Tom Schultz

Tom Schultz

Professor Emeritus
Position Type
- Present
B.A., University of Chicago; Ph.D., University of Texas

Learning & Teaching

  • First Year Studies: Animal Talk
  • Introduction to the Science of Biology
  • Ecology & Evolution
  • Biology of Insects
  • Animal Behavior
  • Senior Research
Academic Positions
  • Professor of Biology, Department of Biology at Denison University, 2000 - present
  • Chair, Department of Biology at Denison University, 1998-2001
  • Director, Denison University Biological Reserve, 1994-2003
  • Visiting Lecturer, Department of Biology at Yale University, 1988-1990
  • Post-doctoral Research Associate, Department of Zoology at Arizona State University, 1985-1987
  • Lecturer, Department of Zoology at The University of Texas at Austin, 1983-1984


I am interested in how animals, especially insects, produce specific colorations and use them in intraspecific communication or anti-predator defense. I use electron microscopy and histochemistry to identify the biophotonic structures that produce colors and optical spectrometry to determine their contrast with visual backgrounds to function as conspicuous signals, or blend with background noise and produce camouflage. Currently, I am exploring the function of intra- and intersexual signals of damselflies and how they are produced in the integument. I am also returning to previous work focused on how habitat preferences and tradeoffs shape the evolution of color patterns in tiger beetles and other cryptic insects.

Communication signals have both content, information intended to change the behavior of a receiver, and structural properties that determine how the information is transmitted from sender to receiver. My research concerns how the structural properties of visual signals evolve and are shaped by the ecology and environment of animals, whether they are detected by mates, rivals, or predators.

The males of many animals exhibit conspicuous colors that attract mates, advertise fitness, and mitigate conflict between rivals. However, visual signals may be intercepted inadvertently by other species and intentionally by predators. Color patterns evolve in response to some or all of these selection pressures within the limits of an animal's ability to see and produce color. Damselflies are an excellent group of organisms for studying these tensions, as they are highly visual, sexually dimorphic, and vulnerable to visual predators. Some species exhibit courtship displays, territorial behavior, or occur in assemblages of closely related species where signals may be confused. The learning and behavioral repertoires of damselflies are limited and their visual environments are relatively simple to characterize. These qualities make it possible to focus on the properties of color signals that make them more or less easy to detect, and their role in transmitting information.

I am also especially interested in the function of structural colors in insects, which have unique optical properties that may be tuned to certain viewing conditions. Insects produce structural colors through ultrastructural modifications of their exoskeleton. In combination with pigments, structural colors have the capacity to produce a wide variety of adaptive color patterns ranging from the flashing iridescence of some damselflies to the camouflage of tiger beetles.


I find it very satisfying and fun to explore the natural world, but my real passion is for the learning process. The challenge of being exposed to a new idea, questioning it, evaluating it, and even testing it, is very fulfilling for me. I especially enjoy making connections between seemingly disparate ideas or concepts in different disciplines (an important ability in a time when boundaries between scientific disciplines are becoming blurred). The courses I teach all involve integrating different approaches and levels of organization. Sometimes I think I have the perfect job in that I am paid to learn new things and share them with students, and to help them to develop a "Swiss Army Knife" of critical thinking skills. With these skills, they can become good leaders and thoughtful citizens in any field, and better able to face the uncertainty of the future.



* Student co-authors

  • Schultz,T.D. & O. M. Fincke. 2009. Structural colors create a flashing cue for sexual recognition and mate quality in a Neotropical giant damselfly . Functional Ecology. v. 23 p. 724-732
  • Seago, A., Brady, P., Vigneron, J-P. & Schultz, T.D.. 2009. Gold bugs and beyond: A review of iridescence and structural color mechanisms in beetles (Coleoptera) . Journal of the Royal Society Interface. v. 6 p. S165-S184
  • Schultz, T.D.. 2009. Diversity and habitats of a prairie assemblage of Odonata at Lostwood National Wildlife Refuge, North Dakota. Journal of Kansas Entomological Society. v. 82 p. 91-102
  • Schultz, T.D., C.N. Anderson*, & L. B. Symes*. 2008. The conspicuousness of colour cues in male pond damselflies depends on ambient light and visual system. Animal Behaviour. v. 76 p. 1357-1364
  • Fincke, O.M., A. Fargevielle, & T. D. Schultz. 2007. Lack of innate preference for morph and species identity in mate-searching Enallagma damselflies. Behavioral Ecology and Sociobiology. v. 61 p. 1121-1131
  • Fincke, O.M., R. Jodicke, D. Paulson, & T. D. Schultz. 2005. The frequency of female-specific color polymorphisms in Holarctic Odonata: why are male-like females typically the minority? . International Journal of Odonatology. v. 8 p. 183-212
  • Schultz, T. D. 2001. Tiger beetle defenses revisited: alternative defense strategies and colorations in two neotropical tiger beetles, Odontocheila nicaraguensis and Pseudoxycheila tarsalis (Carabidae: Cicindelinae). Coleopterists Bulletin. v. 55 p. 153-163
  • Schultz, T. D. & J. Puchalski *. 2001. Chemical Defenses in the Tiger Beetle Pseudoxycheila tarsalis Bates (Carabidae: Cicindelinae). Coleopterists Bulletin. v. 55 p. 164-166
  • Kirkton, S. D.* & T. D. Schultz. 2001. Age-specific behavior and habitat selection of adult male damselflies, Calopteryx maculata (Odonata: Calopterygidae). Journal of Insect Behavior. v. 14 no. 4 p. 545-556
  • Schultz, T. D. 1998. The utilization of patchy thermal microhabitats by the ectothermic insect predator, Cicindela sexguttata. Ecological Entomology. v. 23 p. 444-450
  • Knisley, C. B. & T. D. Schultz. 1997. The Biology of Tiger Beetles and a Guide to the Species of the South Atlantic States. Virginia Museum of Natural History, Martinsville, VA. p. 236 pp
  • Schultz, T. D., M. Quinlan & N. F. Hadley. 1992. Preferred body temperature, metabolic physiology, and water balance of adult Cicindela longilabris: a comparison of populations from boreal habitats and climatic refugia. Physiological Zoology. v. 65 p. 226-242
  • Hadley, N. F., A. Savill, & T. D. Schultz. 1992. Coloration and its thermal consequences in the New Zealand tiger beetle Neocicindela perhispida. J. Thermal Biology. v. 17 p. 55-61
  • Schultz, T. D. 1991. Tiger Hunt. Natural History. p. 38-44
  • Schultz, T. D.& G. Bernard. 1990. Pointillistic mixing of interference colors in cryptic tiger beetles. Nature. v. 337 p. 72-73
  • Hadley, N. F., T. D. Schultz, & A. C. Savill. 1988. Spectral reflectances of three subspecies of the tiger beetle Neocicindela perhispida: correlations with their respective habitat substrates. New Zealand J. of Zoology. v. 15 p. 343-346
  • Schultz, T. D. & N. F. Hadley. 1987. Microhabitat segregation and physiological differences in co-occurring tiger beetle species, Cicindela oregona and Cicindela tranquebarica. Oecologia. v. 73 p. 363-370
  • Schultz, T. D. & N. F. Hadley. 1987. Structural colors of tiger beetles and their role in heat transfer through the integument. Physiological Zoology. v. 60 p. 737-745
  • Schultz, T. D. 1986. The role of structural colors in predator avoidance by tiger beetles of the genus Cicindela. Bulletin of the Entomological Society of America. v. 32 p. 142-146
  • Schultz, T. D. & M. A. Rankin. 1985. The ultrastructure of epicuticular interference reflectors of tiger beetles (Cicindela). J. Experimental Biology. v. 117 p. 88-110
  • Schultz, T. D. & M. A. Rankin. 1985. Developmental changes in the interference reflectors and colorations of tiger beetles (Cicindela). J. Experimental Biology. v. 117 p. 111-118


Professional Memberships
  • Animal Behavior Society
  • Coleopterist’s Society
  • Dragonfly Society of America
  • Ecological Society of America
  • International Society of Behavioral Ecology
  • Ohio Odonata Society
  • Society for the Study of Evolution


Student Collaborations

One of the best aspects of being at a small college is the opportunity for close faculty-student collaboration on independent research. I have had the pleasure of advising a number of undergraduate students who have conducted a variety of outstanding research projects, many of which have been presented at national scientific meetings. As a research advisor, I involve students in my studies of insect behavioral ecology or enlist students interested in conservation biology in conducting inventories and monitoring studies at the Bio Reserve and other sites in Licking County. In almost all cases, these projects require a summer of field work prior to the senior year. In the past, my summer research students have been supported with Anderson Research Fellowships or stipends provided through the Denison University Research Foundation.

Research Projects Supervised

  • Brindle, A. 2008. Differing social environments between primate populations may generate false positive evidence of cultural variation. *
  • Gorsich, E. 2008. Ommochrome signaling in male Enallagma damselflies: can long wavelength coloration be correlated with territorial behavior? *
  • Bring, B. 2007. A study of odonate community development, habitat preferences, and colonization among ponds and artificial wetlands at Dawes Arboretum.
  • Horn, J. 2007. Suburban habitat fragmentation: effects on migratory and residential songbirds in Central Ohio. *
  • Symes, L. 2007. Polychromatism and sex identity signals in the damselfly genus Enallagma. *
  • Symes, L. 2006. Polychromatism in the damselfly Enallagma civile and an assessment of the Male-Mimicry Hypothesis. (Poster presented at 2006 Meeting of the Ecological Society of America).
  • Hughes, D. 2005. The interspecific roosting behaviors of the Turkey Vulture (Cathartes aura) and Black Vulture (Corapgys atratus).
  • Dunlevy, J. 2003. Investigations of summer bird residents at five sites within Licking County.
  • Anderson, C. 2002. Enallagma damselfly colors as visual signals in relation to ambient light and visual backgrounds. * (Poster presented at 2002 Meeting of the Ecological Society of America).
  • Bucci, L. 2002. Effects of vegetation and landscape on butterfly diversity and abundance. *
  • Clark, E. 2002. Corellations of odonate diversity with lotic habitat characteristics.
  • Menninger, H. 2000. Examining the ecology of an indicator taxon: damselfly species diversity and the role of habitat heterogeneity. * (Paper presented at 2000 Meeting of Ohio Odonata Society).
  • Hauck, A. 1999. Correlation between male damselfly colorations and the light environments of courtship arenas. *
  • Menninger, H. 1999. Initiation of a long-term monitoring program for Odonata at the Denison University Biological Reserve. (Poster presented at 1999 Meeting of the Entomological Society of America).
  • Kirkton, S. 1997. Babes in the woods: age-specific dispersal in the territorial damselfly, Calopteryx maculata. * (Paper presented at 1997 Meeting of the Ecological Society of America).
  • Scheub, C. 1997. Diversity and abundance of the Papilionoidea at the Denison University Biological Reserve.* (Poster presented at 1996 Meeting of the Entomological Society of America).
  • Kirkton, S. 1996. Why do male ebony jeweling damselflies (Calopteryx maculata) aggregate far from territorial breeding sites? (Poster presented at 1996 Meeting of the Entomological Society of America).
  • Casey, J. 1996. The significance of interference colors and visual communication in Phidippus audax, the daring jumping spider. *
  • Godfrey, P. 1996. Optimal site choice and foraging posture of the ambush predator Phymata fasciatus. *
  • Forbes, B. 1995. Substrate matching and cryptic defenses in the toad bug Gelastocoris oculatus. *
  • Stocker, E. 1995. Spectral sensitivity of the visual system in the praying mantis, Sphrodomantis lineola. (Paper presented at the 1994 Meeting of the Entomological Society of America).
  • Angalich, L. 1994. Correlation between conspicuousness and escape flight behavior among species of tiger beetles. * (Poster presented at the 1993 Meeting of the Entomological Society of America).
  • Gallo, T. 1994. A review of the history and efficacy of the Endangered Species Act and prospect for its renewal. *
  • Price, C. 1994. A comparative study of insect colonization and decomposition of pig carrion in central Ohio. * (Paper presented at the 1994 Meeting of the Ohio Academy of Science).
  • Puchalski, J. 1994. Comparative chemical analysis of the defensive secretions of tiger beetles (Cicindelidae) and leave beetles (Chrysomelidae). *
  • Van Antwerp, A. 1994. Light gap utilization and behavioral thermoregulation by the green forest tiger beetle, Cicindela sexguttata. * (Poster presented at the 1993 Meeting of the Entomological Society of America).

* denotes Honors Project

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