• Ardia, Dan
Associate Professor of Biology



Office: LSP232B


Associate Professor, Franklin and Marshall College, Lancaster, PA; Department of Biology 2012-Present; Assistant Professor 2006-2012; Chair, Biological Foundations of Behavior Program 2012-2014

Visiting Scholar, Department of Ecology & Evolutionary Biology, Cornell University, Ithaca, NY 2012-Present

Research Associate, Migratory Bird Center, Smithsonian Institution, 2014-2015

Faculty Fellow, Ecosystems Center, Marine Biological Laboratory, 2009


B.S. Tufts University: Biology, Environmental Studies, & Political Science

M.S. SUNY-College of Environmental Science and Forestry: Environmental and Forest Biology

Ph.D. Cornell University: Ecology and Evolutionary Biology

Postdoc: Darwin Fellow: UMass-Amherst & Amherst College


Organism-environment interactions primarily of birds and mammals; evolutionary biology, behavioral ecology, physiological ecology 

Course Information

  • Evolution
  • Behavioral Ecology
  • Conservation Biology
  • Principles of Evolution
  • Ecology & Heredity



! = undergraduate author
 authors contributed equally

Rivers, J., Newberry, G., Schwarz, C and D.R. Ardia. Success despite the stress: violet-green swallows increase glucocorticoids and maintain reproductive output following experimental increases in flight costs. In Press Functional Ecology

Lathi, D.C. and D.R. Ardia. 2016 Shedding light on bird egg color: pigment as parasol and the hot car effect. American Naturalist. 187:

Wischhoff, U, F. Marques-Santos, D.R. Ardia, and J.J. Roper. 2015. White-rumped swallows prospect while they are actively nesting. Journal of Ethology: 33:145-150.

E Ospina, C Cooper, M Liljesthröm, D.R. Ardia, D.W. Winkler. 2014  Biparental nest attendance in Chilean Swallows (Tachycineta meyeni) breeding in Ushuaia, Argentina. Emu: 155:76-79.

Martin, L.B., D.H. Hawley, and D.R. Ardia. 2014. A New Division of Ecoimmunology and Disease Ecology. Integrative and Comparative Biology 54:338-339

Winkler, D. W., K. M. Ringelman, … D.R. Ardia. 2014. Latitudinal variation in clutch size–lay date regressions in Tachycineta swallows: effects of food supply or demography? Ecography 37:670-678.

Windsor!, R.E., J.E. Fegely!, and D.R. Ardia. 2013. Factors affecting the thermal insulation properties of tree swallow nests. Journal of Avian Biology. 44:305-310

Butler, M., Stahlschmidt, Z.R., Ardia, D.R., Davies, S., Guilette, L, Johnson, N., McCormick, S., McGraw, K.J., and D. DeNardo.  2013. Thermal sensitivity of immune function: evidence against a generalist-specialist tradeoff among endothermic and ectothermic vertebrates. American Naturalist. 181:761-784

Ardia, D.R. 2013. The effects of nestbox thermal environment on fledging success and haematocrit in Tree Swallows. Avian Biology Research 6:1-6.

Ardia, D.R., J.E. Gantz!, B.C. Schneider!, and S. Strebel!. 2012 Costs of immunity: an induced immune response increases metabolic rate and decreases antimicrobial activity in insects. Functional Ecology 26:732-739.

Stager, M., E. Lopresti, F.  Angulo, D. R. Ardia, D. Caceres, C.B. Cooper, E.E. Iñigo-Elias, J. Molina, N. Taylor, & D.W. Winkler. 2012. Reproductive biology of a narrowly endemic Tachycineta swallow in dry, seasonal forest in coastal Peru. Ornitología Neotropical 23:95-112.

Ardia, D.R., L. Vogel, and H. Parmentier. 2011. The role of constraints and limitation in driving individual variation in immune response. Special Issue of Functional Ecology on Ecological Immunology (L.B. Martin, D.R. Ardia, D.M. Hawley, editors.). 21:61-73.

Martin, L.B., D.M. Hawley, and D.R. Ardia. 2011. An introduction to ecological immunology. Functional Ecology 21:1-4.

Cooper, C.B., M.A. Voss, D.R. Ardia, S.B. Austin, W.D. Robinson. 2011. Light increases the rate of embryonic development: implications for latitudinal trends in incubation period. Functional Ecology. 25:769-776.

Ardia, D.R., J.H. Peréz!, and E.D. Clotfelter. 2010. Experimental cooling during incubation leads to reduced innate immunity and body condition in nestling tree swallows. Proceedings of the Royal Society of London B. 277:1881-1888.

Ardia, D.R., D.R. Broughton!, and M. Gleicher!. 2010. Short-term exposure to testosterone increases dominance interactions and enhances bill color in zebra finches. Hormones and Behavior 58:526-532.

Lambrechts, M.M., Adriaensen, F., D.R. Ardia, ….53 others. 2010. The design of artificial nestboxes for the study of secondary hole-nesting birds: a review of methodological inconsistencies and potential biases. Acta Ornithologica. 45:1-26.

Evans!, L.E., Ardia, D.R. & J.E.C. Flux. 2009. Breeding synchrony through social stimulation in a spatially-segregated population of European starling (Sturnus vulgaris). Animal Behaviour 78:671-675.

Morrison!, E., Ardia, D.R. & E.D. Clotfelter. 2009. Cross-fostering reveals sources of variation in innate immunity and hematocrit in nestling tree swallows. Journal of Avian Biology 40:573-576.

Stenzler, L. M., C.A. Makarewich, A. Coulon, D.R. Ardia, I.J. Lovette and D.W Winkler. 2009. Subtle edge-of-range genetic structuring in transcontinentally distributed North American Tree Swallows. Condor 111:470-478.

Ardia, D.R., J.H. Peréz!, E.K. Chad!, M.A. Voss, and E.D. Clotfelter. 2009. Temperature and life history: Experimental heating leads female tree swallows to modulate egg temperature and incubation behavior. Journal of Animal Ecology. 78:4-13.

Peréz!, J.H., Ardia, D.R., E.K. Chad!, and E.D. Clotfelter. 2008. Experimental heating reveals nest temperature affects nestling condition in tree swallows (Tachycineta bicolor). Biology Letters 4:468-471. Authors contributed equally.

Ardia, D.R. and K.A. Schat. 2008. Ecoimmunology. In: The Immunology of Birds (editors F. Davison, B. Kaspers and K. A. Schat). Elsevier.

McGraw, K.J. and D.R. Ardia. 2007. Do carotenoids buffer testosterone-induced immunosuppression? An experimental test in a colorful song bird. Biology Letters 3:375-378.

Clotfelter, E.D., D.R. Ardia, & K.J. McGraw. 2007. Red fish, blue fish: trade-offs between pigmentation and immunity in Betta splendens. Behavioral Ecology. 18:1139-1145. Authors contributed equally.

Ardia, D.R. 2007. Site and sex-level differences in adult feeding behaviour and its consequences to offspring quality in tree swallows (Tachycineta bicolor) following brood size manipulation. Canadian Journal of Zoology. 85:847-854.

Ardia, D.R. and E.D. Clotfelter. 2007. Individual quality and age affect responses to an energetic constraint in a cavity-nesting bird. Behavioral Ecology. 18:259-266.

Ardia, D.R. 2007. The ability to mount multiple immune responses simultaneously varies across the range of the tree swallow. Ecography 30:23-30.

Ardia, D.R. and E.B. Rice. 2006. Variation in heritability of immune function in the tree swallow. Evolutionary Ecology. 20:491-500.

Ardia, D.R. 2006. Geographic variation in the trade-off between nestling growth rate and body condition in the Tree Swallow. Condor. 108:601-611.

Ardia, D.R., J.H. Peréz!, and E.D. Clotfelter. 2006. Nest box orientation affects internal temperature and nest site selection by Tree Swallows. Journal of Field Ornithology.

Ardia, D.R., M.W. Wasson, and D.W. Winkler. 2006. Prey abundance and individual quality influence yolk and egg size and egg composition in tree swallows. Journal of Avian Biology. 37:252-259.

Ardia, D.R. and E.D. Clotfelter. 2006. A novel application of an immunological technique reveals the immunosuppressive effects of phytoestrogens in Betta splendens . Journal of Fish Biology. 68a:144-149.

Ardia, D.R., C.B. Cooper, and A.A. Dhondt. 2006. Warm temperatures lead to early onset of incubation, shorter incubation and greater hatching asynchrony in tree swallows at the extremes of their range. Journal of Avian Biology. 37:137-142.

Ardia, D.R. 2006. Glycated hemoglobin and albumin reflect nestling growth and condition in American kestrels. Comparative Biochemistry and Physiology, Part A . 143:62-66.  

Ardia, D.R. 2005. Tree swallows tradeoff immune function and reproductive effort differently across their range. Ecology. 86:2040-2046.  

Ardia, D.R. 2005. Individual quality mediates tradeoffs between reproductive effort and immune function in tree swallows. Journal of Animal Ecology. 74:517-524.  

Ardia, D.R. 2005. Cross-fostering reveals an effect of spleen size and body condition on immune responses in nestling European starlings. Oecologia. 145:326-333.  

Ardia, D.R. 2005. Super size me: an experimental test of factors affecting lipid content and the ability of residual body mass to predict lipid levels in nestling European starlings. Functional Ecology 19:414-420.  

Ardia, D.R. 2005. Natural selection and variation: a lesson using museum specimens. American Biology Teacher. 67:468-473.  

McGraw, K.J. and D.R. Ardia. 2005. Sex differences in carotenoid status and immune performance in zebra finches. Evolutionary Ecology Research. 7:251-262.

McGraw, K.J. and D.R. Ardia. 2004. Immunoregulatory activity of different dietary carotenoids in zebra finches. Chemoecology. 14:21-29.  

Ardia, D.R., K.A. Schat, and D.W. Winkler. 2003.  Reproductive effort decreases long-term immunity in breeding tree swallows. Proceedings of the Royal Society of London B. 270:1679-1683.  

McGraw, K.J. and D.R. Ardia. 2003. Carotenoids, immunocompetence, and the information content of sexual colors: an experimental test. American Naturalist. 162:704-712.  

Ardia, D.R. 2002. Energetic consequences of sex-related habitat segregation in wintering American kestrels. Canadian Journal of Zoology. 80:516-523.  

Ardia, D.R. and K.L. Bildstein. 2001. Sex-related differences in habitat use in wintering American Kestrels. Auk 118:746-749.  

Ardia, D.R. 2001. Winter roosting behavior of American kestrels. J. Raptor Research 35:58-60.

Ardia, D.R. and K.L. Bildstein. 1997. Sex-related differences in habitat selection in wintering American kestrels (Falco sparverius ). Animal Behaviour 53: 1305-1311.