• F&M Associate Professor of Biology Daniel Ardia
Charles A. Dana Professor of Biology, Associate Dean of the Faculty



Office: LSP232B


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

Visiting Scholar, Danish Institute of Study Abroad, Copenhagen, Denmark, 2018 

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; urban ecology, evolutionary biology, behavioral ecology, physiological ecology 

Course Information

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



! = undergraduate author
 authors contributed equally


Handler, A, E. Lonsdorf, D.R. Ardia. In Press. Evidence for red fox (Vulpes vulpes) exploitation of anthropogenic food sources along an urbanization gradient using stable isotope analysis. Canadian Journal of Zoology.

Zimmer, C., C. Taff, D.R. Ardia, D.W. Winkler, M.N. Vitousek. 2019. On again, off again: acute stress response and negative feedback together predict resilience to experimental stressors. Functional Ecology. 33:619-628.

Vitousek, M., C. Taff, D.R. Ardia, J. Stedman, C. Zimmer, T. Salzman, and D.W. Winkler. 2018. The lingering impact of stress: brief acute glucocorticoid exposure has sustained, dose-dependent effects on reproduction. Proceedings of the Royal Society of London B. 285:20180722.

Hardt, B.M, D.R. Ardia, M.J. Bashaw, and J.W Rivers. 2018. Experimental brood enlargement differentially influences the magnitude of the corticosterone stress response in two closely related and co-occurring songbirds Functional Ecology. 32:2008-2018.

Kilvitis, H.J., D.R. Ardia, M. Thiam, L.B. Martin. 2018. Corticosterone is correlated to mediators of neural plasticity and epigenetic potential in the hippocampus of Senegalese house sparrows (Passer domesticus). General and Comparative Endocrinology 269:177-183.

George!, D.R, B.S. Schneider!, K.J. McGraw, and D.R. Ardia. 2017. Carotenoids buffer the acute phase response on fever, sickness behavior, and rapid bill color change in zebra finches. Journal of Experimental Biology 220:2957-2964.

Brace, A., M. Lajeuness, J. Adelman, D.R. Ardia, D. Hawley, K. Buchanan, J. Fair, J. Grindstaff, K. Matson, and L.B. Martin. 2017. Costs of immune responses are related to body mass and life history. Journal of Experimental Zoology A 327:254-261.

Martin, L.B., Kilvitis, H., Thiam, M. and D.R. Ardia. 2017. Stress hormone regulation in house sparrows invading Senegal. General and Comparative Endocrinology 250:15-20

Martin, L.B., Kilvitis, H., Brace, A., Haussmann, M., Mutati, A., Fasanello, V., O'Brien, S. and D.R. Ardia. 2017. Collateral damage, more than resource costs of immunity, tracks range expansion of an introduced bird. Journal of Experimental Biology 220: 2228-2235.

Rivers, J., Newberry, G., Schwarz, C and D.R. Ardia. 2017. Success despite the stress: violet-green swallows increase glucocorticoids and maintain reproductive output following experimental increases in flight costs. Functional Ecology 31:234-244.

Akresh, M.E., D.R. Ardia, and D.I. King. 2017. Effect of nest characteristics on thermal properties, clutch size, and reproductive performance for an open-cup nesting songbird. Avian Biology Research 10:107-118.

Ardia, D.R. 2017. Nests, Eggs, and Incubation: New Ideas about Avian Reproduction [Book review] Auk: Ornithological Advances 134:262-263.

Lathi DC and DR Ardia. 2016. Shedding light on bird egg color: pigment as parasol and the hot car effect. American Naturalist. 187: 547-63.

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

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

Martin LB, DH Hawley, and DR Ardia. 2014. A New Division of Ecoimmunology and Disease Ecology. Integrative and Comparative Biology 54: 338-339.

Winkler DW, KM Ringelman, … DR Ardia. 2014. Latitudinal variation in clutch size–lay date regressions in Tachycineta swallows: effects of food supply or demography? Ecography 37: 670-678.

Windsor! RE, JE Fegely!, and DR Ardia. 2013. Factors affecting the thermal insulation properties of tree swallow nests. Journal of Avian Biology 44: 305-310.

Butler M, ZR Stahlschmidt, DR Ardia, S Davies, L Guilette, N Johnson, S McCormick, KJ McGraw, 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 DR. 2013. The effects of nestbox thermal environment on fledging success and haematocrit in Tree Swallows. Avian Biology Research 6: 1-6.

Ardia DR, JE Gantz!, BC 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, DR Ardia, D Caceres, CB Cooper, EE Iñigo-Elias, J Molina, N Taylor, & DW Winkler. 2012. Reproductive biology of a narrowly endemic Tachycineta swallow in dry, seasonal forest in coastal Peru. Ornitología Neotropical 23: 95-112.

Ardia DR, 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 (LB Martin, DR Ardia, DM Hawley, editors.) 21: 61-73.

Martin LB, DM Hawley, and DR Ardia. 2011. An introduction to ecological immunology. Functional Ecology 21: 1-4.

Cooper CB, MA Voss, DR Ardia, SB Austin, WD Robinson. 2011. Light increases the rate of embryonic development: implications for latitudinal trends in incubation period. Functional Ecology 25: 769-776.

Ardia DR, JH Peréz!, and ED 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 DR, DR 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 MM, F Adriaensen, DR 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! LE, DR Ardia and JEC Flux. 2009. Breeding synchrony through social stimulation in a spatially-segregated population of European starling (Sturnus vulgaris). Animal Behaviour 78: 671-675.

Morrison! E, DR Ardia and ED 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 LM, CA Makarewich, A Coulon, DR Ardia, IJ Lovette and DW Winkler. 2009. Subtle edge-of-range genetic structuring in transcontinentally distributed North American Tree Swallows. Condor 111: 470-478.

Ardia DR, JH Peréz!, EK Chad!, MA Voss, and ED 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! JH, DR Ardia, EK Chad!, and ED Clotfelter. 2008. Experimental heating reveals nest temperature affects nestling condition in tree swallows (Tachycineta bicolor). Biology Letters 4: 468-471. Authors contributed equally.

Ardia DR and KA Schat. 2008. Ecoimmunology. In: The Immunology of Birds (editors F Davison, B Kaspers and KA Schat). Elsevier.

McGraw KJ and DR Ardia. 2007. Do carotenoids buffer testosterone-induced immunosuppression? An experimental test in a colorful song bird. Biology Letters 3: 375-378.

Clotfelter ED, DR Ardia, & KJ McGraw. 2007. Red fish, blue fish: trade-offs between pigmentation and immunity in Betta splendens. Behavioral Ecology 18: 1139-1145. Authors contributed equally.

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

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

Ardia DR and EB Rice. 2006. Variation in heritability of immune function in the tree swallow. Evolutionary Ecology. 20: 491-500.

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

Ardia DR, JH Peréz!, and ED Clotfelter. 2006. Nest box orientation affects internal temperature and nest site selection by Tree Swallows. Journal of Field Ornithology.

Ardia DR, MW Wasson, and DW 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 DR and ED 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 DR, CB Cooper, and AA 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 DR. 2006. Glycated hemoglobin and albumin reflect nestling growth and condition in American kestrels. Comparative Biochemistry and Physiology, Part A 143: 62-66.  

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

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

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

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

McGraw KJ and DR Ardia. 2005. Sex differences in carotenoid status and immune performance in zebra finches. Evolutionary Ecology Research 7: 251-262.

McGraw KJ and DR Ardia. 2004. Immunoregulatory activity of different dietary carotenoids in zebra finches. Chemoecology 14: 21-29.  

Ardia DR, KA Schat, and DW Winkler. 2003.  Reproductive effort decreases long-term immunity in breeding tree swallows. Proceedings of the Royal Society of London B 270: 1679-1683.  

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

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

Ardia DR and KL Bildstein. 2001. Sex-related differences in habitat use in wintering American Kestrels. Auk 118: 746-749.  

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

Ardia DR and KL Bildstein. 1997. Sex-related differences in habitat selection in wintering American kestrels (Falco sparverius ). Animal Behaviour 53: 1305-1311.