Dr. Patricia Kelley

University of North Carolina Wilmington
Department of Geography and Geology

Deloach 117
601 South College Road
Wilmington, NC 28403

Tel: 910 962-7406
Fax: 910 962-7077

Hampton Beach, NH

Research Interests

My research focuses on the evolution and paleoecology of Coastal Plain molluscs.  I am especially interested in the evolutionary process and the factors that control it, and I believe that the fossil record contains data essential for deciphering this process.  My initial studies, a test of the hypothesis of punctuated equilibrium, investigated tempo and mode of evolution of the Miocene mollusc fauna of Maryland.  More recent and current work focuses on evolutionary paleoecology, with a particular interest in the role of ecological factors, such as predation, in evolution.  A major thrust of this work has involved testing Vermeij’s hypothesis of escalation, which proposes that biological hazards such as predation, along with adaptation to those hazards, have increased through geological time.  Thor Hansen (Western Washington University) and I have compiled a database on predation by shell-drilling naticid gastropods in the Gulf and Atlantic Coastal Plain; our collections now include >150,000 mollusc specimens of Cretaceous through Pleistocene age.  We have used Kelleythese data to test a variety of hypotheses about the dynamics and evolutionary history of predator-prey interactions, predator-prey coevolution and escalation, the ecology of mass extinctions and recoveries of mollusc faunas, and spatial variation in predation (in part in collaboration with middle school teachers and students in “The Moonsnail Project”). 

I enjoy collaborating on my research with students at all levels, from undergraduate to PhD. I have coauthored papers or abstracts with 65 graduate students and more than 160 undergraduate students. Recent and current work in collaboration with graduate students includes: the ecological response of the Iceland molluscan fauna to the invasion of predatory gastropods during the Pliocene; taphonomy of an ophiuroid mass mortality bed in Mexico; geographic variation in drilling predation in North America, Brazil and Argentina; relationship between drilling predation and prey diversity in the US Coastal Plain and Europe; conditions under which cannibalism by drilling predators occurs; metabolism and extinction susceptibility in Plio-Pleistocene bivalves; escalation, coevolution, and drilling predation on bivalves and gastropods; factors affecting durophagous predation in ammonites, bivalves, and gastropods; live-dead comparisons of molluscan assemblages as tools in conservation paleobiology; life span bias in the fossil record; season of mortality due to drilling and crushing predation on Pleistocene bivalves.










College of Wooster        B.A., Geology                 1975
Harvard University        A.M., Geology                1977
Harvard University        Ph.D., Geology               1979



Books Published


Kelley, P.H., J. Bryan, and T.A. Hansen (eds). 1999. The Evolution-Creationism Controversy II:Perspectives on Science, Religion, and Geological Education. The Paleontological Society Papers, vol. 5 (series editor, W. Manger).

Kowalewski, M., and P.H. Kelley (eds). 2002.The Fossil Record of Predation.The Paleontological Society Papers, vol. 8, 398 p.

Kelley, P.H., M. Kowalewski, and T.A. Hansen (eds). 2003.Predator-Prey Interactions in the Fossil Record.Kluwer Academic/Plenum Press, 464 p.

Kelley, P.H., and R.K. Bambach (eds). 2008.From Evolution to Geobiology: Paleontology at the Start of a New Century.Paleontological Society Papers, 374 p.

Allmon, W.D., P.H. Kelley, and R.D. Ross (eds).2009. Stephen Jay Gould: Reflections on His View of Life.Oxford University Press, 400 p.

Kelley, P.H., G.P. Dietl, and L.W. Ward. 2011. Field Trip 401 – Plio-Pleistocene Stratigraphy and Paleontology of Southeastern North Carolina. In Taylor, K.B., and J.C. Reid (eds), Field Trip Guidebook – 60th Annual Meeting Southeastern Section, Geological Society of America, 28 p. + Appendix.


Recent Articles


Kelley, P.H. 2000. Studying evolution and keeping the faith. Geotimes 45(12):22-23,41.

Kelley, P.H., and T.A. Hansen. 2001. The role of ecological interactions in the evolution of naticid gastropods and their molluscan prey. Pp. 149-170. In W. Allmon and D. Bottjer, Evolutionary Paleoecology. Columbia University Press, New York.

Kelley, P.H., and T.A. Hansen. 2001. The Mesozoic marine revolution, p. 94-97. In D.E.G. Briggs and P.R. Crowther, Palaeobiology II. Blackwell Science, Oxford.

Kelley, P.H., T.A. Hansen, S.E. Graham, and A.G. Huntoon. 2001. Temporal patterns in the efficiency of naticid gastropod predators during the Cretaceous and Cenozoic of the United States Coastal Plain. Palaeogeography, Palaeoclimatology, Palaeoecology 166(1/2):165-176.

Kelley, P.H. and T.A. Hansen. 2001. Extinction. New Book of Knowledge, 5:425-426. (also published online at

Dietl, G.P. and P.H. Kelley. 2001. Mid-Paleozoic latitudinal predation gradient: Distribution of brachiopod ornamentation reflects shifting Carboniferous climate. Geology 29:111–114.

Dietl, G.P., P.H. Kelley, R. Barrick and W. Showers. 2002. Escalation and extinction selectivity: morphology versus isotopic reconstruction of bivalve metabolism. Evolution 56(2):284-291.

Hansen, T.A., and P.H. Kelley. 2002. Moonsnail project: roles of technology and research in learning science. pp. 57-64, In Chambers, J.A. (ed.) Selected Papers from the Thirteenth International Conference on College Teaching and Learning. The Center for the Advancement of Teaching and Learning.

Dietl, G. P. and P.H. Kelley. 2002. The fossil record of predator-prey arms races: coevolution and escalation hypotheses, p. 353-374. In M. Kowalewski and P.H. Kelley (eds). The Fossil Record of Predation. The Paleontological Society Papers, vol. 8.

Kelley, P.H., and T.A. Hansen. 2003. The fossil record of drilling predation on bivalves and gastropods, p. 113-139. In P.H. Kelley, M. Kowalewski, and T.A. Hansen (eds), Predator-Prey Interactions in the Fossil Record. Kluwer Academic/Plenum Press.

Hansen, T.A., P.H. Kelley, and J.C. Hall. 2003. Moonsnail Project: a scientific collaboration with middle school teachers and students. Journal of Geoscience Education 50(1):35-38.

Hansen, T.A., P.H. Kelley, and D.M. Haasl. 2004. Paleoecological patterns in molluscan extinctions and recoveries: Comparison of the Cretaceous-Tertiary and Eocene-Oligocene extinctions in North America. Palaeogeography, Palaeoclimatology, Palaeoecology 214(3):233-242.

Reinhold, M.E., and P.H. Kelley. 2005. The influence of anti-predatory morphology on survivorship of the Owl Creek Formation molluscan fauna through the end-Cretaceous extinction. Palaeogeography, Palaeoclimatology, Palaeoecology 217:143-153.

Kelley, P.H., and T.A. Hansen. 2006. Comparisons of class- and lower taxon-level patterns in naticid gastropod predation, Cretaceous to Pleistocene of the U.S. Coastal Plain. Palaeogeography, Palaeoclimatology, Palaeoecology 236(3/4):302-320.

Dietl, G.P., and P.H. Kelley. 2006. Can naticid gastropod predators be discriminated by the holes they drill? Ichnos 13:1-6.

Kelley, P.H., and T.A. Hansen. 2007. A case for cannibalism:  Confamilial and conspecific predation by naticid gastropods, Cretaceous through Pleistocene of the United States Coastal Plain, p. 151-170.  In Elewa, A.M.T. (ed.), Predation in Organisms: A Distinct Phenomenon. Springer Verlag.

Kelley, P.H., and T.A. Hansen. 2007. Latitudinal patterns in naticid gastropod predation along the east coast of the United States: a modern baseline for interpreting temporal patterns in the fossil record, p. 284-299. In: Bromley, R.G., Buatois, L.A., Mángano, M.G., Genise, J.F., and Melchor, R.N. (eds.), Sediment-Organism Interactions: A multifaceted Ichnology. SEPM Special Publications, v. 88.

Kelley, P.H. 2008. Role of bioerosion in taphonomy: effect of predatory drillholes on preservation of mollusc shells, p. 451-470. In Tapanila, L., and M. Wisshak (eds), Current Developments in Bioerosion. Springer.

Kelley, P.H., 2008. The View from the Top: A Century of PS Presidents’ Perspectives on the Paleontological Society and Paleontology, p. 1-15.  In Kelley, P.H., and R.K. Bambach (eds), From Evolution to Geobiology: Paleontology at the Start of a New Century.  Paleontological Society Papers.

Bruce, K., J. Horan, P.H. Kelley, and M. Galizio. 2009.  Teaching Evolution in the Galapagos.  Journal of Effective Teaching 9(2):13-28.

Kelley, P.H. 2009.  Stephen Jay Gould’s Winnowing Fork: Science, Religion, and Creationism, p. 171-188.  In Allmon, W.D., P.H. Kelley, and R.D. Ross (eds), Stephen Jay Gould: Reflections on His view of Life.  Oxford University Press.

Kelley, P.H. 2009.  Teaching Evolution During the Week and Bible Study on Sunday:  Perspectives on Science, Religion, and Intelligent Design, P. 163-179.  In Schneiderman, J.S., and W.D. Allmon (eds), For the Rock Record: Geologists On Intelligent Design Creationism.  University of California Press.

Kelley, P.H. 2009.  A College Honors Seminar on Evolution and Intelligent Design: Successes and Challenges. Journal of Effective Teaching 9(2):29-37.

Dietl, G., S. Durham, and P. Kelley.  2010.  Shell repair as a reliable indicator of bivalve predation by shell-wedging gastropods in the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 296:174-184.

Ottens, K.J., G.P. Dietl*, P.H. Kelley*, and S.D. Stanford. 2012. A comparison of analyses of drilling predation on fossil bivalves: bulk- vs. taxon-specific sampling and the role of collector experience. Palaeogeography, Palaeoclimatology, Palaeoecology 319-320:84-92. * contributed equally.

Harper, E.M., and P.H. Kelley. 2012. Predation of bivalves. Treatise Online. Part N, Revised, Bivalvia, Volume 1, Chapter 22, p. 1-21.

Carter, J.G., P.J. Harries, N. Malchus, A.F. Sartori, L.C. Anderson, R. Bieler, A.E. Bogan, E.V. Coan, J.C.W. Cope, S. Cragg, J.R. García-March, J. Hylleberg, P. Kelley, K. Kleemann, J. Kříž, C. McRoberts, P.M. Mikkelsen, J. Pojeta, Jr., I. Tëmkin, T. Yancey, and A. Zieritz. 2012. Illustrated Glossary of the Bivalvia. Treatise on Invertebrate Paleontology. Part N, Revised, Bivalvia, Volume 1, Chapter 31. Treatise Online 48:1–209. Published online 7/13/12.

Kelley, P.H. 2012. Strategies for teaching evolution in a high-enrollment introductory paleontology course for non-science majors, p. 77-92. In Yacobucci, M.M., and R. Lockwood, Teaching Paleontology in the 21st Century. Paleontological Society.

Kelley, P.H., and C.C. Visaggi. 2012. Learning paleontology through doing: Integrating an authentic research project into an invertebrate paleontology course, p. 181-197. In Yacobucci, M.M., and R. Lockwood, Teaching Paleontology in the 21st Century. Paleontological Society.

Visaggi, C.V., G.P. Dietl, and P.H. Kelley. 2013. Influence of sediment depth and prey health on drilling behaviour of Neverita duplicata (Gastropoda: Naticidae) with a review of alternative modes of predation. Journal of Molluscan Studies 79:310-322. Published online 7/16/13: doi: 10.1093/mollus/eyt023

Surge, D., Ting Wang, I. Gutiérrez-Zugasti, and P.H. Kelley. 2013. Isotope sclerochronology and season of annual growth line formation in limpet shells (Patella vulgata) from warm- and cold-temperate zones in the eastern north Atlantic. Palaios 28:386-393.

Kelley, P.H., D.E. Fastovsky, M.A. Wilson, R.A. Laws, and A. Raymond. 2013. From paleontology to paleobiology: A half-century of progress in understanding life history, p. 191-232. In Bickford, M.E., ed., The Web of Geological Sciences: Advances, Impacts, and Interactions: Geological Society of America Special Paper 500. doi:10.1130/2013.2500(06)

Korpanty, C.A., and P.H. Kelley. 2014. Live-Dead Molluscan Fidelity in Anthropogenically Impacted Seagrass Habitats: Siliciclastic versus Carbonate Environments. Palaeogeography, Palaeoclimatology, Palaeoecology 410:113-125. doi: 10.1016/j.palaeo.2014.05.014

Kelley, P.H. 2014. Controversial topics in the classroom: (how) do we dare discuss them?, p. 133-143. In Ashe, D., and C. Clements, Best Practices in University Teaching (2d ed). University of North Carolina Wilmington.

Hattori, K.E., P.H. Kelley, G.P. Dietl, N.O. Moore, S.L. Simpson, A.M. Zappulla, K.J. Ottens, C.C. Visaggi. 2014. Validation of taxon-specific sampling by novice collectors for studying drilling predation in fossil bivalves. Palaeogeography, Palaeoclimatology, Palaeoecology 412:199-207.

Klompmaker, A., and P.H. Kelley. 2015. Shell ornamentation as an exaptation: evidence from predatory drilling on Cenozoic bivalves. Paleobiology 41(1):187-201. DOI: 10.1017/pab.2014.12

Visaggi, C.C., and P.H. Kelley. 2015. Equatorward increase in naticid gastropod drilling predation on infaunal bivalves from Brazil with paleontological implications. Palaeogeography, Palaeoclimatology, Palaeoecology 438:285-299.

Kerr, J.P., and P.H. Kelley. 2015. Assessing the influence of escalation during the Mesozoic Marine Revolution: shell breakage and adaptation against enemies in Mesozoic ammonites. Palaeogeography, Palaeoclimatology, Palaeoecology 440:632-646. DOI:10.1016/j.palaeo.2015.08.047.