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Simon Levin

Adjunct Professor and George M. Moffett Professor of Biology, Princeton University


  • Ecology and Evolutionary Biology

Graduate Fields

  • Applied Mathematics
  • Ecology and Evolutionary Biology


  • Understanding how macroscopic patterns and processes are maintained at the level of ecosystems and the biosphere, in terms of ecological and evolutionary mechanisms that operate primarily at the level of organisms
  • Evolution of diversification, the mechanisms sustaining biological diversity in natural systems, and the implications for ecosystem structure and functioning
  • Development of macroscopic descriptions for the collective behavior of large and heterogeneous ensembles that are subject to continual evolutionary modification
  • Evolution and ecology of dispersal in plant communities, marine open-ocean and intertidal systems
  • Self-organization and evolution of strain structure in influenza A
  • Conservation implications of the basic ecological work, with emphasis on reserve design and on economic linkages


  • Moore, S.A., T.J. Wallington, R.J. Hobbs, P.R. Ehrlich, C.S. Holling, S.A. Levin, D. Lindenmayer, C. Pahl-Wostl, H. Possingham, M.G. Turner, and M. Westoby. 2006. Diversity in current ecological thinking: implications for environmental management. Environmental Management. DOI 10.1007/S00267-008-9187-2.
  • Satake, A., Y. Iwasa, and S.A. Levin. 2008. Comparison between perfect information and passive adaptive social learning models of forest harvesting. Theoretical Ecology. DOI 10.1007/S12080-008-0019-z.
  • Bartumeus, F., P. Fernandez, M.G.E. daLuz, J. Catalan, R.V. Sole and S. Levin. 2008. Superdiffusion and encounter rates in diluted, low dimensional worlds. European Physics Journal Special Topics 157: 157-166.
  • Levin, S.A. and J. Lubchenco. 2008. Resilience, robustness, and marine ecosystem-based management. Bioscience 58(1): 27-32.
  • Buchman, T.G., J. Dushoff, M.B. Effron, P.R. Ehrlich, S. Fitzpatrick, R. Laxminarayan, B. Levin, S.A. Levin, M. Lipsitch, A. Malani, C. Nemeroff, S.P. Otto, V.L. Patel, and J.S. Solomkin. 2008. Antibiotic overuse: the influence of social norms. Journal of the American College of Surgeons. In press.
  • May, R.M., S.A. Levin, and G. Sugihara. 2008. Ecology for bankers. Nature 451: 893-895.
  • Klausmeier, C.A., E. Litchman, T. Daufresne, and S.A. Levin. 2008. Phytoplankton stoichiometry. Ecological Research 23: 479-485.
  • Menge, D.N.L., S.A. Levin, and L.O. Hedin. 2008. Evolutionary tradeoffs can select against nitrogen fixation and thereby maintain nitrogen limitation. PNAS 105(5): 1573-1578.
  • Nadell, C.D., J. Xavier, S.A. Levin, and K.R. Foster. 2008. The evolution of quorum sensing in bacterial biofilms. PLoS Biology 6(1): 171-179.
  • Perring, M.P., L. Hedin, S.A. Levin, M. McGroddy, and C. de Mazancourt. 2008. Increased plant growth from nitrogen addition should conserve phosphorus in terrestrial ecosystems. PNAS. 105(6): 1971-1976.
  • Stock, C.A., T.M. Powell, and S.A. Levin. 2008. Bottom-up and Top-down forcing in a simple size structured plankton dynamics model. Journal of Marine Systems. In press.
  • Wiegand, K., D. Saltz, D. Ward, and S.A. Levin. 2008. The role of size inequality in self-thinning: a pattern-oriented simulation model for arid savannas. Ecological Modelling 210: 431-445.
  • Baskett, M., J. Weitz and S.A. Levin. 2007. The evolution of dispersal in reserve networks. American Naturalist 170(1): 59-78.
  • Baskett, M., F. Micheli, and S.A. Levin. 2007. Designing marine reserves for interacting species: Insights from theory. Biological Conservation 137(2): 163-179.
  • Cullen, J.J., W.F. Doolittle, S.A. Levin, and W.K.W. Li. 2007. Patterns and predictions in microbial oceanography. Oceanography 20(2): 32-44.
  • Klausmeier, C.A., E. Litchman, and S.A. Levin. 2007. A model of flexible uptake of two essential resources. Journal of Theoretical Biology 246(2): 278-289.
  • Kryazhimskiy, S., U. Dieckmann, S.A. Levin, and J. Dushoff. 2007. On state-space reduction in multistrain
  • pathogen models, with an application to antigenic drift in influenza A. PloS Computational Biology.
  • Levin, S.A. 2007. Introduction: Infectious diseases. Environment and Development Economics 12: 625-626.
  • Levin, S.A. 2007. Book review. Remodeled Foundations. Theoretical ecology: principles and applications. 3rd ed. Bob May and Angela McClean, eds. Science 316: 1699-1700.
  • Levin, S.A. 2007. Book review. Evolutionary Dynamics: Exploring the Equations of Life, by M.A. Nowak. Quarterly Review of Biology.
  • Lichstein, J.W., J. Dushoff, S.A. Levin, and S.W. Pacala. 2007. Intraspecific variation and species coexistence. American Naturalist 170: 807-818.
  • Ma, J., L. Worden and S.A. Levin. 2007. Evolutionary branching of single traits. (K. McCann, D. Noakes, N. Rooney, eds.). From Energetics to Ecosystems: The Dynamics and Structure of Ecological Systems. Chapter 10, p. 191-212.
  • Moon, S.J., B. Nabet, N. E. Leonard, S.A. Levin, and I.G. Kevrekidis. 2007. Heterogeneous animal group models and their group-level alignment dynamics; an equation-free approach. Journal of Theoretical Biology 246: 100-112.
  • Muneepeerakul, R., S.A. Levin, A. Rinaldo, and I. Rodriguez-Iturbe. 2007. On Biodiversity in River Networks: a trade-off metapopulation model and comparative analysis. Water Resources Research 43(7): W07426, doi:10.1029/2006WR005857.
  • Muneepeerakul, R., J. S. Weitz, S.A. Levin, A. Rinaldo, and I. Rodriguez-Iturbe. 2007. A neutral metapopulation model of biodiversity in river networks. Journal of Theoretical Biology 245(2): 351-363.
  • Pulliam, J.R.C., J. Dushoff, S.A. Levin, and A.P. Dobson. 2007. Epidemic enhancement in partially immune populations. PLoS ONE 2(1): e165. doi:10.1371/journal.pone.0000165.
  • Satake, A., M.A. Janssen, S.A. Levin, and Y. Iwasa. 2007. Synchronized deforestation induced by social learning under uncertainty of forest-use value. Ecological Economics 63(2-3): 452-462.
  • Satake, A., H.M. Leslie, Y. Iwasa, and S.A. Levin. 2007. Coupled ecological-social dynamics in a forested landscape: spatial interactions and information flow. Journal of Theoretical Biology 246(4): 695-707.
  • Scanlon, T., K. Caylor, S.A. Levin, and I. Rodriguez-Inturbe. 2007. Positive feedbacks promote power-law clustering of Kalahari vegetation. Nature 449: 209-212 (doi:10.1038/nature06060).
  • Worden, L. and S.A. Levin. 2007. Evolutionary escape from the prisoner’s dilemma. Journal of Theoretical Biology 245: 411-422.
  • Bazykin, G.A., J. Dushoff, S.A. Levin, and A.S. Kondrashov. 2006. Bursts of non-synonymous substitutions in HIV-1 evolution reveal instances of positive selection at conservative protein sites. Proceedings of the National Academy of Sciences, USA 103(51): 19396-19401.
  • Buchman, T.G., V.L. Patel , J. Dushoff, P.R. Ehrlich, M. Feldman, M. Feldman, B. Levin, D.T. Miller, P. Rozin, S.A. Levin, and S. Fitzpatrick. 2006. Enhancing the use of clinical guidelines: A social norms perspective. Journal of the American College of Surgeons 202(5): 826-836.
  • Buchman, T.G., J. Dushoff, P.R. Ehrlich, M. Feldman, M. Feldman, S. Fitzpatrick, B. Levin, S.A. Levin, D.T. Miller, V.L. Patel, and P. Rozin. 2006. Battling bad behavior: how do you convince people to do what’s in their best interest? The Scientist. 20(2): 51-57.
  • Casagrandi, R., L. Bolzoni and S.A. Levin. 2006. The SIRC model for the ecology and evolution of drifting influenza A in seasonal environments. Mathematical Biosciences. 200(2): 152-169.
  • Chapin, F.S., Hoel, Michael, Carpenter, Steven R., Lubchenco, Jane, Walker, Brian, Callaghan, Terry V., Folke, Carl, Levin, Simon A., Mäler, Karl-Göran, Nilsson, Christer, Barrett, Scott, Berkes, Fikret, Crépin, Anne-Sophie, Danell, Kjell, Rosswall, Thomas, Starrett, David, Xepapadeas, Anastasios, Zimov, Sergey A. 2006. Building Resilience and Adaptation to Manage Arctic Change. AMBIO: A Journal of the Human Environment: 35(4): 198–202.
  • De Leenheer, P., S.A. Levin, E.D. Sontag and C.A. Klausmeier. 2006. Global stability in a chemostat with multiple nutrients. J. Mathematical Biology 52(4): 419-438.
  • Earn, D.J.D. and S. A. Levin. 2006. Global asymptotic coherence in discrete dynamical systems. Proceedings of the National Academy of Sciences, USA 103(11): 3968-3971. Erratum December 19, 2006, 103(51): 19605; originally published December 8, 2006.
  • Levin, S.A. 2006. On Karl Hadeler becoming 70. Journal of Mathematical Biology 53(4): 496-498.
  • Levin, S.A. 2006. Unity from division: in search of a collective kokoro. In Quest of Kokoro/Human Minds for this Planet. Kyoto International Culture Forum, Kyoto University, October 2006.
  • Levin, S.A. 2006. Learning to live in a Global Commons: Socioeconomic challenges for a sustainable environment. Ecological Research. Special Feature 21(3): 328-333.
  • Levin, S.A. 2006. Fundamental Questions in Biology. PLoS Biology 4(9): 1471-1472.
  • Ma, J. and S.A. Levin. 2006. The evolution of resource adaptation: how generalist and specialist consumers evolve. Bulletin of Mathematical Biology 68: 1111-1123.
  • Nabet, B., Leonard, N.E., Couzin, I. and S.A. Levin. 2006. Leadership in animal group motion: A bifurcation analysis. Proceedings of the 17th International Symposium on Mathematical Theory of Networks and Systems (MTNS), Kyoto, Japan, July 24-28, 2006.
  • Pascual, M., J.A. Dunne and S.A. Levin. 2006. Challenges for the future: Integrating ecological structure and dynamics. Ecological Networks: Linking Structure to Dynamics in Food Webs. Pp. 351-371. Eds. M. Pascual and J.A. Dunne. Oxford University Press, New York.
  • Sterner, T., M. Troell, J. Vincent, S. Aniyar, S.Barrett, W. Brock, S. Carpenter, K. Chopra, P. Ehrlich, M. Hoel, S. Levin, K, Goran Maler, J. Norberg, L. Pihl, T. Soderquist, J. Wilen and A. Xepapadeas. 2006. Quick Fixes for environment: part of the solution, or part of the problem? Environment 48(10): 20-27.
  • Weitz, J. and S.A. Levin. 2006. Size and scaling of predator-prey dynamics. Ecology Letters 9: 548-557.
  • Wingreen, N.S. and S.A. Levin. 2006. Cooperation among microorganisms. PLoS Biology 4(9): 1486-1488.
  • Zea-Cabrera, E., Y. Iwasa, S. Levin and I. Rodriguez-Iturbe. 2006. Tragedy of the commons in plant water use. Water Resources Research 42. W06D02, doi:10.1029/2005WR004514. (figure correction)