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Paul Harnik

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Paul Harnik

Assistant Professor of Earth and Environmental Geosciences

Department/Office Information

Earth and Environmental Geoscience
218 Ho Science Center
  • M 12:30pm - 2:30pm (218 Ho Science Center)
  • W 10:15am - 12:15pm (218 Ho Science Center)

Dr. Paul Harnik is a paleobiologist, interested in how organisms respond to environmental change, their capacities to adapt and move, as well as their vulnerabilities to extinction. He focuses on marine animals in oceans past and present to better understand the origins and maintenance of biodiversity on Earth and the impacts of human activities on coastal ecosystems.

To learn more about the courses that I teach, the research that my students and I engage in, and how to get involved in research, see the .

Ph.D., Evolutionary Biology, University of Chicago (2009)

B.A., Geology (Minor: History), Oberlin College (1998)

Assistant Professor, Department of Geology, vlog (2020-)

Associate Professor, Department of Earth and Environment, Franklin and Marshall College (2019-20)

Assistant Professor, Department of Earth and Environment, Franklin and Marshall College (2013-19)

Postdoctoral Fellow, National Evolutionary Synthesis Center (2011-13)

Postdoctoral Researcher, Department of Geological and Environmental Sciences, Stanford University (2009-2011)

Museum Educator, Paleontological Research Institution (1999-2003)

National Science Foundation – CAREER: Environmental drivers of life history variation in coastal ecosystems (2018-24)

National Academies of Sciences, Engineering, and Medicine – Gulf Research Program Early-Career Research Fellowship (2017-20)

National Science Foundation – Historical ecology of marine mollusks in the northern Gulf of Mexico: Investigating anthropogenic impacts in coastal ecosystems through live-dead analysis; Subaward to "The Keck Geology Consortium: Year-long Collaborative and Comprehensive Research Experiences for Undergraduates.” (2016-17)

National Evolutionary Synthesis Center – Determinants of extinction in ancient and modern seas working group (w/ S. Finnegan & R. Lockwood) (2010-12)

Peer-Reviewed Journal Articles

Mamo, B.L., J.D. Cybulski, Y. Hong, P.G. Harnik, A. Chao, A. Tsujimoto, C.-L. Wei, D.M. Baker, and M. Yasuhara. 2023. Modern biogeography of benthic foraminifera in an urbanized tropical marine ecosystem. The Geological Society of London Special Publications 529: SP529-2022-188.

Hong, Y., M. Yasuhara, H. Iwatani, P.G. Harnik, A. Chao, J.D. Cybulski, Y. Liu, Y. Ruan, X. Li, C.-L. Wei. 2022. Benthic ostracod diversity and biogeography in an urbanized seascape. Marine Micropaleontology 174: 102067.

Hong, Y., M. Yasuhara, H. Iwatani, A. Chao, P.G. Harnik, C-L Wei. 2021. Ecosystem turnover in an urbanized subtropical seascape driven by climate and pollution. Anthropocene 36:100304.

Harnik, P.G., H. Maherali, J.H. Miller, and P. Manos. 2018. Geographic range velocity and its association with phylogeny and life history traits in North American woody plants. Ecology and Evolution 8:2632-2644.

Harnik, P.G., M.L. Torstenson, and M.A. Williams. 2017. Assessing the effects of anthropogenic eutrophication on marine bivalve life history in the northern Gulf of Mexico. Palaios 32:678-688.

Hoehn, K.B., P.G. Harnik, and V.L. Roth. 2016. A framework for detecting natural selection on traits above the species level. Methods in Ecology and Evolution 7:331-339.

Finnegan, S., S.C. Anderson, P.G. Harnik, C. Simpson, D.P. Tittensor, J.E. Byrnes, Z.V. Finkel, D.R. Lindberg, L.H. Liow, R. Lockwood, H.K. Lotze, C.M. McClain, J.L. McGuire, A. O’Dea, and J.M. Pandolfi. 2015. Paleontological baselines for evaluating extinction risk in the modern oceans. Science 348:567-570. 

Liow, L.H., T. Reitan, and P.G. Harnik. 2015. Ecological interactions on macroevolutionary time scales: clams and brachiopods are more than ships that pass in the night. Ecology Letters 18:1030-1039.

Orzechowski, E.A., R. Lockwood, J. Byrnes, S.C. Anderson, S. Finnegan, Z.V. Finkel, P.G. Harnik, D.R. Lindberg, L.H. Liow, H.K. Lotze, C.R. McClain, J.L. McGuire, A. O’Dea, J.M. Pandolfi, C. Simpson, and D.P. Tittensor. 2015. Marine extinction risk shaped by trait-environment interactions over 500 million years. Global Change Biology 21:3595-3607.

Harnik, P.G., P.C. Fitzgerald, J.L. Payne, and S.J. Carlson. 2014. Phylogenetic signal in extinction selectivity in Devonian terebratulide brachiopods. Paleobiology 40:675-692.

Harnik, P.G., C. Simpson, and J.L. Payne. 2012. Long-term differences in extinction risk among the seven forms of rarity. Proceedings of the Royal Society B – Biological Sciences 279:4969-4976.

Harnik, P.G., H.K. Lotze, S.C. Anderson, Z.V. Finkel, S. Finnegan, D.R. Lindberg, L.H. Liow, R. Lockwood, C.M. McClain, J.L. McGuire, A. O’Dea, J.M. Pandolfi, C. Simpson, and D.P. Tittensor. 2012. Extinctions in ancient and modern seas. Trends in Ecology and Evolution 27:608-617.

Harnik, P.G. 2011. Direct and indirect effects of biological factors on extinction risk in fossil bivalves. Proceedings of the National Academy of Sciences USA 108:13594-13599.

Harnik, P.G., and R. Lockwood. 2011. Part N, Revised, Volume 1, Chapter 24: Extinction in the marine Bivalvia. Treatise Online 29:1–24 (an online journal for the Treatise on Invertebrate Paleontology).

Harnik, P. G., D. Jablonski, A. Z. Krug, and J. W. Valentine. 2010. Genus age, provincial area and the taxonomic structure of marine faunas. Proceedings of the Royal Society B - Biological Sciences 277:3427-3435.

Harnik, P. G. 2009. Unveiling rare diversity by integrating museum, literature, and field data. Paleobiology 35:190-208.

Simpson, C., and P. G. Harnik. 2009. Assessing the role of abundance in marine bivalve extinction over the post-Paleozoic. Paleobiology 35:631-647.

Harnik, P. G., and R. M. Ross. 2004. Models of inquiry-based science outreach to urban schools. Journal of Geoscience Education 52:420-428.

Harnik, P. G., and R. M. Ross. 2003. Developing effective K-16 geoscience research partnerships. Journal of Geoscience Education 51:5-8.

Harnik, P. G., and R. M. Ross. 2003. Assessing data accuracy when involving students in authentic paleontological research. Journal of Geoscience Education 51:76-84.

Ross, R. M., P. G. Harnik, W. D. Allmon, J. M. Sherpa, A. M. Goldman, P. L. Nester, and J. J. Chiment. 2003. The Mastodon Matrix Project as an experiment with large-scale collaboration in paleontological research. Journal of Geoscience Education 51:39-47.

Simonson, B. M., and P. G. Harnik. 2000. Have distal impact ejecta changed through geologic time? Geology 28:975-978.

Edited Volumes

Harnik, P.G., and Ross, R.M., eds., 2003. Geoscience research partnerships. Journal of Geoscience Education 51(1).