My research
focuses on the
evolution of colorful plumage. In the nineteenth century, Charles
Darwin and Alfred
Wallace first recognized
the challenge that brightly colored
plumage
posed to the theory of evolution by natural selection. Why would
small vulnerable creatures like songbirds be brilliantly and
conspicuously
colored? The existence of colorful plumage presented one of the
strongest
challenges to natural selection theory.
Darwin proposed and argued for the idea
that female
mate choice drives the evolution of colorful plumage. Wallace
rejected
this idea and proposed various forms of natural selection - crypsis,
mimicry,
and species recognition - to explain colorful plumage. Until the
late 1980s, however, the ideas of Darwin and Wallace remained
untested.
My research program picks up where Darwin and Wallace left off.
My
students and I are experimentally testing the role of female mate
choice
in the evolution of various types of ornamental plumage. We are
also
studying the proximate control of variation in expression of colorful
plumage
to better understand the signal content of such ornamental display.
My research program is aimed at
understanding the function and evolution of the three types of
ornamental coloration of feathers: carotenoid pigmentation, melanin
pigmentation, and structurally based coloration. We currently
study two local species of birds - House Finches and Eastern Bluebirds
– and one species in Costa Rica - Long-tailed Manakins - that have
various combinations of these color displays in their plumage.
House Finches have carotenoid-based coloration; Eastern Bluebirds have
both structural blue and rust phaeomelanin coloration; and, Long-tailed
Manakins have structural blue, carotenoid red, and eumelanin black
coloration. Follow the links below for brief descriptions of ongoing
research projects.
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| House
Finches:
parasite interactions and maternal effects |
Studies of House Finches build on
substantial previous work in which we showed that male carotenoid-based
plumage coloration is an honest signal of male condition that is used
by females in choosing mates. Early studies were summarized in a book
in 2002. In recent studies with grad students, I have shown that
dietary access to carotenoid pigments, nutritional condition, and
parasites can each have a significant affect on expression of plumage
coloration.
Dr. Alex Badyaev
has been a long-time collaborator with me. Alex
and I have also been working on variation in sexual dimorphism and
selection on dimorphism across North America and particularly between
our focal study populations in Montana and Alabama. We have
shown
that males and females vary extensively and independently in size and
shape across N. American populations, and that the sizes and shapes of
males and females conform closely with the local optimum dictated by
survival, fecundity, and pairing selection. However, selection on
adult phenotypes cannot achieve the observed degree of dimorphism
evolution, so we have begun to look at ontogeny as a source of
divergence in dimorphism among populations. Most recently, Alex
and I found that House Finch chicks in different positions in the hatch
order grow and develop differently and that females in different
populations allocate males and females to positions in the hatch order
such that they create males and females that match the local optimum in
size and shape. We are now following up this study with a
detailed investigation of maternal effects.
With Kristen Navara, a
doctoral student who works jointly in my lab and
in the lab of Dr.
Mary Mendonca, we are
examining allocation patterns
of hormones and carotenoids into the eggs of House Finches and Eastern
Bluebirds. We are attempting to interpret allocation patterns in
the context of female investment strategies by examining their
physiological and immunological effects on the resulting offspring.
My interest in plumage coloration and disease led to a collaboration
with Dr.
Sharon Roberts, a
microbiologist at Auburn, and Dr.
Scott Edwards, a
population geneticist at the Harvard University.
Scott, Sharon, and I are studying the co-evolution of a new House Finch
disease, Mycoplasma gallicepticum (MG), and eastern populations of
House Finches, which have been exposed to the disease since 1994.
My primary goal in this project is to test whether expression of
carotenoid-based plumage coloration might be associated with specific
genes for disease resistance. The infection experiments that we
are conducting to assess change in resistance in the House Finch and
change in virulence in the microbe are being conducted by Kristy
Farmer, a doctoral student working jointly in my lab and the lab of Dr.
Roberts. In a recent study, Kristy and I found that redder male
were more resistant to MG infection than drabber males. We will
now attempt to find genetic correlates to MG resistance.
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| Eastern
Bluebirds: from nanostructure to macrocolor |
| A major focus of my research concerns the
function and evolution of
structurally based blue plumage coloration. Current research on
structural coloration is being conducted by my doctoral students, Lynn
Siefferman and Mark Liu, on the Eastern Bluebird, which has brilliant
blue/UV coloration. Lynn Siefferman’s work focuses on testing the
hypothesis that the structurally based blue
coloration of bluebirds
might serve as a condition-dependent signal of quality like
carotenoid-based coloration. So
far, she has found that plumage
blueness is positively related to nest box acquisition and feeding rate
of males. We are currently testing the relative importance of genes and
environment on expression of plumage blueness and looking at mate
guarding and male paternity in relation to plumage blueness. Mark Liu
is assessing the relative importance of ornamentation versus genetic
compatibility in mate choice in the bluebird. Mark will look at
mating patterns related to both structural blue coloration and genetic
complementarily and will assess the fitness of offspring produced by
ornamented males versus males that were genetically compatible with
their mates. |
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Long-tailed
Manakins: an excursion to the tropics
|
Stephanie Doucet, another doctoral
student, is studying the function
and evolution of elaborate
male ornaments in Neotropical
manakins. Stephanie’s project investigates the causes and
consequences of variation in ornamentation in Long-tailed
Manakins. Male manakins have elaborate, sexually selected black,
red, and blue coloration that results from melanin pigments, carotenoid
pigments, and structural elements, respectively. Stephanie is
conducting careful observations of the leks of these birds to determine
how plumage coloration affects male mating success. Because these
birds go through a long series of transitional plumage before attaining
definitive plumage in their fifth year, Stephanie will also look at the
ontogeny of the different color displays to gain insight into how the
different components of plumage function.
|

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Feathers at a fine scale: an integrative approach
|
| My doctoral student, Matt
Shawkey, is studying
feather color at a very
small scale, examining the nanostructural basis of variation in
coloration and the ways in which feather-degrading microbes might
affect this color. In bluebirds, he has found a number of nanoscale
morphological variables that predict color variation quite well.
Additionally, he has thoroughly characterized the microbial communities
on their feathers and has shown that some of these microbes can disrupt
feather morphology and cause drastic changes in color. He has
recently expanded these studies to species with carotenoid-based
coloration. |

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Song: the
forgotten ornament
|
| Although much of the research in my lab
group focuses on plumage
coloration, my students and I are also interested in the use of song as
a signal of condition, especially relative to female choice of mates.
With Dr. Dan
Mennill, a postdoc in my
lab, and Alex Badyaev, I am
studying how song functions as a signal in a Montana population of
House Finches. This Montana population is particularly
appropriate for such a study because Alex has detailed information on
pairing success, nesting success, provisioning rate, extra-pair
paternity etc. that can compared to song parameters. Dan also
studies the ecology and evolution of vocal duetting in neotropical
Thryothorus wrens and female eavesdropping on male song contests in
both temperate and tropical territorial songbirds. |

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Molecular Ecology: the new frontier
|
| Dr.
Herman Mays joined my lab
group
as a postdoc in 2001, and he helped
us establish a molecular ecology lab where we can conduct genetic
sexing, paternity analysis, and molecular systematics. Herman is
currently working with my former master's student, Michelle Beck (now
at Washington State University), on an
analysis of House Finch paternity, with my Ph.D. students, Lynn
Siefferman and Mark Liu, on Eastern Bluebird paternity, and with my
former master's students Barb Ballentine (now at Duke University) and
Amber
Keyser (recently finished her Ph.D. at the University of Georgia) on
Blue Grosbeak
paternity. Bill Ashwander, an undergraduate student at Auburn, has been
working with
Herman on a phylogeny of House Finch subspecies and populations using
mitochrondrial DNA. The genetic expertise that Herman brings to
my research group has provided exciting new opportunities for the study
of sexual selection, mating systems, and plumage coloration. |


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Arboviruses:
applied biology
|
| Although I am primarily a behavioral and
evolutionary biologist,
I also have been involved in a number of applied projects looking at
bird habitat use. I am currently working on a study of the
ecology of encephalitis virus in a swamp forest in Alabama. This
research is being conducted with Dr. Ed Cupp, an entomologist at Auburn
who is conducting companion sampling of the mosquitoes of this area,
and Dr. Tom Unnasch, who is identifying the blood meals of
mosquitoes. A primary goal of this study is to identify the
feeding preferences of bird-feeding mosquitoes, and this requires a
null hypothesis based on random bird encounters. Toward this end, Laura
Estep, a doctoral student in my lab, is using her skills in spatial
ecology and geostatistics to create Kriged maps representing the
abundances of all birds within the feeding range of mosquitoes.
From this spatial abundance data set, Laura will develop a model of
mosquito movement and construct a null model for host feeding.
Laura’s null model can then be compared to observed blood-meal
composition. This is a novel approach to the study of the ecology
of arboviruses and holds great promise for helping to better understand
these diseases. |

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Last Revised: January 2005