Timothy C Roth Associate Professor of Psychology




2021-                 Program Chair,  Biological Foundations of Behavior (Neuroscience and Animal Behavior)

2018-                 Associate Professor, Franklin and Marshall College

2012-2018      Assistant Professor, Franklin and Marshall College

2011-2012      Visiting Assistant Professor, Kenyon College

2007-2011      Postdoctoral Research Associate, University of Nevada, Reno

2006-2007     Postdoctoral Research Associate, Indiana State University



2006     Doctor of Philosophy. Indiana State University. Terre Haute, IN. Phi Kappa Phi

1999     Master of Science. Missouri State University. Springfield, MO.

1997     Bachelor of Arts, Magna Cum Laude. Cornell College. Mt. Vernon, IA. Phi Beta Kappa, Beta Beta Beta 



The overarching goal of my research is to understand the processes by which natural selection influences the use of space in animals. I integrate the study of neurobiology, animal behavior, and ecology to ask questions ranging from the neurological mechanisms of memory and behavior, to the large-scale effects of movement on avian communities, to the evolution of cognitive spatial processes.




*Denotes undergraduate authors

Johnsson, R.D. , F. Connelly, A.L. Vyssotski, T.C. Roth, and J.A. Lesku. 2021. Homeostatic regulation of NREM sleep, but not REM sleep, in Australian magpies. SLEEP

Roth, A.D.*, A.R. Krochmal, and T.C. Roth. 2021. Context-specific cue use in the Eastern Painted Turtle (Chrysemys picta) and its effects on decision making. Behaviour

Krochmal, A.R., T.C. Roth, and N. Simmons* 2021. My way is the highway: the role of plasticity in learning complex migration routes. Animal Behaviour.

Krochmal, A.R., T.C. Roth, and N. Simmons* 2021. Geomagnetic field does not appear to influence navigation in Eastern painted turtles. Ethology. 127:246-252.

Roth, T.C., M. Rosier*, A.R. Krochmal, and L. Clark*. 2020. A multi-trait, field-based examination of personality in a semi-aquatic turtle. Ethology.

Roth, T.C., and M. Thaker. 2020. “Predator-prey Interactions” In Oxford Bibliographies in Ecology. Ed.  D. Gibson. New York: Oxford University Press.

 Roth, T.C., A.R. Krochmal, and L.D. LaDage. 2020. Reptilian cognition: A more complex picture via integration of neurological mechanisms, behavioral constraints, and evolutionary context. BioEssasys. https://doi.org/10.1002/bies.201900033

Krochmal, A.R., T.J. LaDuc, A.J. Place, and T.C. Roth. 2019. Phylogenetic patterns in learning and decision making in pit vipers.  Animal Behaviour  https://www.sciencedirect.com/science/article/pii/S0003347218302951

Roth, T.C., and A.R. Krochmal. 2019. Turtles navigating during migration rely on the formation and recollection of spatial memories facilitated by M1 acetylcholine receptors. Proceeding of the Royal Society B . Featured in Scientific American.  https://blogs.scientificamerican.com/observations/how-a-painted-turtle-finds-its-way/

Roth, T.C., and A.R. Krochmal. 2019. Testudines Life History. In: Encyclopedia of Animal Cognition and Behavior, J. Vonk and T.K. Shackelford, eds. Springer. 

 McQuillan, M.A., T.C. Roth, A.V. Huynh, and A.M. Rice. 2018. Hybrid Songbirds are Deficient in Learning and Memory. Evolution 72:1155-1164. Featured in Audubon.

Vitale, A., R. Calisi, C. Carere, T. Carter, J.C. Ha, R. Hubrecht, D. Jennings, N. Metcalfe, A.G. Ophir, J.M. Ratcliffe, T.C. Roth, A. Smith, and L. Sneddon. 2018. Guidelines for the treatment of animals in behavioural research and teaching (revised). Animal Behaviour 135:I-X.

Roth, T.C., and A.R. Krochmal. 2018. Where the wild things are: a naturalistic approach to studying cognition in Chelonia. Box for: Wilkinson, A., and E. Glass. Cold-blooded cognition: How to get a tortoise out of its shell. In: Animal Cognition, F. Amici and N. Bueno eds., Cambridge University Press.

Krochmal, A.R., T.C. Roth, and H. O’Malley*. 2018. The role of learning in conservation: an assessment of soft releases for translocation. Animal Conservation 21:36-44.

 LaDage, L.D., T.C. Roth, C.J. Downs, B. Sinervo, and V.V. Pravosudov. 2017. Increased testosterone causes decreased medial cortical volume and neurogenesis in territorial side-blotched lizards (Uta stansburiana). Frontiers in Neuroscience. In press.

Krochmal, A.R., and T.C. Roth. 2017. From Comfort to Confidence: Modeling Science as a Process of Risk-Taking in the Classroom. In: Risk-Taking in Higher Education: The Importance of Negotiating Intellectual Challenge in the College Classroom. R. Kelty and B.A. Bunton, Eds. Rowman & Littlefield, MD.

Roth, T.C., A.R. Krochmal, W.B. Gerwig*, S. Rush*, N. Simmons*, J.D. Sullivan*, and K. Wachter*. 2016. Using pharmacological manipulation and high-precision radio telemetry to study the spatial cognition in free-ranging animals. Journal of Visual Experiments. 117: doi:10.3791/54790. 

Aulsebrook, A.E., T.M Jones, N.C. Rattenborg, T.C. Roth, and J.A. Lesku.  2016. Sleep ecophysiology: a case for neuroscience in ecology. Trends in Ecology and Evolution. In press.

Roth, T.C., K. Stocker*, and R.A. Mauck. 2016. Morphological changes in the hippocampus as a function of space use in birds. Developmental NeurobiologyIn press.

LaDage, L.D., T.C. Roth, B. Sinervo, and V.V. Pravosudov. 2016. Heritability and cortical volume in territorial and non-territorial side-blotched lizards (Uta stansburiana). Animal Behaviour. In press.

Roth, T.C., and A.R. Krochmal. 2016. Pharmacological Evidence is Consistent with a Prominent Role of Spatial Memory in Complex Navigation. Proceedings of the Royal Society B. 283:20152548. Cover article. 

Roth, T.C., and A.R. Krochmal. 2016. Cognition-centered conservation as a means of advancing integrative animal behavior. Current Opinion in Behavioral Sciences. In press.

Krochmal, A.R., T.C. Roth, S. Rush*, K.Wachter*. 2015. Turtles outsmart rapid environmental change: the role of cognition in navigation. Communicative and Integrative Biology. In press.

Rubenstein, D.R., H. Hofmann, E. Akcay, S. Alonzo, E. Archie, A. Beery, R. Calisi-Rodríguez, K. Carleton, B. Chow, J. Dubnau, C. Grozinger, E. Ketterson, A. Leifer, T. Linksvayer, M. MacManes, L. Martin, K. McGraw, L. McGraw, T. Mendelson, L. O'Connell, A. Ophir, L. Remage-Healey, S. Renn, T. Roth, J. Tung, and S. Woolley. 2015. New Frontiers for the Integrative Study of Animal Behavior. A white paper report to the National Science Foundation.

Pravosudov, V.V., T.C. Roth, L.D. LaDage, and C.A. Freas. 2015. Environmental influences on spatial memory and the hippocampus in food-caching chickadees. Comparative Cognition & Behavior Reviews. 10:25-43.

Croston, R., C.L. Branch, D.Y. Kozlovsky, T.C. Roth, L.D. LaDage, C.A. Freas, and V.V. Pravosudov. 2015. Climate-related variation in spatial memory and the hippocampus – what are the mechanisms driving population-level differences? Integrative and Comparative Biology 55:354-371.

Roth, T.C., A.R. Krochmal, and Z. Nemeth. 2015. Thinking about change: an integrative approach for examining cognition in a changing world. Integrative and Comparative Biology 55:347-353.

Roth, T.C., and A.R. Krochmal. 2015. The role of age-specific learning and experience for turtles navigating a changing landscape. Current Biology 25:333-337. DOI: http://dx.doi.org/10.1016/j.cub.2014.11.048; Cover article and cover photo. 

Kozlovsky, D.Y., S.L. Brown, C.L. Branch, T.C. Roth, and V.V. Pravosudov. 2014. Chickadees with bigger brains have smaller stomachs: a multi-population comparison. Brain Behavior and Evolution 84:172-180.

Freas, C; T.C. Roth. L.D. LaDage, and V.V. Pravosudov. 2013. Hippocampal neuron soma size is associated with population differences in winter climate severity in food-caching chickadees. Functional Ecology 27:1341-1349.

LaDage, L.D., M.V. Forney*, R.M. Maged*, T.C. Roth, B. Sinervo, and V.V. Pravosudov. 2013. Interaction between territoriality, spatial environment, and hippocampal neurogenesis in male side-blotched lizards, Uta stansburiana. Behavioral Neuroscience  127:555-565.

Pravosudov, V.V., and Roth, T.C. 2013. Food hoarding and the evolution of memory and the hippocampus. Annual Reviews in Ecology, Evolution and Systematics 44: 18.1-18.21.

Roth, T.C., D. Chevalier*, L. D. LaDage, and V.V. Pravosudov. 2013.  Variation in hippocampal glial cell numbers in different climates. Developmental Neurobiology 73:480-485.

Roth, T.C. 2013. “Predator-prey Interactions” In Oxford Bibliographies in Ecology. Ed.  D. Gibson. New York: Oxford University Press.

 Freas, C., L.D. LaDage, T. C. Roth, and V.V. Pravosudov. 2012. Elevation-related differences in memory and the hippocampus in Mountain Chickadees. Animal Behaviour. In press. 

Roth, T.C., C. Gallagher*, L.D. LaDage, and V.V. Pravosudov. 2012. Variation in brain regions associated with learning in contrasting climates. Brain Behavior and Evolution 79: 181-190.

Roth, T.C., L.D. LaDage, C. Freas, and V.V. Pravosudov. 2012. Variation in memory and the hippocampus across populations from different climates: a common garden approach. Proceedings of the Royal Society B 279: 402-410.

Roth, T.C., L.D. LaDage, and V.V. Pravosudov. 2012. Long-term memory in a food-caching parid. Animal Cognition 15:149-154.

Roth, T.C., L.D. LaDage, and V.V. Pravosudov. 2011. Variation in hippocampal morphology along an environmental gradient: controlling for the effects of latitude. Proceedings of the Royal Society B 278:2662-2667.

Chancellor, L.V.*, T.C. Roth, L.D. LaDage, and V.V. Pravosudov. 2011. The effect of environmental harshness on neurogenesis: a large-scale comparison. Developmental Neurobiology 71:246-252.

Rattenborg, N.C., D. Martinez-Gonzalez, T.C. Roth, and V.V. Pravosudov. 2011. Hippocampal memory consolidation during sleep: a comparison of mammals and birds. Biological Reviews 86:658-691.

Roth, T.C., L.D. LaDage, and V.V. Pravosudov. 2010. Learning capabilities enhanced in harsh environments: a common garden study. Proceedings of the Royal Society B 277:3187-3193. 

Pravosudov, V.V., T.C. Roth, and L.D. LaDage. 2010. Chickadees are selfish group members when it comes to food caching. Animal Behaviour 80:175-180.  

LaDage, L.D., T.C. Roth, and V.V. Pravosudov. 2010. Hippocampal neurogenesis is associated with migratory behavior in adult but not juvenile white-crowned sparrows (Zonotrichia leucophrys ssp.). Proceedings of the Royal Society B 278:138-143.

Roth, T.C., A. Brodin, T.V. Smulders, L.D. LaDage, and V.V. Pravosudov. 2010. Is bigger always better? A critical appraisal of the use of volumetric analysis in the study of the hippocampus. Philosophical Transactions of the Royal Society B 365:915-931. Erratum: 2010; 365:2423.

Roth, T.C., N.C. Rattenborg, and V.V. Pravosudov. 2010. The ecological relevance of sleep:the trade-off between sleep, memory, and energy conservation. Philosophical Transactions of the Royal Society B 365:945-959.

Fox, R.A., T.C. Roth, L.D. LaDage, and V.V. Pravosudov. 2010. Effects of social group composition and size on hippocampal formation morphology and neurogenesis in mountain chickadees (Poecile gambeli). Developmental Neurobiology 70:538-547.

Steury, T.D., J.E. McCarthy, T.C. Roth, S.L. Lima, and D.L. Murray. 2010. Evaluation of root-n bandwidth selectors for kernel density estimation. Journal of Wildlife Management 74:539-548.

LaDage, L.D., T.C. Roth, R.A. Fox, and V.V. Pravosudov. 2010. Ecologically-relevant spatial memory use modulates hippocampal neurogenesis. Proceedings of the Royal Society B 277:1071-1079.

Roth, T. C. and V. V. Pravosudov. 2009. Hippocampal volume and neuron numbers increase along a gradient of environmental harshness: a large-scale comparison. Proceedings of the Royal Society B  276:401-405

LaDage, L.D., T.C. Roth, and V.V. Pravosudov. 2009. Biases in measuring the brain: the trouble with the telencephalon. Brain, Behavior, and Evolution 73:253-258.Erratum: 2010; 76:70.

Lesku, J.A., T.C. Roth, N.C. Rattenborg, C.J. Amlaner, and S.L. Lima. 2009. History and future of comparative analyses in sleep research. Neuroscience and Biobehavioral Reviews 33:1024-1036.

oth, T. C. and V. V. Pravosudov. 2009. Tough times call for bigger brains. Communicative and Integrative Biology 2(3):1-3. Invited article.

Fox, R.A., L.D. LaDage, T.C. Roth, and V.V. Pravosudov. 2009. Temperament traits predict dominance status in mountain chickadees. Animal Behaviour 77:1441-1448.

LaDage, L.D., T.C. Roth, R.A. Fox, and V.V. Pravosudov. 2009. Flexible cue use in food-caching birds?  Animal Cognition 12:419-426.

LaDage, L.D., T.C. Roth, R.A. Fox, and V.V. Pravosudov. 2009. Effects of captivity and memory-based experiences on the hippocampus of mountain chickadees. Behavioral Neuroscience 123:284-291. 

Roth, T.C., J.G. Cox, and S.L. Lima. 2008. Can foraging birds assess predation risk by scent? Animal Behaviour 76:2021-2027. 

Roth, T.C., J.G. Cox, and S.L. Lima. 2008. The use and sharing of information about predation risk by flocks of wintering finches. Ethology 114:1218-1226.

Roth, T.C., W.E. Vetter, and S.L. Lima. 2008. Spatial ecology of wintering Accipiter hawks: home range, habitat use, and the influence of feeders. Condor 110:260-268.

Roth, T.C., and W.E. Vetter. 2008. The effect of feeder hotspots on the predictability and home range use of a small bird in winter. Ethology 114: 398-404.

Lesku, J.A., T.C. Roth, N.C. Rattenborg, C.J. Amlaner, and S.L. Lima. 2008. Phylogenetics and the correlates of mammalian sleep: a reappraisal. Sleep Medicine Reviews 12:229-244

Angilletta, M.J., T.C. Roth, R.S. Wilson, A.C. Niehaus, and P.L. Ribeiro. 2008. The fast and the fractalous: thermal sensitivity of speed and tortuosity in running ants. Functional Ecology 22:78-83.

Roth, T.C., and S.L. Lima. 2007. The predatory behavior of wintering Accipiter hawks: temporal patterns in activity of predators and prey. Oecologia 152 (1): 169-178. 

Roth, T.C., and S.L. Lima. 2007. Use of predictable prey hotspots by an avian predator: purposeful randomness? The American Naturalist 169 (2): 264-273. 

Roth, T.C., and S.L. Lima. 2006. Predatory behavior and diet of wintering male Cooper’s hawks in a rural habitat.  Journal of Raptor Research 40 (4): 287-290.

Roth, T.C., J.A. Lesku, C.J. Amlaner, and S.L. Lima. 2006. A phylogenetic analysis of the correlates of sleep in birds.  Journal of Sleep Research 15 (4): 395-402

Lesku, J.A., T.C. Roth, C.J. Amlaner, and S.L. Lima. 2006. A phylogenetic analysis of sleep architecture in mammals: the influence of anatomy, physiology, and ecology.  The American Naturalist 168 (4): 441-453. 

Roth, T.C., and B.D. Greene. 2006. Movement patterns and home range use of the northern watersnake (Nerodia sipedon). Copeia 2006 (3): 543-550.

Roth, T.C., S.L. Lima, and W.E. Vetter. 2006. Determinants of predation risk in small wintering birds: a hawk's perspective. Behavioral Ecology and Sociobiology 60 (2):195-204.

Roth, T.C., S.L. Lima, and W.E. Vetter. 2005. Survival and causes of mortality in wintering Sharp-shinned Hawks and Cooper’s Hawks.  Wilson Journal of Ornithology 117 (3): 237-244.

Roth, T.C., and S.L. Lima. 2003. Hunting behavior and diet of Cooper’s Hawks: An urban view of the small-bird-in-winter paradigm. Condor 105 (3): 474-483. 

Lima, S.L., W.A. Mitchell, and T.C. Roth. 2003. Predators feeding on behaviorally responsive prey: some implications for the classical models of optimal diet choice. Evolutionary Ecology Research 5 (7): 1083-1102.


Current Student Collaborations


Abi Buell – neural changes as a function of hybridization 

Josephine Cleverdon – behavioral response to rattlesnake rattles

Andrea Contreras Diaz - neural structure in hybridizing chickadees

Eva Ish – neural changes as a function of hybridization 

Ann Medina - neural changes as a function of hybridization

Saisha Ramdour – DCX expressionin in hybridizing chickadees 

Abimael Ramos – neural structure in hybridizing chickadees

Tati Zhang – neural structure in hybridizing chickadees


Course Information


PSY 230. Experimental Design and Statistics lab 

PSY 270. Neuroethics (Natural Sciences in Perspective) 

PSY/BFB 302. Biopsychology with lab

PSY/BFB 306. Evolution of Mind and Intelligence 

PSY 487. Collaborative in Biopsychology with lab