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Indiana University Bloomington

Department of Biology

Faculty & Research

Faculty Profile

P. David Polly

Photo of P. David Polly
Research Images
Research photo by P. David Polly

Average digitigrady in mammalian carnivore communities, measured using proportions of the calcaneum bone in the ankle.  The geographic distribution of this feature, which is related to locomotion, is correlated with mean annual temperature, macrovegetation cover, and ecological region.  (from Polly, 2009)

Research photo by P. David Polly

Reconstruction of Titanoboa cerrejonensis, giant snake from the Paleocene of Colombia.  Titanoboa was more than 13 meters long, a size that is only attainable in very hot climates.  The fossil remains of Titanoboa were recovered from the Cerrejon Coal Mine and described by my colleagues and I (Head et al., 2009).  We used morphometric models of morphological variation to identify the vertebral bones of the snake and to estimate its body size, from which the paleotemperature of the tropical Cerrejon forest in the Paleocene could be estimated. (reconstruction by Jason Bourque, (c) 2009).

Adjunct: Professor of Geological Sciences

IU Affiliations
Geological Sciences

Contact Information
By telephone: 812-855-7994
Geology Building, 513


Polly Lab website

Citations on GoogleScholar

Society of Vertebrate Paleontology

Research Areas
  • Ecology
  • Evolution

Ph.D., 1993, Department of Integrative Biology, University of California–Berkeley
B.A, 1987, Plan II Honors Program, University of Texas–Austin


2014-2018 - Vice-President, President, and Past President, Society of Vertebrate Paleontology.

2009-2014 - Research Associate, Department of Zoology, Field Museum of Natural History, Chicago.

2011 - James Philiop Holland Award for Exemplary Teaching and Service to Students.  Indiana University.

2011 - McCormick Science Grant (with A. Michelle Lawing) for faculty/graduate student team whose research is judged most creative, visionary, and innovative.  College of Arts and Sciences, Indiana University.

2004 - Draper\\\\\\\'s Prize for Excellence in Teaching, Queen Mary, University of London.

2001 - Joseph T. Gregory Award for outstanding service to the welfare of the Society of Vertebrate Paleontology.

2001.  Best Teacher of Biomedical Science (Dentistry), Barts and the London Medical and Dental Students Association, Queen Mary, University of London.

1994-1996 - Michigan Society Fellowship, Michigan Society of Fellows, University of Micigan, Ann Arbor.

Research Description

I work in the field of vertebrate paleontology, specifically with fossil mammals from the Cenozoic Era (65 million years ago to the present).  Paleontology is an interdisciplinary science, rooted firmly in evolution biology, anatomy, ecology, and geology.  Through paleontology, these disciplines have many connections, and I have intentionally structured my research to demonstrate how the rich historical context of geology – my primary discipline – can be used to improve understanding life on Earth, past and present.

Quantitative Functional Paleontology

This research is concerned with developing new approaches for measuring the complex morphological structures of fossils and the many processes that govern their evolution and geographic distribution through geological time.  Some of the most important properties of fossils -- their shape, their function, their relationship to sediments and stratigraphy, their geographic distributions through time, their variety, and their evolutionary trajectories – are difficult to capture with ordinary measurements.  One of my contributions has been to help develop the evolutionary side of geometric morphometric methods.  This work has included modeling evolutionary processes on multivariate phenotypic traits in GMM space, measuring the empirical patterns in teeth or limb bones from living populations and modeling their evolution over millions of generations to test how well competing hypotheses about the mode of evolution explain the observed fossil record.  With my former post-doc Anjali Goswami, I have extended this approach to simulate the evolutionary consequences of patterns of correlation between structures (such as those observed between the functionally integrated bones of the skull in living species) so that their relevance to evolution can be tested against the varieties of shape that are documented in the fossil record.  Norman MacLeod and I developed the “Eigensurface” method for measuring complete three-dimensional structures and modeling their evolution.  

Response of Vertebrates to Changing Climates and Environments

My research also addresses the response of vertebrates to changing climates and environments.  The geography of vertebrate species is influenced by climate and vegetation.  The sorting process is important for understanding the evolutionary processes that are driven by climatic and environmental change, as well as for understanding past climates and environments from fossil vertebrates.  Using mammalian species we have estimated paleotemperature and precipitation patterns during the last 2.6 million years of Earth’s history.  With former post-doc Jason Head and other collaborators, we used the scaling between ambient temperature and body size in poikilothermic (“cold-blooded”) vertebrates to estimate the mean annual temperatures of the tropics in the Paleocene (65-55 million years ago) using Titanoboa, the giant Paleocene snake. 

Software and Data

I have made available add-in packages for Mathematica to perform geometric morphometrics, evolutionary modeling, species distribution modeling, modularity analysis, and phylogenetic tree manipulation, which are available here.  Geographic grids, paleoclimate data, and phylogenetic matrices are available here.


Select Publications
Polly, P. D. 2014. Issues in Paleobiology. In: M. Sánchez-Villagra and N. MacLeod (eds), Issues in Paleobiology. Scidinge Hall Verlag: Zürich, Switzerland.  [article]
Goswami, A., J. B. Smaers, C. Soligo, and P. D. Polly.  2014.  The macroevolutionary consequences of phenotypic integration:  from development to deep time.  Philosophical Transactions of the Royal Society B, 369: 20130254   [article]
Polly, P.D. and S. Sarwar.  2014.  Extinction, extirpation, and exotics: effects on the correlation between traits and environment at the continental level. Annales Zoologici Fennici, 51: 209-226.  [article]
Polly, P. D. 2014. Trait-based extinction catches the Red Queen napping during the Cambrian. Proceedings of the National Academy of Sciences USA, doi: 10.1073/pnas.1419138111.  [article]
Grossnickle, D.M. and P.D. Polly. 2013. Mammal disparity decreases during the Cretaceous angiosperm radiation. Proceedings of the Royal Society, B, 280 doi: 10.1098/rspb.2013.2110  [article]
Cardini, A. and P.D. Polly. 2013. Larger mammals have longer faces because of size-related constraints on skull form. Nature Communications4, art. 2458: 1-7.  [article]
Polly, P. D.,  A. M. Lawing, A.-C. Fabre, and A. Goswami. 2013. Phylogenetic Principal Components Analysis and Geometric Morphometrics. Hystrix, 24: 1-9.  [article]
Uhen, M.D., A.D. Barnosky, B. Bills, J. Blois, M.T. Carrano, M.A. Carrasco, G.M. Erickson, J.T. Eronen, M. Fortelius, R.W. Graham, E.C. Grimm, M.A. O'Leary, A. Mast, W.H. Piel, P.D. Polly, and L.K. Säilä.  2013.  From card catalogs to computers:  databases in vertebrate paleontology.  Journal of Vertebrate Paleontology, 33: 13-28.   [article]
Polly, P. D. 2013. Evolution: Stuck between the teeth. Nature, 497: 325–326.  [article]
Polly, P.D., A.V. Polyakov, V.B. Ilyashenko, S.S. Onischenko, T.A. White, N.S. Bulatova, S. Pavlova, P.M. Borodin, and J.B. Searle. 2013. Phenotypic variation across chromosomal hybrid zones of the Common shrew (Sorex araneus) indicates reduced gene flow. PLoS One, 8(7): e67455  [article]
Smith, M.R. and P.D. Polly.  2013.  A reevaluation of the Harrodsburg Crevice Fauna (Late Pleistocene of Indiana, USA) and the climatic implication of its mammals.  Journal of Vertebrate Paleontology, 33: 410-420.    [article]
Rödder D., A. M. Lawing, M. Flecks, F. Ahmadzadeh, J. Dambach, J. O. Engler, J. C. Habel, T. Hartmann, D. Hörnes, F. Ihlow, K. Schidelko, D. Stiels, and P. D. Polly. 2013 Evaluating the Significance of Paleophylogeographic Species Distribution Models in Reconstructing Quaternary Range-Shifts of Nearctic Chelonians. PLoS ONE 8(10): e72855.  [article]
Gómez-Robles, A., J. M. Bermúdez de Castro, J.-L. Arsuaga, E. Carbonell, and P. D. Polly. 2013. No known hominin species matches the expected dental morphology of the last common ancestor of Neanderthals and modern humans. Proceedings of the National Academy of Sciences, USA, 110: 18196-18201.   [article]
Gómez-Robles, A. and P.D. Polly.  2012.  Morphological integration in the hominin dentition:  evolutionary, developmental, and functional factors.  Evolution, 66: 1024-1043.   [article]
Goswami, A., P.D. Polly, O.B. Mock, and M.R. Sánchez Villagra.  2012.  Shape, variance, and integration during craniogenesis: constrasting marsupial and placental mamamls.  Journal of Evolutionary Biology, 25: 862-872.  [article]
Polly, P.D. 2012. Measuring the evolution of body size in mammals.  PNAS, 109: 4027-4028.  [article]
Lawing, A.M., J.J. Head, and P.D. Polly.  2012.  The ecology of morphology:  the ecometrics of locomotion and macroenvironment in North American snakes.  Pp. 117-146 in J. Louys (ed), Paleontology in Ecology and Conservation.  Springer-Verlag, Berlin and Heidelberg  [article]
Polly, P.D., J.T. Eronen, M. Fred, G.P. Dietl, V. Mosbrugger, C. Scheidegger, D.C. Frank, J. Damuth, N.C. Stenseth & M. Fortelius. 2011. History matters: ecometrics and integrative climate change biology. Proceedings of the Royal Society B, 278: 1121-1130.   [article]
Polly, P.D. and J.T. Eronen.  2011.  Mammal associations in the Pleistocene of Britain:  implications of ecological niche modelling and a method for reconstrucing palaeoclimate. Pp. 279-304 in N. Ashton, S. G. Lewis, and C. Stringer (eds.), The Ancient Human Occupation of Britain.  Elsevier.   [article]
Lawing, A. M. and P.D. Polly.  2011.  Pleistocene climate, phylogeny, and climate envelope models: an integrative approach to better understand species' response to climate change.  PLoS ONE, 16: e28554  [article]
Johnson, L.J., J.A. Cotton, C.P. Lichtenstein, G.S. Elgar, R.A. Nichols, P.D. Polly, and S.C. Le Comber. 2011.  Stops making sense: translational trade-offs and stop codon reassignment.  BMC Evolutionary Biology, 11: 227.  [article]
Goswami, A. and P.D. Polly. 2010 . Methods for studying morphological integration, modularity and covariance evolution. Pp. 213-243 in J. Alroy and G. Hunt (eds.), Quantitative Methods in Paleobiology. Paleontological Society Short Course, October 30th, 2010. The Paleontological Society Papers, Volume 16.   [article]
Lawing, A.M. and P.D. Polly. 2010. Geometric morphometrics:  recent applications to the study of evolution and development.  Journal of Zoology, 280: 1-7.  [article]
Goswami, A. and P.D. Polly. 2010. The influence of character correlations on phylogenetic analyses: a  case study of the carnivoran craniu. Pp. 141-164 in A. Goswami and A. Friscia (eds.), Carnivoran Evolution: New Views on Phylogeny, Form, and Function. Cambridge University Press.  [article]
Polly, P.D.  2010.  Tiptoeing through the trophics: geographic variation in carnivoran locomotor ecomorphology in relation to environment.   In A. Goswami and A. Friscia (eds.), Carnivoran Evolution: New Views on Phylogeny, Form, and Function.  [article]
Head, J.J., J.I. Bloch, A.K. Hastings, J.R. Bourque, E. Cadena, F. Herrera, P.D. Polly, and C.A. Jaramillo.  2009.  Giant boine snake from a Paleocene Neotropical rainforest indicates hotter past equatorial temperatures.  Nature, 457: 715-718.  [article]
Polly, P.D.  2008.  Developmental dynamics and G-matrices:  Can morphometric spaces be used to model evolution and development?  Evolutionary Biology, 35: 83-96.  [article]
Polly, P. D.  2008.  Adaptive Zones and the Pinniped Ankle: A 3D Quantitative Analysis of Carnivoran Tarsal Evolution.  Pp. 165-194 in (E. Sargis and M. Dagosto, Eds.) Mammalian Evolutionary Morphology: A Tribute to Frederick S. Szalay.  Springer: Dordrecht, The Netherlands.  [article]
Polly, P.D.  2007.  Development with a Bite.  News & Views.  Nature, 449: 413-415.  [article]
Gündüz, İ, M Jaarola, C. Tez, C. Yeniyurt, P.D. Polly, and J. B. Searle.  2007.  Multigenic and morphometric differentiation of ground squirrels (Spermophilus, Scuiridae, Rodentia) in Anatolia, with a description of a new species.  Molecular Phylogenetics and Evolution 43: 916-935.  [article]
Polly, P. D., G. D. Wesley-Hunt, R. E. Heinrich, G. Davis, and P. Houde. 2006.  Earliest known carnivoran auditory bulla and support for a recent origin of crown-group Carnivora (Eutheria, Mammalia).  Palaeontology, 49: 1019-1027.
Caumul, R. and P. D. Polly.  2005.  Comparative phylogenetic and environmental components of morphological variation:  skull, mandible and molar shape in marmots (Marmota, Rodentia). Evolution, 59: 2460-2472.
Polly, P. D.  2004.  On the simulation of the evolution of morphological shape:  multivariate shape under selection and drift.  Palaeontologia Electronica, 7.2.7A: 28pp, 2.3MB.   [article]

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