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Kraig Adler

Professor Emeritus

Kraig Adler

Seeley G Mudd Bio Science Wing, Room 342


  • Neurobiology and Behavior


My major research program over the last two decades has been the sensory basis of long-distance orientation and navigation. These studies have been conducted both in field situations (Ithaca and Arizona) and in the laboratory (Mudd Hall and Liddell Field Station), depending upon the nature of the questions. Most of our work has focused on amphibians and reptiles as model systems, since these organisms are often locally abundant and easy to manipulate, but we have also conducted experiments with humans. We were the first to discover that amphibians and reptiles can detect and use skylight polarization patterns for orientation (polarotaxis) and that the critical receptors for this cue are extraocular (pineal and related structures). We were also first to show that amphibians can detect and use the earth's magnetic field for orientation, although the critical receptor remains unknown.

Other studies have dealt with the influence of slope on orientation, the ability to learn a new homesite, the importance of individual variability, time-dependent orientation mechanisms, the role of environmental stress in orientation, and the function of melatonin in re-setting internal clocks which are then used for orientation. Recently, we were able to demonstrate for the first time that amphibians are capable of true navigation. Most of these studies are conducted using the red-spotted newt, an abundant species in the Ithaca area that has both aquatic and terrestrial life stages. Our studies on human orientation have dealt primarily with tests for the possible use of magnetic cues, which we failed to find despite an earlier claim. Recently, I have initiated a comparative study of electroreception in salamanders. In addition to these studies on behavior, I have also been involved in studies on the evolution and systematics of amphibians and reptiles. Most of this work is centered on the herpetofaunas of China and Central America.


  • Contributions to the History of Herpetology, vol. 2. Contr. Herpetol., Saint Louis, vol. 21, 2007, 389 pages.
  • The evolution of amphibian systematics.  In Harold Heatwole (ed.), Amphibian Biology, vol. 7.  Surrey Beatty, Chipping Norton (Australia), pp. 2407–2435 [W. E. Duellman and K. Adler].
  • The Development of Systematic Reviews of the Turtles of the World.  Vert. Zool., Dresden, 57(2): 139–148, 2007.
  • Foreword, p. x, in M. S. Khan, Amphibians and Reptiles of Pakistan. Krieger Publ. Co., Malabar, Florida, 2006.
  • An American’s Appreciation of Professor Ding Hanbo, p. 26-28 In Y.-Z. Ding (ed.), A Memorial to Professor Ding Hanbo. Fujian Univ., Fuzhou, PRC, 2005.
  • Charles Boewe (ed.) Profiles of Rafinesque. Isis, 96: 285-286, 2005 (book review).
  • America’s first herpetological expedition: William Bartram’s travels in southeastern United States (1773–1777).  In A. Bauer (ed.), Herpetological Voyages. Bonner Zool. Beiträge, 52: 275-295, 2004.
  • In Memoriam: Roger Conant (1909–2003) with reflections by some of Roger’s colleagues.  Herpetol. Rev., 35(2): 101–107, 2004.
  • Henri Clerét Seibert (1916–2003). Herpetol. Rev., 35(1): 3, 2004.
  • Cynthia Kagarise Sherman (1950–2002).  Herpetol. Rev., 34(4):289, 2003.
  • On the dates of publication and correct citation of Olivier’s Voyage dans l’Empire Othoman and its herpetological content.  International Society for the History of Bibliography of Herpetology, Newsletter bulletin, 4(2): 7–18, 2003.  (with Aaron M. Bauer).
  • An American’s Appreciation of Professor Ding Hanbo.  In Y.-D. Zhang (ed.) A memorial to Professor Ding Hanbo.  Fujian Normal University, Fujian, Fuzhou, PRC, 2003.
  • Amphibians and humans, p. 51–55.  In M. Hutchins, W. E. Duellman, and N. Schlager (eds.), Grzimek’s Animal Life Encyclopedia, 2nd ed., vol. 6,  Amphibians.  Gale Group, div. Thomson Learning, Farmington Hills, Michigan, 2003.
  • Salamander classification and reproductive biology.  An historical overview, p. 1­–29.  In D. M. Sever (ed.), Reproductive Biology and Phylogeny of Urodela (general series: Reproductive Biology and Phylogeny, ed. By B. G. M. Jamieson).  Science Publishers, Enfield, New Hampshire and Plymouth, UK, 2003.
  • The publishing history of Pitman’s “Snakes of Uganda.”  Newsl. Bull. Internatl. Soc. Hist. Biblio. Herpetol., 4(1): 5–14, 2003.
  • Illustrating amphibians and reptiles, p. 385–400.  In E. Hodges (ed.), Guild Handbook of Scientific Illustration, second edition, 2003 [with David M. Dennis].
  • Classification and taxonomy.  Evolutionary relationships of amphibians and reptiles, p. 20–21. In Tim Halliday and Kraig Adler (eds.), The New Encyclopedia of reptiles and Amphibians.  Oxford University Press, Oxford, UK. 2002.
  • Amphibians; A key amphibian event; Kaleidoscopic adaptation; Conscientious parents, pp. 10–19, 22–29.  In Tim Halliday and Kraig Adler (eds.), The New Encyclopedia of reptiles and Amphibians.  Oxford University Press, Oxford, UK. 2002.
  • Encyclopaedia of Reptiles and Amphibians, second edition.  Oxford Univ. Press, Oxford, 2002. [with Tim R. Halliday].
  • Revised checklist of Chinese Amphibia & Reptilia.  Sichuan Jour. Zool., 19(3): 196–207, 2000 [with Er-mi Zhao, Hsueh-Wen Chang, and Hui Zhao].
  • Myth versus reality: the challenges of illustrating amphibians and reptiles, p. 5–15.  In S. Haines, Slithy Toves: Illustrated Classic Herpetological Books at the Univ. Kansas in Pictures and Conversations.  Soc. Study Amphib. Rept., Ithaca, NY, Contrib. Herpetol. 16, 2000.
  • Russell’s viper: Daboia russelii, not Daboia russellii, due to Classical Latin rules.  Hamadryad (Madras), 25(3): 83–85, 2000 [and H.M. Smith, S.H. Prince, P. David, and D. Chiszar].Revised checklist of Chinese Amphibia & Reptilia, p. 215–238.  In E.-M. Zhao, H.-W. Chang and H. Zhao (eds.), Taxonomic Bibliography of Chinese Amphibia and Reptilia, including Karyological Literature.  Fu-Wen Publ. Co., Kaohsiung, Taiwan, 2000 [with E. Zhao, H.W. Chang, and H. Zhao].
  • Foreword to the reproduction.  In Thomas Bell, A Monograph of the Testudinata.  Octavo Editions, Oakland, California, on CD.  1999.
  • Firefly toxicosis in lizards.  Jour. Chem. Ecol., 25(9): 1981–1986, 1999 [with M. Knight, R. Glor, S.R. Smedley, A. González, and T. Eisner].
  • Nikolai Nikolaevich Szczerbak (31 October 1927–27 January 1998).  Gekko, 1(1): 36-37, 1999.
  • Deciphering Latin dates and place names on the title pages of old herpetological books.  Internat. Soc. Hist. Biblio. Herpetol., 1(2): 34–35, 2000 [date on issue “1999”].
  • Organizing scientific meetings, by August Epple (book review).  Amer. Zool., 38(2): 410.  1998.
  • Biology as history, by G. Pinna and M.T. Ghiselin (eds.) (review).  Isis, 89(3): 584–585.  1998.
  • Herpetology vs. herpetoculture.  Cooperation or competition?  Reptiles, 5(3): 72-79, 1997 (J.B. Murphy, K. Adler, and J.T. Collins).
  • The salamanders of Guerrero, Mexico, with descriptions of five new species of Pseudoeurycea (Caudata: Plethodontidae).  Occas. Pap. Nat. Hist. Mus. Univ. Kansas, 177: 1-28, map, 1996.
  • An annotated checklist of West Indian amphibians and reptiles, p. 51-94, plates 1-8.  In R. Powell and R.W. Henderson (eds.), Contributions to West Indian Herpetology.  Soc. Study Amphib. Rept., Contrib. Herpetol., 1996 (R. Powell, R.W. Henderson, K. Adler, H.A. Dundee).
  • The proper generic name for the Asian wolf snakes: Lycodon (Serpentes: Colubridae).  Sichuan Jour. Zool., 14: 74-75, 1995 (K. Adler and Zhao Er-mi).
  • Revised Catalogue of the Herpetological Collection in Turin University I. Amphibia, by E. Gavetti and F. Andreone (review).  Herpetol. Rev., 26: 162, 1995.True navigation by an amphibian.  Anim. Behav., 50: 855-858, 1995 (J.B. Phillips, K. Adler, S.C. Borland).
  • Captive Management and Conservation of Amphibians and Reptiles.  Soc. Study Amphib. Reptiles, Contrib. Herpetol., 11:408pp., frontispiece, 1994 (J.B. Murphy, K. Adler, and J.T. Collins, eds.)
  • The remarkable career of Roger Conant, p. 16-23.  In J. B. Murphy, K. Adler, and J. T. Collins (eds.), Captive Management and Conservation of Amphibians and Reptiles.  Soc. Study Amphib. Reptiles, Contrib. Herpetol. 11, 1994.
  • Snake Life History in Kyoto, by Hajime Fukada (review).  Herpetol. Rev., 24: 113-114, 1993.Herpetology of China.  Soc. Study. Amphib. Reptiles, Oxford, Ohio, Contrib. Herpetol., 10: 522 pages, frontispiece, 48 plates, folding map, 1993 (Zhao Ermi and K. Adler).
  • Herpetological investigations in China by foreign naturalists, p. 331.  In E.-M. Zhao, B.-H. Chen, and T. J. Papenfuss (eds.), Proceedings of the 1st Asian Herpetological Meeting, Huangshan. China Forestry Press, Beijing, 1993.
  • First records of the pipe snake (Cylindrophis) in China.  Asiatic Herpetol. Res., 4: 37-41, 1992.  (K. Adler, Zhao Ermi, and Ilya S. Darevsky).
  • Handbook to Middle East Amphibians and Reptiles.  Soc. Study Amphib. Reptiles, Oxford, Ohio, Contr. Herpetol., 8: vii, 252 pages, 33 plates, 1992. (A. E. Leviton, S. C. Anderson, K. Adler, and S. A. Minton).
  • Herpetology: Current Research on the Biology of Amphibians and Reptiles.  Proceedings of the First World Congress of Herpetology.  Soc. Study Amphib. Reptiles, Oxford, Ohio, Contrib. Herpetol., 9: v, 245 pages, 1992 (K. Adler, editor).
  • Studies on hynobiid salamanders, with description of a new genus.  Asiatic Herpetol. Res., 3: 37-45, 1990 (K. Adler and Zhao Ermi).
  • Cheng-chao Liu from an American perspective, p. 26-30.  In E. Zhao (ed.), From Water Onto Land [commemorating 90th birthday of C.-C. Liu].  China Forestry Press, Beijing, 1990.
  • Atlas Anima, volume 3, Amphibia and Reptilia, by Hiroshi Aramata. (book review). Herpetol. Rev., 22: 32, 1990.
  • Foreword: Dedication to Charles F. Walker, p. xv-xviii. In R. A. Pfingsten and F. L. Downs (eds.), Salamanders of Ohio. Bull. Ohio Biol. Surv., new ser., vol. 7, no. 2, Columbus, 1989.
  • [Hynobius chinensis, its 100th Anniversary: A Redescription] (in Chinese). Sichuan Jour. Zool., 8(2): 18-20, 1plate. (K. Adler and Zhao Ermi).
  • [Cylindrophis ruffus, a New Family Record for Chinese Snakes](in Chinese). Sichuan Jour. Zool., 8(2): 26, 1989. (K. Adler and Zhao Ermi).
  • Contributions to the History of Herpetology.  Soc. Study Amphib. Rept., Contr. Herpetol., 5: 1-202, 1989 (K. Adler, editor).
  • Herpetologists of the past, p. 5-141.  In Kraig Adler (ed.), Contributions to the History of Herpetology. 1989.
  • Orientation mechanisms in a desert lizard, Uma notata, p. 583-588. In Z. Rôcek (ed.), Studies in Herpetology, Proc. Europ. Herpetol. Meet., Charles Univ., Prague, 1988.
  • Chapter 17: Illustrating amphibians and reptiles, p. 321-337, 553-554. In Elaine R. S. Hodges (ed.), Guild Handbook of Scientific Illustration. Van Nostrand Reinholt, New York, 576 p., 1988. (David M. Dennis and K. Adler).
  • The Encyclopaedia of Reptiles and Amphibians.  Unwin Animal Library, vol. V.  (T. R. Halliday and K. Adler, eds.), George Allen and Unwin, London and Sydney, xvi, 144p., 1986 (Foreign language editions in Japanese, Korean, and most European languages). 
  • Amphibians: Introduction; Metamorphosis; Colors; Parental Care.  In T. R. Halliday and K. Adler (eds.), The Encyclopaedia of Reptiles and Amphibians, p. 2-15, 1986.
  • Polarotaxis in desert lizards (Uma notata).  Jour. Comp. Physiol., A, 156: 547-552, 1985.  (K. Adler and J. B. Phillips).
  • Human homing orientation: a critique and alternative hypotheses, p. 573-593.  In J. L. Kirschvink et al. (eds.), Magnetite Biomineralization and Magnetoreception in Organisms.  Plenum Press, New York, 1985.  (K. Adler and C. Pelkie).
  • Salamanders of the genus Bolitoglossa from the Sierra Madra del Sur of southern Mexico.  Jour. Herpetol., 17: 295-307, 1983  (T. J. Papenfuss, D. B. Wake, and K. Adler).
  • Herpetological activities in the People's Republic of China.  Herpetol. Rev., 14: 107, 1983.
  • Sensory aspects of amphibian orientation.  Vertebrata Hungar., 21: 7-18, 1982.
  • Toad orientation: variability of response and its relationship to individuality and environmental parameters.  Jour. Comp. Physiol., A, 144: 45-51, 1981   (K. Adler and D. H. Taylor).
  • Individuality in the use of orientation cues by green frogs.  Anim. Behav., 28: 413-425, 1980.
  • Melatonin and thyroxine: influence on compass orientation in salamanders.  Jour. Comp. Physiol., A, 136: 235-241, 1980  (K. Adler and D. H. Taylor).
  • The mysteries of amphibian migration (review).  Bull. N.Y. Herpetol. Soc., 15: 2-6, 1980.
  • A brief history of herpetology in North America before 1900.  Soc. Study Amphib. Rept., Herpetol. Circ., 8: 1-40, 1979.
  • Toad tadpoles associate preferentially with siblings.   Nature, 282: 611-613, 1979  (B. Waldman and K. Adler).
  • The pineal body: site of extraocular perception of celestial cues for orientation in the tiger salamander.  Jour. Comp. Physiol., A, 124: 357-361, 1978 (K. Adler and D. H. Taylor).
  • Directional and discriminatory responses of salamanders to weak magnetic fields, p. 325-333.  In K. Schmidt-Koenig and W. T. Keeton (eds.), Animal Migration, Navigation, and Homing.  Springer-Verlag, Berlin, 1978  (J. B. Phillips and K. Adler).
  • Extraocular photoreception in amphibians.  Photochem. Photobiol., 23: 275-298, 1976.
  • Territorial and vocal behavior in neotropical frogs of the genus Centrolenella.  Herpetologica, 30: 75-78, 1974  (R. W. McDiarmid and K. Adler).
  • The function of extraocular photoreception in amphibian behavior.  Amer. Soc. Photobiol., Vancouver, pp. 126-127, 1974 (abstract).
  • Understanding the science of biology, p. 10-27.  In K. Adler, Readings and Exercises for Introductory Biology.  Cornell University, Ithaca, New York, 264 p.
  • Careers in biological systematics.  Joint Publ. System. Zool.-Amer. Soc. Plant Taxon.  16 pages, 1973  (T. J. Crovello and K. Adler).
  • Two methods for using period length to study rhythmic phenomena.  Jour. Comp. Physiol., 83: 63-71, 1973 (S. A. Binkley, K. Adler and D. H. Taylor).
  • Spatial orientation by salamanders using plane-polarized light.  Science, 181: 285-287, 1973 (D. H. Taylor and K. Adler).
  • Extraocular perception of polarized light by orienting salamanders.  Jour. Comp. Physiol., 87: 203-212, 1973 (K. Adler and D. H. Taylor).
  • Shoreline imprinting in the tiger salamander (Ambystoma tigrinum).  Herpetol. Rev., 4: 128, 1972 (abstract; D. H. Taylor and K. Adler).
  • Perception of the plane of polarized light and its use by orienting salamanders.  Proc. Indiana Acad. Sci., 81: 339, 1972 (abstract; K. Adler and D. H. Taylor).
  • New tree frogs of the genus Hyla from the cloud forests of western Guerrero, Mexico.  Occas. Pap. Mus. Nat. Hist. Univ. Kansas, (7): 1-19, pls. 1-2, 1972  (K. Adler and D. M. Dennis).
  • Function of extraoptic photoreceptors in amphibian behavior: I. Manipulation of the biological clock.  Proc. Indiana Acad. Sci., 80: 486, 1971 (abstract).
  • Pineal end organ: role in extraoptic entrainment of circadian locomotor rhythm in frogs,  p. 342-350.  In M. Menaker (ed.), Biochronometry.  Publ. U.S. Natl. Acad. Sci., Washington, 1971.
  • Polarized light: detection via extraoptic receptor and its use by orienting salamanders.  Herpetol. Rev., 3: 105, 1971 (abstract; K. Adler and D. H. Taylor).
  • How do trained frogs find a goal box?  Jour. Herpetol., 3: 198, 1970 (abstract).
  • The influence of prehistoric man on the distribution of the box turtle.  Ann. Carnegie Mus., 41: 263-280, 1970.
  • The role of extraoptic photoreceptors in amphibian rhythms and orientation: a review.  Jour. Herpetol., 4: 99-112, 1970.
  • Extraoptic phase-shifting of circadian rhythm in salamanders. Science, 164: 1290-1292, 1969.
  • Environmental control of locomotor activity in a salamander (Plethodon glutinosus).  Dissert. Abst., 30: 882-B, 1969  (abstract).
  • Extraoptic entrainment of circadian rhythm in salamanders (Plethodon glutinosus). Jour. Herpetol., 2: 176 (abstract).
  • Synonymy of the Pliocene turtles Pseudemys hilli Cope and Chrysemys limnodytes Galbreath. Jour. Herpetol., 1: 32-38, 1968.
  • Turtles from archeological sites in the Great Lakes region.  Michigan Archaeol., 14: 147-163, 1968.
  • Three new frogs of the genus Hyla from the Sierra Madre del Sur of Mexico.  Occas. Pap. Mus. Zool. Univ. Michigan, (642): 1-18, 1965.
  • An ecological study of the snake Natrix erythrogaster, with remarks on its post-glacial dispersal.  Jour. Ohio Herpetol. Soc., 4: 61-62, 1963  (abstract).
  • Plethodon longicrus, a new salamander (Amphibia: Plethodontidae) from North Carolina.  Spec. Publ., Ohio Herpetol. Soc., (4): 1-14, pls. I-II, 1962  (K. Adler and D. M. Dennis).
  • Egg-laying behavior in the spotted turtle, Clemmys guttata (Schneider).  Ohio Jour. Sci., 61(3): 180-182, 1961.