK
Magazin, GIGA
(Denmark), Newswise, Mindful-Things,
SexNews Daily, The
KCRA Channel (California), Grinning
Idiot, Venezuela
Innovadora, Tiscali
(Netherlands), Tercera
(Chile), Shape Magazine, Allure Magazine, Cornell Communique, Cornell Alumni Magazine, YupiMSN,
Avisa Norland (Norway), Aftonbladet
(Sweden), 3sat
Online (Denmark), Modern Bride magazine,
Most Frequently Dowloaded
Ethology paper in 2003
17.
McGraw, K. J., G. E.
Hill, R. Stradi, and R. S. Parker. 2002. The effect of dietary carotenoid
access on sexual dichromatism and plumage pigment composition in the American
goldfinch. Comp. Biochem. Physiol. B 131:261-269. PDF
16. Badyaev, A. V.,
Hill, G. E., Beck, M. L., Dervan, A. A., Duckworth, R. A., McGraw, K. J., Nolan,
P. M., and Whittingham, L. A.
2002. Sex-biased hatching order and adaptive population
divergence in a passerine bird. Science 295:316-318. PDF, News of the
Week, USA
Today, National
Geographic News.
15. McGraw, K. J. and G. E. Hill. 2002. Testing reversed sexual dominance from an
ontogenetic perspective: juvenile female House Finches Carpodacus mexicanus
are dominant to juvenile males. Ibis 144:39-42. PDF
14. McGraw, K.
J., P. M. Nolan, A. M. Stoehr,
and G. E. Hill. 2001. Intersexual differences in age-specific parental effort
in the house finch. Etologia 9:35-41. PDF,
Journal cover
13. McGraw, K. J., G. E. Hill, R. Stradi, and R. S. Parker. 2001. The influence of
carotenoid acquisition and utilization on the maintenance of species-typical
plumage pigmentation in male American goldfinches (Carduelis tristis)
and northern cardinalis (Cardinalis cardinalis). Physiol. Biochem. Zool. 74:843-852. PDF,
Online
version with color photos
12. McGraw, K. J. and G. E. Hill. 2001. Carotenoid access and intraspecific variation in
plumage pigmentation in male American goldfinches (Carduelis tristis)
and northern cardinals (Cardinalis cardinalis). Funct. Ecol. 15:732-739. PDF, Journal cover
11. McGraw, K. J., A. M. Stoehr, P. M. Nolan, and G. E. Hill. 2001. Plumage redness
predicts breeding onset and reproductive success in the house finch: a validation
of Darwin's theory. J. Avian Biol. 32:90-94. PDF
10. Stoehr, A. M. and K. J. McGraw. 2001. Ultraviolet reflectance of color patches in
male Sceloporus undulatus and Anolis carolinensis. J. Herpetol. 35:168-171. PDF
9. Stoehr, A. M, K. J. McGraw, P. M. Nolan, and G. E. Hill. 2001. Parental care
in relation to brood size in the house finch. J. Field Ornithol. 72:412-418. PDF
8. Nolan, P. M., A. M. Stoehr, G. E. Hill and K.
J. McGraw. 2001. The number of
provisioning visits by house finches predicts the mass of food delivered.
Condor 103:851-855. PDF
7. McGraw, K. J. and G. E. Hill. 2000. Plumage brightness and breeding-season dominance
in the house finch: a negatively correlated handicap? Condor 102:457-462. PDF
6. McGraw, K. J. and G. E. Hill. 2000. Differential effects of endoparasitism on the
expression of carotenoid- and melanin-based ornamental coloration. Proc. R.
Soc. Lond. B. 267:1525-1532. PDF, NPR Radio Show, Birdwatcher’s Digest
5. McGraw, K. J. and G. E. Hill. 2000. Carotenoid-based ornamentation and status
signaling in the house finch. Behav. Ecol. 11: 520-527. PDF
4. Stoehr, A. M., P. M. Nolan, G. E. Hill, and K.
J. McGraw. 2000. Nest mites (Pellonyssus
reedi) and the reproductive biology of the house finch (Carpodacus
mexicanus). Can. J. Zool. 78:2126-2133. PDF
3. Badyaev, A. V., G. E. Hill, A. M. Stoehr, P. M.
Nolan, and K. J. McGraw. 2000. The
evolution of sexual dimorphism in the house finch: II. Population divergence in
relation to local selection. Evolution 54:2134-2144. PDF
2. McGraw, K. J. and G. E. Hill. 1999. Induced homosexual behaviour in male house
finches (Carpodacus mexicanus): the "Prisoner Effect".
Ethol. Ecol. Evol. 11:197-201. PDF
1. McGraw, K. J., G. E. Hill and A. J. Keyser. 1999. Ultraviolet reflectance of
colored plastic leg bands. J. Field Ornithol. 70:236-243. PDF
Popular
articles
McGraw, K. J. 2005. Polly want a
pigment? Cracking the chemical code to red coloration in parrots. Australian Birdkeeper Magazine 18:608-611. PDF
McGraw, K. J. 2004. What flamingos
might look like if they ate blueberries.
ASK magazine (Arts and Sciences for Kids)
3:33. PDF
McGraw, K. J.
and G. E. Hill. 2003. Why birds wear bright feathers. Highlights for Children
58:24-25. PDF, Rochester
Democrat and Chronicle (24 July 2003), Families.com
McGraw, K. J.
2003. Color me chemically: an analysis of feather pigments in colorful birds.
The Sample (Retsch Inc. newsletter) 23:6-7. PDF
1) Societies
American Association
for the Advancement of Science (AAAS)
Association of
Field Ornithologists, Inc.
Federation of
American Societies for Experimental Biology (FASEB)
International
Carotenoid Society
International
Society for Behavioral Ecology
Society for
Integrative and Comparative Biology
2) Journals
Avian ecology and
evolutionary biology journals
Others not on that page =
Ethology
Ecology and Evolution
Physiological
and Biochemical Zoology
3) Graduate Student Research Grants
Animal
Behavior Society (Student Research Grants)
American
Ornithologists Union (Research Awards)
Association
of Field Ornithologists (E. Alexander Bergstrom Memorial Research Award)
BIRDNET Grants, Awards,
and Prizes in Ornithology
American Museum of
Natural History (Frank F. Chapman Memorial Grant)
Wilson Ornithological
Society (Research Grants)
Environmental
Protection Agency (Science-to-Achieve-Results (STAR) fellowship)
4) Sports
Cornell
University baseball program
Arizona
State University athletics
5) Others
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Protection Agency
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Created: 23-Sept-98
Last Revised: 4-Feb-10
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