Linda A. Whittingham    


Department of Biological Sciences
University of Wisconsin-Milwaukee
P.O. Box 413
Milwaukee, WI 53201
Phone: 414-229-2252
Fax: 414-229-3926


1992 Ph.D. (Biology) Queen's University, Canada

1988 M.S. (Biology) University of Michigan

1983 B.S. (Biology) University of Michigan

Ecology and Evolution of Parental Care
Evolution and Ecology of Birds (BioSci 523) 
Behavioral and evolutionary ecology of birds 
Survey of Zoology (BioSci 100) 
Sexual selection, Mating systems  
Found. of Biol. Sci. I (BioSci 150) 
Dr. Whittingham is not accepting new graduate students at this time

 Tree Swallows

 Offspring sex ratio and

female body condition

 Repeatability of

extrapair mate choice

Collaborators: _Peter Dunn (UWM)



 Alternative mating strategies in

House Wrens
 Mate switching
 Male mating strategies and reproductive success
 Mate guarding and paternity

   Common Yellowthroat _  
Male mask size and mating success
Paternity and male parental care

Male traits and female choice:

aviary experiments

 Male traits and offspring sex




Evolution of sexual dimorphism and mating systems in birds
Comparative analyses of sexual dimorphism in relation to 
social mating systems (e.g., monogamy, polygyny) and sperm competition 
(indexed by testes size).
 Publications: Evolution 55: 161-175
 Collaborators: _Peter Dunn (UWM) and Trevor Pitcher (Univ. of Toronto)




Councilor, International Society for Behavioral Ecology, 2000-2004.


Board of Directors, Bluebird Restoration Association of Wisconsin, 2001-2004


TAFF, C.C. , C.R. FREEMAN-GALLANT, P.O. DUNN & L.A. WHITTINGHAM. 2013. Spatial distribution of nests constrains the strength of
sexual selection in a warbler. Journal of Evolutionary Biology, in press.

DUNN, P.O., J.L. BOLLMER, C.R. FREEMAN-GALLANT & L.A. WHITTINGHAM. 2013. MHC variation is related to a sexually selected ornament,
survival, and parasite resistance in the common yellowthroat. Evolution 67: 679-687.

EIMES, J.A., K. REED, K. MENDOZA, J.L. BOLLMER, L.A. WHITTINGHAM, Z.W. BATESON & P.O. DUNN. 2013. Greater prairie-chickens have a
compact MHC-B with a single class IA locus. Immunogenetics 65: 133-144.

BOLLMER, J.L., P.O. DUNN, C.R. FREEMAN-GALLANT & L.A. WHITTINGHAM. 2012. Social and extra-pair mating in relation to MHC
variation in common yellowthroats. 2012. Proceedings of the Royal Society of London B 279: 4778-4785.

TAFF, C.C., C.R. FREEMAN-GALLANT, P.O. DUNN , L.A. WHITTINGHAM, K. Belinsky, D. Steinberger, C. Clark, H. Sacks. 2012. Multi-modal sexual
selection in a warbler: plumage and song are related to different fitness components. Animal Behaviour 84: 813-821.

M. STANBACK. 2012. Spatiotemporal Patterns in Nest Box Occupancy by Tree Swallows Across North America. Avian Conservation and Ecology 7
(1): 3.

HESS, B.D., P.O. DUNN & L.A. WHITTINGHAM. 2012. Females choose multiple mates in the lekking greater prairie-chicken. Auk 129: 133-139.

EIMES, J.A., J.L. BOLLMER, L.A. WHITTINGHAM, J.A. JOHNSON, C. VAN OOSTERHOUT & P.O. DUNN. 2011. Rapid loss of MHC class II variation in
a bottlenecked population is explained by drift and loss of copy number variation. Journal of Evolutionary Biology 24: 1847-1856.

TAFF, C.C., C.R. FREEMAN-GALLANT, P.O. DUNN & L.A. WHITTINGHAM. 2011. Relationship between brood sex ratio and male
ornaments depends on male age in a warbler. Animal Behaviour 81: 619-625.

DUNN, P.O., D.W. WINKLER, L.A. WHITTINGHAM, S.J. HANNON, & R.J. ROBERTSON. 2011. A test of the mismatch hypothesis:
how predictable is the timing of food abundance for aerial insectivores? Ecology 92: 450-461.

WHITTINGHAM, L.A., & P.O. DUNN. 2010. Fitness benefits of polyandry for experienced females. Molecular Ecology 19:2328-2335.

BOLLMER, J.L. , P.O. DUNN, L.A. WHITTINGHAM, & C. WIMPEE. 2010. Extensive MHC class II B gene duplication in a passerine, the
Common Yellowthroat (Geothlypis trichas). Journal of Heredity 101: 448-460.

FREEMAN-GALLANT, C.R., C.C. TAFF, D.F. MORIN, P.O. DUNN, L.A. WHITTINGHAM, & S.M. TANG. 2010. Sexual selection, multiple ornaments,
and age- and condition-dependent signalling in the common yellowthroat. Evolution. 64: 1007-1017.

EIMES, J.A., J.L. BOLLMER, P.O. DUNN. L.A. WHITTINGHAM, & C. WIMPEE. 2010. MHC class II diversity and balancing selection
in greater prairie-chickens. Genetica 138: 265-271.

DUNN. P.O., J. GARVIN, L.A. WHITTINGHAM, C. FREEMAN-GALLANT, & D. HASSELQUIST. 2010. Carotenoid and melanin-based ornaments
signal similar aspects of male quality in two populations of the common yellowthroat. Functional Ecology 24:149-158.

DUNN. P.O., J.T. LIFJELD, & L.A. WHITTINGHAM. 2009. Multiple paternity and nestling quality in tree swallows. Behavioral Ecology
and Sociobiology 63:911-922.

ARMENTA, J., P.O. DUNN, & L.A. WHITTINGHAM. 2008. Effects of specimen age on plumage color. Auk 125: 803-808.

ARMENTA, J., P.O. DUNN, & L.A. WHITTINGHAM. 2008. Quantifying avian sexual dichromatism: a comparison of methods.
Journal of Experimental Biology 211:2423-2430.

DUNN, P.O., L.A. WHITTINGHAM, C. FREEMAN-GALLANT, & J. DeCOSTE. 2008. Geographic variation in the function of ornaments in the
common yellowthroat. Journal of Avian Biology 39:66-72.

GARVIN, J., P.O. DUNN, L.A. WHITTINGHAM, D. STEEBER, & D. HASSELQUIST. 2008. Do male ornaments signal immunity in
common yellowthroats? Behavioral Ecology 19: 54-60.

DALE, J. . P. O. DUNN, J. FIGUEROLA, T. LISLEVAND, T. SZEKELY & L. A. WHITTINGHAM. 2007. Sexual selection explains
Rensch's rule of allometry for sexual size dimorphism. Proceedings of the Royal Society of London B 274: 2971-2979.

WHITTINGHAM, L.A., P.O. DUNN, & J.T. LIFJELD. 2007. Egg size is related to nestling quality, but there is no differential
allocation in tree swallows. Condor 109: 585-594.

MITCHELL, D.P., P. O. DUNN, L.A. WHITTINGHAM & C. FREEMAN-GALLANT. 2007. Attractive males provide less parental care in
two populations of the common yellowthroat. Animal Behaviour 73: 165-170

ABROE, B., GARVIN, J., PEDERSEN, M., WHITTINGHAM, L.A., & P.O. DUNN. 2007. Brood sex ratios are related to male size, but
not attractiveness in the common yellowthroat. Auk 124: 176-184.

DUNN. P. O. & L. A. WHITTINGHAM. 2007. Search costs influence the spatial distribution, but not the level of extra-pair mating
in tree swallows. Behavioral Ecology and Sociobiology 61: 449-454.

PEDERSEN, M., P. O. DUNN, & L. A. WHITTINGHAM. 2006. Extra-territorial forays are related to a male ornamental trait in
common yellowthroats. Animal Behaviour 72: 479-486.

GARVIN, J., B. ABROE, B., M. PEDERSEN, P.O. DUNN & L.A. WHITTINGHAM. 2006. Immune response of nestling warblers varies
with extra-pair paternity and temperature. Molecular Ecology 15: 3833-3840.

WHITTINGHAM, L.A., P.O. DUNN, & STAPLETON, M. 2006. Repeatability of extra-pair mate choice by female tree swallows.
Molecular Ecology 15: 841-849.

DUNN, P.O. & L. A. WHITTINGHAM. 2005. Radio-tracking of female tree swallows prior to egg-laying. Journal of
Field Ornithology 76: 259-263.

NOOKER, J., P. O. DUNN, & L. A. WHITTINGHAM. 2005. Effects of food abundance, weather and female condition on
reproduction in tree swallows. Auk 122: 1225-1238.

WHITTINGHAM, L.A., P.O. DUNN, & NOOKER, J. 2005. Maternal influences on brood sex ratios: an experimental study
 in tree swallows. Proceedings of the Royal Society of London B 272: 1775-1780.

TAROF, S., P.O. DUNN & L. A. WHITTINGHAM. 2005. Dual functions of a melanin-based ornament in the common
yellowthroat. Proceedings of the Royal Society of London B 272: 1121-1127.

PITCHER, T., P.O. DUNN, & L.A. WHITTINGHAM. 2005. Sperm competition and the evolution of avian testes size.
Journal of Evolutionary Biology 18: 557-567.

Whittingham, L. A. and P.O. Dunn. 2004. Effects of extra-pair and within-pair reproductive success on the opportunity for selection in birds. Behavioral Ecology in press.

Brylawski, A.M.Z. and L.A. Whittingham. 2004. An experimental study of mate guarding and paternity in house wrens. Animal Behaviour in press.

Poirier, NE, LA Whittingham and PO Dunn. 2004. Males achieve greater reproductive success through multiple broods than through extra-pair mating in house wrens. Animal Behaviour 67: 1109-1116.

Poirier, NE, LA Whittingham and PO Dunn. 2003. Effects of paternity and mate availability on mate switching in house wrens. Condor 105: 816-821.

Badyaev A. V., M. L. Beck, G. E. Hill, L. A. Whittingham. 2003. The evolution of sexual size dimorphism in the house finch: V. Maternal effects. Evolution 57: 384-396.

Whittingham, L. A., Dunn, P. O. and E. D. Clotfelter. 2003. Parental allocation of food to nestling tree swallows: the influence of nestling behavior, sex, and paternity. Animal Behaviour 65: 1203-1210.

Badyaev, A. V., G. E. Hill, L. A. Whittingham. 2002. Population consequences of maternal effects: sex-bias in egg-laying order facilitates divergence in sexual dimorphism between bird populations. Journal of Evolutionary Biology 15: 997-1003.

Whittingham, L. A., S. M. Valkenaar, N. E. Poirier and P. O. Dunn. 2002. Maternal condition and nestling sex ratio in house wrens. Auk 119: 125-130.

Whittingham, L. A., B. Slikas, F. H. Sheldon and D. W. Winkler. 2002. Phylogeny of the tree swallow genus (Aves: Hirundinidae: Tachycineta) by Bayesian analysis of mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 22: 430-441.

Whittingham, L. A. and H. Schwabl. 2002. Maternal testosterone in tree swallow eggs varies with female aggression. Animal Behaviour 63: 63-67.

Badyaev, A.V, G. E. Hill, M. L. Beck, A. A. Dervan, R. A. Duckworth, K.J. McGraw, P. M. Nolan, and L. A. Whittingham. 2002. Sex-biased hatching order and adaptive population divergence in a passerine bird. Science 295: 316-318.

Lifjeld, J.T., P.O. Dunn and L.A. Whittingham. 2001. Short-term fluctuations in cellular immunity of adults tree swallows feeding nestlings. Oecologia 130: 185-190.

Stapleton, M., P.O. Dunn, J. McCarty, A. Secord and L.A. Whittingham. 2001. Polychlorinated biphenyl contamination and minisatellite DNA mutation rates of tree swallows. Environmental Toxicology and Chemistry 20: 2263-2267.

Thusius, K. J., K. A. Peterson, P. O. Dunn and L. A. Whittingham. 2001. Sexual selection through within-pair and extra-pair mate choice in the common yellowthroat (Geothylpis trichas). Animal Behaviour 62: 435-446..

Thusius, K. J., P. O. Dunn, K. A. Peterson and L. A. Whittingham. 2001. Extrapair paternity is influenced by breeding synchrony and density in the common yellowthroat. Behavioral Ecology 12:633-639.

Whittingham, L. A. and P. O. Dunn. 2001. Male parental care and paternity in birds. Current Ornithology 16: 257-298.

Peterson, K. A., K. J. Thusius, L. A. Whittingham, and P.O. Dunn. 2001. Allocation of male parental care in relation to paternity within and among broods of the common yellowthroat Geothylpis trichas. Ethology 107: 573-586..

Whittingham, L. A. and P. O. Dunn. 2001. Survival of extrapair and withinpair offspring in tree swallows. Behavioral Ecology 12: 496-500.

Badyaev, A.V., L. A. Whittingham and G. Hill. 2001. The Evolution of sexual dimorphism in the house finch: III Developmental basis. Evolution 55: 176-189.

Dunn, P. O., L. A. Whittingham and T. E. Pitcher. 2001. Mating systems, sperm competition and the evolution of sexual dimorphism in birds. Evolution 55: 161-175..

Whittingham, L. A. and P. O. Dunn. 2001. Responses of female tree swallows to intraspecific brood parasitism. Condor 103: 166-170.

Clotfelter, E. D., L. A. Whittingham and P. O. Dunn. 2000. Laying order, hatching order and their effect on body mass in nestling tree swallows (Tachycineta bicolor). Journal of Avian Biology 31: 329-334.

Whittingham, L. A. and P. O. Dunn. 2000. Offspring sex ratios in tree swallows: females in better condition produce more sons. Molecular Ecology. 9: 1123-1129.

Whittingham, L. A., F. H. Sheldon and S. T. Emlen. 2000. Molecular phylogeny of jacanas and its implications for morphologic and biogeographic evolution. The Auk 117: 22-32.

Kirchman, J. J., L. A. Whittingham and F. H. Sheldon. 2000. Relationships among cave swallow (Petrochelidon fulva) populations determined by comparisons of microsatellite and cytochrome-b data. Molecular Phylogenetics and Evolution 14: 107-121.

Sheldon, F. H., L. A. Whittingham and D. W. Winkler. 1999. A comparison of cytochrome-b and DNA hybridization data bearing on the phylogeny of swallows (Aves: Hirundinidae). Molecular Phylogenetics and Evolution 11: 320-331

Whittingham, L. A. and P. O. Dunn. 1998. Male parental effort and paternity in a variable mating system. Animal Behaviour 55: 629-640.

Magrath, R.D. and L.A. Whittingham. 1997. Kinship, cooperation and non-cooperation in white-browed scrubwrens. Behavioral Ecology and Sociobiology 41: 185-192.

Whittingham, L. A., A. Kirkconnell and L. M. Ratcliffe. 1997. The context and function of duets and solo songs in Red-shouldered Blackbirds (Agelaius assimilis). The Wilson Bulletin 109: 279-289

Whittingham, L. A., P. O. Dunn and R. D. Magrath. 1997. Relatedness, polyandry and extra-group paternity in the cooperatively breeding white-browed scrubwren (Sericornis frontalis ). Behavioral Ecology and Sociobiology 40: 261-270.

Whittingham, L. A., A. Kirkconnell and L. M. Ratcliffe. 1996. Breeding behavior, social organization and morphology of Red-shouldered (Agelaius assimilis) and Tawny-shouldered (A. humeralis) blackbirds. The Condor 98: 832-836.

Whittingham, L. A. and J. T. Lifjeld. 1995. Extra-pair fertilizations increase the opportunity for sexual selection in the monogamous house martin Delichon urbica. Journal of Avian Biology 26: 283-288.

Whittingham, L. A. and J. T. Lifjeld. 1995. High paternal investment in unrelated young: extra-pair paternity and male parental care in house martins. Behavioral Ecology and Sociobiology 37: 103-108.

Whittingham, L. A., P. O. Dunn and R. J. Robertson. 1995. Testing the female mate guarding hypothesis. Animal Behaviour 50: 277-279.

Whittingham, L. A. 1994. Additional mating opportunities and male parental care in red-winged blackbirds. Animal Behaviour 48: 875-883.

Whittingham, L. A., P. O. Dunn and R. J. Robertson. 1994. Female response to reduced male parental care in birds: an experimental study in tree swallows. Ethology 96: 260-269.

Whittingham, L. A., P. O. Dunn and R. J. Robertson. 1994. Do female tree swallows guard their mates by copulating frequently ? Animal Behaviour 47: 994-997.

Whittingham, L. A. and R. J. Robertson. 1994. Food availability, male parental care and mating success in red-winged blackbirds (Agelaius phoeniceus). Journal of Animal Ecology 63: 139-150.

Dunn, P. O., L. A. Whittingham, J. T. Lifjeld, R. J. Robertson and P. T. Boag. 1994. Effects of breeding density, synchrony and experience on extra-pair paternity in tree swallows. Behavioral Ecology 5: 123-129.

Quinn, J. S., L. A. Whittingham and R. D. Morris. 1994. Infanticide in skimmers and terns: side effects of territorial attacks or intergenerational conflict ? Animal Behaviour 47 : 363-367.

Whittingham, L. A. and R. J. Robertson. 1993. Nestling hunger and parental care in Red-winged Blackbirds. The Auk 110: 240-246.

Whittingham, L. A., P. O. Dunn and R. J. Robertson. 1993. Confidence of paternity and male parental care: an experimental study in tree swallows. Animal Behaviour 46: 139-147.

Whittingham, L. A. 1993. Effects of nestling provisioning on time-activity budgets of male red-winged blackbirds. Condor 95: 730-734.

Whittingham, L. A., A. Kirkconnell and L. M. Ratcliffe. 1992. Differences in song and sexual dimorphism in Cuban and North American Red-winged Blackbirds (Agelaius phoeniceus). The Auk 109: 928-933.

Whittingham, L. A., P. D. Taylor and R. J. Robertson. 1992. Confidence of paternity and male parental care. American Naturalist 139: 1115-1125.

Whittingham, L. A.. 1989. An experimental study of paternal behavior in red-winged blackbirds. Behavioral Ecology and Sociobiology 25: 73-80.


Book Chapters:

Sheldon, F. H. and L. A. Whittingham. 1997. Using molecular phylogenies to study bird ecology, behavior, and morphology. In: Avian Molecular Evolution and Systematics (D. Mindell, ed.). Academic Press, San Diego. pp. 279-300.